0.2 <documentation xmlns="http://cellml.org/tmp-documentation"> <article> <articleinfo> <title>Biophysical Characteristics of the Pig Kidney Na/Glucose Cotransporter SGLT2 Reveal a Common Mechanism for SGLT1 and SGLT2</title> <author> <firstname>Jonna</firstname> <surname>Terkildsen</surname> <affiliation> <shortaffil>Auckland Bioengineering Institute, University of Auckland</shortaffil> </affiliation> </author> </articleinfo> <sect1 id="sec_status"> <title>Model Status</title> <para>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> </sect1> <sect1 id="sec_structure"> <title>Model Structure</title> <para>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>The original paper reference is cited below:</para> <para> 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>Journal of Biological Chemistry</emphasis> , 271, 32678-32683. <ulink url="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=pubmed&amp;cmd=Retrieve&amp;dopt=AbstractPlus&amp;list_uids=8955098&amp;query_hl=1&amp;itool=pubmed_docsum">PubMed ID: 8955098</ulink> </para> <informalfigure float="0" id="fig_reaction_diagram"> <mediaobject> <imageobject> <objectinfo> <title>model diagram</title> </objectinfo> <imagedata fileref="mackenzie_1996.png" /> </imageobject> </mediaobject> <caption>Schematic diagram of the Mackenzie et al 1996 SGLT2 model. C' represents the external-facing carrier. C'' represents the internal-facing carrier.</caption> </informalfigure> </sect1> </article> </documentation> <rdf:RDF xmlns:rdf="http://www.w3.org/1999/02/22-rdf-syntax-ns#" xmlns:dcterms="http://purl.org/dc/terms/" xmlns:dc="http://purl.org/dc/elements/1.1/" xmlns:vCard="http://www.w3.org/2001/vcard-rdf/3.0#" xmlns:bqs="http://www.cellml.org/bqs/1.0#" xmlns:j.0="http://www.example.org/foo/bar/" xmlns:cmeta="http://www.cellml.org/metadata/1.0#" xmlns:j.1="http://imc.org/vCard/3.0#"> <rdf:Description rdf:about="http://www.cellml.org/cellml/1.0"> <dcterms:created rdf:parseType="Resource"> <dcterms:W3CDTF>2009-10-28</dcterms:W3CDTF> </dcterms:created> <dc:creator rdf:parseType="Resource"> <vCard:ORG rdf:parseType="Resource"> <vCard:Orgname>Auckland Bioengineering Institute</vCard:Orgname> </vCard:ORG> <vCard:EMAIL> <j.1:internet> <rdf:value>j.terkildsen@auckland.ac.nz</rdf:value> </j.1:internet> </vCard:EMAIL> <vCard:N rdf:parseType="Resource"> <vCard:Given>Jonna</vCard:Given> <vCard:Family>Terkildsen</vCard:Family> </vCard:N> </dc:creator> </rdf:Description> <j.0:bazfoo/> <rdf:Description rdf:about="http://www.cellml.org/cellml/1.0#mackenzie_1996"> <bqs:reference rdf:parseType="Resource"> <bqs:JournalArticle rdf:parseType="Resource"> <bqs:last_page>32683</bqs:last_page> <bqs:first_page>32678</bqs:first_page> <bqs:volume>271</bqs:volume> <bqs:Journal rdf:parseType="Resource"> <dc:title>Journal of Biological Chemistry</dc:title> </bqs:Journal> <dc:title>Biophysical Characteristics of the Pig Kidney Na/Glucose Cotransporter SGLT2 Reveal a Common Mechanism for SGLT1 and SGLT2</dc:title> <dc:creator> <rdf:Seq> <rdf:li rdf:parseType="Resource"> <bqs:Person rdf:parseType="Resource"> <vCard:N rdf:parseType="Resource"> <vCard:Given>M</vCard:Given> <vCard:Family>Mackenzie</vCard:Family> </vCard:N> </bqs:Person> </rdf:li> <rdf:li rdf:parseType="Resource"> <bqs:Person rdf:parseType="Resource"> <vCard:N rdf:parseType="Resource"> <vCard:Other>F</vCard:Other> <vCard:Other>D</vCard:Other> <vCard:Given>D</vCard:Given> <vCard:Family>Loo</vCard:Family> </vCard:N> </bqs:Person> </rdf:li> <rdf:li rdf:parseType="Resource"> <bqs:Person rdf:parseType="Resource"> <vCard:N rdf:parseType="Resource"> <vCard:Given>M</vCard:Given> <vCard:Family>Panayotova-Heiermann</vCard:Family> </vCard:N> </bqs:Person> </rdf:li> <rdf:li rdf:parseType="Resource"> <bqs:Person rdf:parseType="Resource"> <vCard:N rdf:parseType="Resource"> <vCard:Other>M</vCard:Other> <vCard:Given>E</vCard:Given> <vCard:Family>Wright</vCard:Family> </vCard:N> </bqs:Person> </rdf:li> </rdf:Seq> </dc:creator> </bqs:JournalArticle> <bqs:Pubmed_id>8955098</bqs:Pubmed_id> </bqs:reference> <bqs:reference rdf:parseType="Resource"> <dc:subject rdf:parseType="Resource"> <rdf:value> <rdf:Bag> <rdf:li>Na/glucose cotransport</rdf:li> <rdf:li>ion transport</rdf:li> </rdf:Bag> </rdf:value> <bqs:subject_type>keyword</bqs:subject_type> </dc:subject> </bqs:reference> <cmeta:comment rdf:parseType="Resource"> <dc:creator rdf:parseType="Resource"> <vCard:FN>Jonna Terkildsen</vCard:FN> </dc:creator> <rdf:value>This is the CellML description of the Mackenzie et al 1996 quantitative model of the Na/glucose cotransporter (SGLT2).</rdf:value> </cmeta:comment> <dc:title>The Mackenzie et al 1996 quantitative model of the Na/glucose cotransporter (SGLT2).</dc:title> </rdf:Description> </rdf:RDF> C:\Users\Christopher\Dropbox\Work Related\Annotated Models\MacKenzie_1996\mackenzie_1996.cellml Model of the pig kidney Na1/Glucose cotransporter SGLT2 mackenzie_1996 mackenzie_1996 thompsct@umich.edu Area of spatial entity http://bhi.washington.edu/OPB#OPB_00295 Charge flow rate http://bhi.washington.edu/OPB#OPB_00318 Chemical concentration http://bhi.washington.edu/OPB#OPB_00340 Chemical molar amount http://bhi.washington.edu/OPB#OPB_00425 Electrical potential http://bhi.washington.edu/OPB#OPB_00506 sodium(1+) http://purl.obolibrary.org/obo/CHEBI_29101 methyl alpha-D-glucopyranoside http://purl.obolibrary.org/obo/CHEBI_320061 sodium-glucose transport protein subtype 2 inhibitor http://purl.obolibrary.org/obo/CHEBI_73273 oocyte http://purl.obolibrary.org/obo/CL_0000023 Plasma membrane http://purl.org/sig/ont/fma/fma63841 Portion of cytosol http://purl.org/sig/ont/fma/fma66836 A physical entity that is defined by linking multiple physical entity terms with structural relations. A mathematical and/or logical expression that determines the value of a data structure. An element of a SemSim model that relates to the mathematical or logical representation of physical quantities or qualities. A physical dependency defined in a specific SemSim model. A physical entity defined in a specific SemSim model. A physical process defined in a specific SemSim model. A physical property defined in a specific SemSim model. A computational element that stores numerical and/or logical information related to the model. A data structure whose value is limited to decimals. A data structure whose value is limited to the whole numbers. A data structure unique to JSim's Mathematical Modeling Language (MML) used to set one or more model input values at runtime. A quantitative relation between values of physical properties that occurs over a span of time and that determines the time-course of a physical process. A physics analytical entity that represents the material, energy, and/or information content of material or immaterial, space-occupying continuants. A SemSim model element used to represent, in whole or in part, the physical meaning of a computational element. A physical processural entity that is a flow, or the control of a flow, of energy and/or information amongst physical continuants that are participants in the physical process. A physical attribute of an instance of a physical entity, an occurrence of a physical process, or a physical dependency that has a quantitative value that is measurable by a physical device or method, or is computable from such values. A data structure whose value is limited to the real numbers. A physical dependency that is defined using a term in an external knowledge resource. A physical entity that is defined using a term from an external knowledge resource. A physical process that is defined using a term in an external knowledge resource. A physical property that is defined using a term in an external knowledge resource. A logical expression that constrains, but does not define, the value of a data structure. Example: x > 0. A computational or physical element of a SemSim model. A grouping of one or more data structures that comprise a computationally or physically distinct part of a model or submodel. A physical quantity chosen as a standard in terms of which other physical quantities may be expressed. sodium(1+) http://purl.obolibrary.org/obo/CHEBI_29101 sodium(1+) http://purl.obolibrary.org/obo/CHEBI_29101 methyl alpha-D-glucopyranoside http://purl.obolibrary.org/obo/CHEBI_320061 methyl alpha-D-glucopyranoside http://purl.obolibrary.org/obo/CHEBI_320061 sodium-glucose transport protein subtype 2 inhibitor http://purl.obolibrary.org/obo/CHEBI_73273 oocyte http://purl.obolibrary.org/obo/CL_0000023 Cell culture medium External Na Bound SGLT2 Cotransporter External Na and Substrate Bound SGLT2 Cotransporter External Unbound SGLT2 Cotransporter Glucose transport across cell membrane via SGLT2 cotransporters 1.0 1.0 1.0 1.0 1.0 Internal Na Bound SGLT2 Cotransporter Internal Na and Substrate Bound SGLT2 Cotransporter Internal Unbound SGLT2 Cotransporter NBC_current false false in true Steady state external unbound cotransporter concentration false false in true Steady state external Na bound cotransporter concentration false false in true Steady state external CNaS concentration false false in true Steady state internal CNaS concentration false false in true Steady state internal CNa concentration false false in true Steady state internal unbound cotransporter concentration false false in true Pre-steady state external unbound cotransporter concentration Chemical molar amount http://bhi.washington.edu/OPB#OPB_00425 false false in true Pre-steady state external Na bound cotransporter concentration Chemical molar amount http://bhi.washington.edu/OPB#OPB_00425 false false in true Pre-steady state external CNaS concentration Chemical molar amount http://bhi.washington.edu/OPB#OPB_00425 false false in true Steady state internal CNaS concentration Chemical molar amount http://bhi.washington.edu/OPB#OPB_00425 false false in true Pre-steady state internal CNa concentration Chemical molar amount http://bhi.washington.edu/OPB#OPB_00425 false false in true Pre-steady state internal unbound cotransporter concentration Chemical molar amount http://bhi.washington.edu/OPB#OPB_00425 false false in true Faraday constant false false out true Na glucose steady state current false false out true Na-glucose pre-steady state current Charge flow rate http://bhi.washington.edu/OPB#OPB_00318 false false out true Sodium ion flow into the cytosol Charge flow rate http://bhi.washington.edu/OPB#OPB_00318 false false in true Fraction of membrane electric field between bulk solution and external face Na binding site. false false in true Fraction of membrane electric field between bulk solution and internal face Na binding site. false false in true Fraction of the membrane field sensed by the empty binding on the carrier durring translocation. false false in true Rate for sodium ion binding to external unbound cotransporter. false false in true Rate for external unbound transport site to internal unbound transport site reaction step false false in true Rate for sodium ion release from external bound cotransporter. false false in true Rate for external CNa to internal CNa reaction step. false false in true Rate for external CNaS to internal CNas reaction step false false in true Rate for internal CNaS to external CNaS reaction step false false in true Rate for internal CNa to external CNa reaction step false false in true Rate for release of Sodium from internal Na bound cotransporter false false in true Rate for internal unbound cotransporter binding site to external unbound cotransporter binding site reaction step false false in true Rate for sodium ion release from internal Na bound cotransporter. false false in true Sodium-glucose coupling coefficent false false in true Na valence false false in true Empty transporter valence <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> <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> <minus /> <apply> <times /> <ci>k_34</ci> <ci>C3</ci> </apply> <apply> <times /> <ci>k_43</ci> <ci>C4</ci> </apply> </apply> </apply> </apply> </math> Sodium ion flow into cytosol via SGLT2 cotransporter 1.0 1.0 1.0 1.0 1.0 1.0 C_per_mol -1.0 J_per_K_per_mol -1.0 -1.0 M -1.0 M2 2.0 M2_per_second -1.0 2.0 M_per_second -1.0 ampere true coulomb true joule true kelvin true litre true metre true mole true per_M2_per_second5 -2.0 -5.0 per_M_per_second -1.0 -1.0 per_mol -1.0 per_second -1.0 per_second5 -5.0 second true uA micro um2 2.0 micro umol micro umol_per_second -1.0 true volt environment false false out true Time - Solution Domain Plasma membrane http://purl.org/sig/ont/fma/fma63841 Portion of cytosol http://purl.org/sig/ont/fma/fma66836 ion_concentrations 10e-3 false false out true Intracellular sodium concentration Chemical concentration http://bhi.washington.edu/OPB#OPB_00340 100e-3 false false out true Sodium concentration in cell culture medium Chemical concentration http://bhi.washington.edu/OPB#OPB_00340 -150e-3 false false out true Electrical potential of the cell membrane Electrical potential http://bhi.washington.edu/OPB#OPB_00506 0e-3 false false out true Intracellular glucose concentration Chemical concentration http://bhi.washington.edu/OPB#OPB_00340 1e-3 false false out true Glucose concentration in experimental medium Chemical concentration http://bhi.washington.edu/OPB#OPB_00340 false false in true Time - Solution Domain <math xmlns="http://www.w3.org/1998/Math/MathML"> <apply> <eq /> <apply> <diff /> <bvar> <ci>time</ci> </bvar> <ci>V</ci> </apply> <cn type="e-notation" xmlns:cellml="http://www.cellml.org/cellml/1.0#" cellml:units="volt_per_second"> 100 <sep /> -6 </cn> </apply> </math> kinetic_equations 3.923e-9 false false out true Chemical molar amount http://bhi.washington.edu/OPB#OPB_00425 1.016e-7 false false out true Pre-steady state external Na bound cotransporter concentration Chemical molar amount http://bhi.washington.edu/OPB#OPB_00425 1.515e-8 false false out true Pre-steady state external CNaS concentration Chemical molar amount http://bhi.washington.edu/OPB#OPB_00425 8.912e-10 false false out true Steady state internal CNaS concentration Chemical molar amount http://bhi.washington.edu/OPB#OPB_00425 1.485e-8 false false out true Pre-steady state internal CNa concentration Chemical molar amount http://bhi.washington.edu/OPB#OPB_00425 false false out true Pre-steady state internal unbound cotransporter concentration Chemical molar amount http://bhi.washington.edu/OPB#OPB_00425 3.06798e-9 false false out true Chemical molar amount http://bhi.washington.edu/OPB#OPB_00425 false false in true Density of functional SGLT2 carriers in the oocyte membrane Chemical molar amount http://bhi.washington.edu/OPB#OPB_00425 false false in true Rate for sodium ion binding to external unbound cotransporter. false false in true Rate for external unbound transport site to internal unbound transport site reaction step false false in true Rate for sodium ion release from external bound cotransporter. false false in true Rate for binding of glucose to external CNa binding site false false in true Rate for external CNa to internal CNa reaction step. false false in true Rate for release of glucose from external CNaS false false in true Rate for external CNaS to internal CNas reaction step false false in true Rate for internal CNaS to external CNaS reaction step false false in true Rate constant for release of glucose from internal CNaS false false in true Rate for internal CNa to external CNa reaction step false false in true Rate for binding of glucose to internal CNa binding site false false in true Rate for release of Sodium from internal Na bound cotransporter false false in true Rate for internal unbound cotransporter binding site to external unbound cotransporter binding site reaction step false false in true Rate for sodium ion release from internal Na bound cotransporter. false false in true Time - Solution Domain <math xmlns="http://www.w3.org/1998/Math/MathML"> <apply> <eq /> <apply> <diff /> <bvar> <ci>time</ci> </bvar> <ci>C_1</ci> </apply> <apply> <minus /> <apply> <plus /> <apply> <times /> <ci>k_21</ci> <ci>C_2</ci> </apply> <apply> <times /> <ci>k_61</ci> <ci>C_6</ci> </apply> </apply> <apply> <times /> <apply> <plus /> <ci>k_12</ci> <ci>k_16</ci> </apply> <ci>C_1</ci> </apply> </apply> </apply> <apply> <eq /> <apply> <diff /> 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<ci>k_45</ci> <ci>k_43</ci> </apply> <ci>C_4</ci> </apply> </apply> </apply> <apply> <eq /> <apply> <diff /> <bvar> <ci>time</ci> </bvar> <ci>C_5</ci> </apply> <apply> <minus /> <apply> <plus /> <apply> <times /> <ci>k_45</ci> <ci>C_4</ci> </apply> <apply> <times /> <ci>k_65</ci> <ci>C_6</ci> </apply> <apply> <times /> <ci>k_25</ci> <ci>C_2</ci> </apply> </apply> <apply> <times /> <apply> <plus /> <ci>k_54</ci> <ci>k_52</ci> <ci>k_56</ci> </apply> <ci>C_5</ci> </apply> </apply> </apply> <apply> <eq /> <apply> <diff /> <bvar> <ci>time</ci> </bvar> <ci>C_6_temp</ci> </apply> <apply> <minus /> <apply> <plus /> <apply> <times /> <ci>k_16</ci> <ci>C_1</ci> </apply> <apply> <times /> <ci>k_56</ci> <ci>C_5</ci> </apply> </apply> <apply> <times /> <apply> <plus /> <ci>k_61</ci> <ci>k_65</ci> </apply> <ci>C_6_temp</ci> </apply> </apply> </apply> <apply> <eq /> <ci>C_6</ci> <apply> <minus /> <ci>C_T</ci> <apply> <plus /> <ci>C_1</ci> <ci>C_2</ci> <ci>C_3</ci> <ci>C_4</ci> <ci>C_5</ci> </apply> </apply> </apply> </math> king_altman_states false false out true Chemical molar amount http://bhi.washington.edu/OPB#OPB_00425 false false true false false out true Steady state external Na bound cotransporter concentration false false true false false out true Steady state external CNaS concentration false false true false false out true Steady state internal CNaS concentration false false true false false out true Steady state internal CNa concentration false false true false false out true Steady state internal unbound cotransporter concentration false false true false false in true Density of functional SGLT2 carriers in the oocyte membrane Chemical molar amount http://bhi.washington.edu/OPB#OPB_00425 false false true false false in true Rate for sodium ion binding to external unbound cotransporter. false false in true Rate for external unbound transport site to internal unbound transport site reaction step false false in true Rate for sodium ion release from external bound cotransporter. false false in true Rate for binding of glucose to external CNa binding site false false in true Rate for external CNa to internal CNa reaction step. false false in true Rate for release of glucose from external CNaS false false in true Rate for external CNaS to internal CNas reaction step false false in true Rate for internal CNaS to external CNaS reaction step false false in true Rate constant for release of glucose from internal CNaS false false in true Rate for internal CNa to external CNa reaction step false false in true Rate for binding of glucose to internal CNa binding site false false in true Rate for release of Sodium from internal Na bound cotransporter false false in true Rate for internal unbound cotransporter binding site to external unbound cotransporter binding site reaction step false false in true Rate for sodium ion release from internal Na bound cotransporter. <math xmlns="http://www.w3.org/1998/Math/MathML"> <apply> <eq /> <ci>C1_sum</ci> <apply> <plus /> <apply> <times /> <ci>k_21</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_21</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_21</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_21</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_25</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_23</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_21</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_25</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_21</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_25</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_21</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_21</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_54</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_21</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_21</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_21</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_52</ci> <ci>k_61</ci> </apply> </apply> </apply> <apply> <eq /> <ci>C2_sum</ci> <apply> <plus /> <apply> <times /> <ci>k_16</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_54</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_52</ci> <ci>k_61</ci> </apply> </apply> </apply> <apply> <eq /> <ci>C3_sum</ci> <apply> <plus /> <apply> <times /> <ci>k_16</ci> <ci>k_25</ci> <ci>k_43</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_25</ci> <ci>k_43</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_23</ci> <ci>k_43</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_23</ci> <ci>k_43</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_21</ci> <ci>k_43</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_23</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_23</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_23</ci> <ci>k_43</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_23</ci> <ci>k_43</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_23</ci> <ci>k_45</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_23</ci> <ci>k_43</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_25</ci> <ci>k_43</ci> <ci>k_54</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_23</ci> <ci>k_43</ci> <ci>k_54</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_23</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_23</ci> <ci>k_43</ci> <ci>k_52</ci> <ci>k_61</ci> </apply> </apply> </apply> <apply> <eq /> <ci>C4_sum</ci> <apply> <plus /> <apply> <times /> <ci>k_16</ci> <ci>k_25</ci> <ci>k_34</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_25</ci> <ci>k_34</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_23</ci> <ci>k_34</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_23</ci> <ci>k_34</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_21</ci> <ci>k_34</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_25</ci> <ci>k_32</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_25</ci> <ci>k_32</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_21</ci> <ci>k_32</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_23</ci> <ci>k_34</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_23</ci> <ci>k_34</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_23</ci> <ci>k_34</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_25</ci> <ci>k_34</ci> <ci>k_54</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_23</ci> <ci>k_34</ci> <ci>k_54</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_25</ci> <ci>k_32</ci> <ci>k_54</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_23</ci> <ci>k_34</ci> <ci>k_52</ci> <ci>k_61</ci> </apply> </apply> </apply> <apply> <eq /> <ci>C5_sum</ci> <apply> <plus /> <apply> <times /> <ci>k_16</ci> <ci>k_25</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_25</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_23</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_23</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_21</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_25</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_25</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_21</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_25</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_25</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_21</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_25</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_23</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_25</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_25</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_61</ci> </apply> </apply> </apply> <apply> <eq /> <ci>C6_sum</ci> <apply> <plus /> <apply> <times /> <ci>k_16</ci> <ci>k_25</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_56</ci> </apply> <apply> <times /> <ci>k_12</ci> <ci>k_25</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_56</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_23</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_56</ci> </apply> <apply> <times /> <ci>k_12</ci> <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> parameters false false out true Density of functional SGLT2 carriers in the oocyte membrane Chemical molar amount http://bhi.washington.edu/OPB#OPB_00425 96485.34 false false out true Faraday constant 6.022e23 false false true Avogadro's number 8.4e10 false false true 8.314 false false out true Boltzmann constant 310 false false out true Temperature false false in true Electrical potential of the cell membrane Electrical potential http://bhi.washington.edu/OPB#OPB_00506 false false out true Fraction of membrane electric field between bulk solution and external face Na binding site. 0 false false out true Fraction of membrane electric field between bulk solution and internal face Na binding site. 1e6 false false true Area of spatial entity http://bhi.washington.edu/OPB#OPB_00295 0.7 false false out true Fraction of the membrane field sensed by the empty binding on the carrier durring translocation. 20000 false false out true Rate constant for sodium ion binding to external unbound cotransporter. 100 false false out true Rate constant for external unbound transport site to internal unbound transport site reaction step 400 false false out true Rate constant for sodium ion release from external bound cotransporter. 1e4 false false out true Rate constant for binding of glucose to external CNa binding site 0.01 false false out true Rate constant for external CNa to internal CNa reaction step. 20 false false out true Rate constant for release of glucose from external CNaS 50 false false out true Rate constant for external CNaS to internal CNas reaction step 50 false false out true Rate constant for internal CNaS to external CNaS reaction step 800 false false out true Rate constant for release of glucose from internal CNaS 0.17 false false out true Rate constant for internal CNa to external CNa reaction step 6.7e6 false false out true Rate constant for binding of glucose to internal CNa binding site 48 false false out true 35 false false out true Rate constant for internal unbound cotransporter binding site to external unbound cotransporter binding site reaction step 50 false false out true Rate constant for sodium ion release from internal Na bound cotransporter. false false out true Relationship between membrane voltage and F*R/T. 1 false false out true Sodium-glucose coupling coefficent 1 false false out true Na valence -1 false false out true Empty transporter valence <math xmlns="http://www.w3.org/1998/Math/MathML"> <apply> <eq /> <ci>alpha_p</ci> <apply> <minus /> <apply> <minus /> <cn xmlns:cellml="http://www.cellml.org/cellml/1.0#" 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 type="e-notation" xmlns:cellml="http://www.cellml.org/cellml/1.0#" cellml:units="umol_per_mol"> 1 <sep /> 6 </cn> <ci>N_C</ci> </apply> <ci>N_Avo</ci> </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> phenomonological_constants false false in true Density of functional SGLT2 carriers in the oocyte membrane Chemical molar amount http://bhi.washington.edu/OPB#OPB_00425 false false in true Faraday constant false false out true Maximum sodium current across cell membrane false false out true Maximum glucose current across cell membrane false false out true Experimental substrate sodium concentration for half maximal current. false false out true Experimental substrate glucose concentration for half maximal current. false false in true Sodium concentration in cell culture medium Chemical concentration http://bhi.washington.edu/OPB#OPB_00340 false false out true false false out true false false out true false false out true false false out true Membrane dislocation false false in true Glucose concentration in experimental medium Chemical concentration http://bhi.washington.edu/OPB#OPB_00340 false false in true Rate constant for binding of glucose to external CNa binding site false false in true Rate for sodium ion binding to external unbound cotransporter. false false in true Rate for external unbound transport site to internal unbound transport site reaction step false false in true Rate for sodium ion release from external bound cotransporter. false false in true Rate for binding of glucose to external CNa binding site false false in true Rate for external CNa to internal CNa reaction step. false false in true Rate for release of glucose from external CNaS false false in true Rate for external CNaS to internal CNas reaction step false false in true Rate for internal CNaS to external CNaS reaction step false false in true Rate constant for release of glucose from internal CNaS false false in true Rate for internal CNa to external CNa reaction step false false in true Rate for binding of glucose to internal CNa binding site false false in true Rate for release of Sodium from internal Na bound cotransporter false false in true Rate for internal unbound cotransporter binding site to external unbound cotransporter binding site reaction step false false in true Rate for sodium ion release from internal Na bound cotransporter. false false in true Sodium concentration independent rate for sodium ion binding to external unbound cotransporter. false false out true Sum of all enzymes. false false out true <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 xmlns:cellml="http://www.cellml.org/cellml/1.0#" 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> <divide /> <cn xmlns:cellml="http://www.cellml.org/cellml/1.0#" cellml:units="dimensionless">1</cn> <ci>lambda</ci> </apply> <apply> <plus /> <apply> <times /> <ci>k0_23</ci> <ci>k_16</ci> <ci>k_43</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k0_23</ci> <ci>k_16</ci> <ci>k_34</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k0_23</ci> <ci>k_16</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k0_23</ci> <ci>k_16</ci> <ci>k_43</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k0_23</ci> <ci>k_16</ci> <ci>k_34</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k0_23</ci> <ci>k_16</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k0_23</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k0_23</ci> <ci>k_16</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_56</ci> </apply> </apply> </apply> </apply> <apply> <eq /> <ci>alpha</ci> <apply> <times /> <apply> <divide /> <cn xmlns:cellml="http://www.cellml.org/cellml/1.0#" cellml:units="dimensionless">1</cn> <ci>lambda</ci> </apply> <apply> <plus /> <apply> <times /> <ci>k_21</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_25</ci> <ci>k_43</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_21</ci> <ci>k_43</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_25</ci> <ci>k_34</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_21</ci> <ci>k_34</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_25</ci> <ci>k_32</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_21</ci> <ci>k_32</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_21</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_21</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_21</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_25</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_21</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_25</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_21</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_25</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_21</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_25</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_21</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_25</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_21</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_25</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_21</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_21</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_54</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_21</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_21</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_21</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_52</ci> <ci>k_61</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_25</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_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_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> <apply> <eq /> <ci>gamma</ci> <apply> <times /> <apply> <divide /> <cn xmlns:cellml="http://www.cellml.org/cellml/1.0#" cellml:units="dimensionless">1</cn> <ci>lambda</ci> </apply> <apply> <plus /> <apply> <times /> <ci>k_16</ci> <ci>k_21</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_54</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_21</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_21</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> <apply> <times /> <ci>k_16</ci> <ci>k_21</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_52</ci> <ci>k_65</ci> </apply> </apply> </apply> </apply> <apply> <eq /> <ci>phi</ci> <apply> <times /> <apply> <divide /> <cn xmlns:cellml="http://www.cellml.org/cellml/1.0#" cellml:units="dimensionless">1</cn> <ci>lambda</ci> </apply> <apply> <minus /> <apply> <minus /> <apply> <times /> <apply> <minus /> <ci>ks_12</ci> </apply> <ci>k_25</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_25</ci> <ci>k_32</ci> <ci>k_45</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> </apply> <apply> <times /> <ci>ks_12</ci> <ci>k_25</ci> <ci>k_32</ci> <ci>k_43</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> </apply> </apply> </apply> <apply> <eq /> <ci>epsilon</ci> <apply> <times /> <apply> <divide /> <cn xmlns:cellml="http://www.cellml.org/cellml/1.0#" cellml:units="dimensionless">1</cn> <ci>lambda</ci> </apply> <apply> <minus /> <ci>ks_12</ci> </apply> <ci>k0_23</ci> <ci>k_34</ci> <ci>k_45</ci> <ci>k_56</ci> <ci>k_61</ci> </apply> </apply> <apply> <eq /> <ci>Imax_gluc</ci> <apply> <divide /> <apply> <times /> <cn xmlns:cellml="http://www.cellml.org/cellml/1.0#" cellml:units="dimensionless">2</cn> <ci>F</ci> <ci>C_T</ci> <ci>epsilon</ci> <ci>Na_o</ci> </apply> <apply> <plus /> <ci>beta</ci> <ci>Na_o</ci> </apply> </apply> </apply> <apply> <eq /> <ci>Imax_Na</ci> <apply> <divide /> <apply> <times /> <cn xmlns:cellml="http://www.cellml.org/cellml/1.0#" cellml:units="dimensionless">2</cn> <ci>F</ci> <ci>C_T</ci> <apply> <plus /> <ci>phi</ci> <apply> <times /> <ci>epsilon</ci> <ci>glucose_o</ci> </apply> </apply> </apply> <apply> <plus /> <ci>chi</ci> <ci>glucose_o</ci> </apply> </apply> </apply> <apply> <eq /> <ci>Khalf_gluc</ci> <apply> <divide /> <apply> <plus /> <ci>alpha</ci> <apply> <times /> <ci>chi</ci> <ci>Na_o</ci> </apply> </apply> <apply> <plus /> <ci>beta</ci> <ci>Na_o</ci> </apply> </apply> </apply> <apply> <eq /> <ci>Khalf_Na</ci> <apply> <divide /> <apply> <plus /> <ci>alpha</ci> <apply> <times /> <ci>beta</ci> <ci>glucose_o</ci> </apply> </apply> <apply> <plus /> <ci>chi</ci> <ci>glucose_o</ci> </apply> </apply> </apply> </math> rate_constants false false in true Density of functional SGLT2 carriers in the oocyte membrane Chemical molar amount http://bhi.washington.edu/OPB#OPB_00425 false false in true Intracellular sodium concentration Chemical concentration http://bhi.washington.edu/OPB#OPB_00340 false false in true Sodium concentration in cell culture medium Chemical concentration http://bhi.washington.edu/OPB#OPB_00340 false false in true Fraction of membrane electric field between bulk solution and external face Na binding site. false false in true Fraction of membrane electric field between bulk solution and internal face Na binding site. false false in true Fraction of the membrane field sensed by the empty binding on the carrier durring translocation. false false in true Intracellular glucose concentration Chemical concentration http://bhi.washington.edu/OPB#OPB_00340 false false in true Glucose concentration in experimental medium Chemical concentration http://bhi.washington.edu/OPB#OPB_00340 false false in true Rate constant for sodium ion binding to external unbound cotransporter. false false in true Rate constant for external unbound transport site to internal unbound transport site reaction step false false in true Rate constant for sodium ion release from external bound cotransporter. false false in true Rate constant for binding of glucose to external CNa binding site false false in true Rate constant for external CNa to internal CNa reaction step. false false in true Rate constant for release of glucose from external CNaS false false in true Rate constant for external CNaS to internal CNas reaction step false false in true Rate constant for internal CNaS to external CNaS reaction step false false in true Rate constant for release of glucose from internal CNaS false false in true Rate constant for internal CNa to external CNa reaction step false false in true Rate constant for binding of glucose to internal CNa binding site false false out true Glucose independent rate constant for binding of glucose to internal CNa binding site false false in true Rate constant for release of Sodium from internal Na bound cotransporter false false in true Rate constant for internal unbound cotransporter binding site to external unbound cotransporter binding site reaction step false false in true Rate constant for sodium ion release from internal Na bound cotransporter. false false out true Rate for sodium ion binding to external unbound cotransporter. false false out true Rate for external unbound transport site to internal unbound transport site reaction step false false out true Rate for sodium ion release from external bound cotransporter. false false out true Rate for binding of glucose to external CNa binding site false false out true Rate for external CNa to internal CNa reaction step. false false out true Rate for release of glucose from external CNaS false false out true Rate for external CNaS to internal CNas reaction step false false out true Rate for internal CNaS to external CNaS reaction step false false out true Rate for release of glucose from internal CNaS false false out true Rate for internal CNa to external CNa reaction step false false out true Glucose concentration independent rate for internal CNa to external CNa reaction step false false out true Rate for binding of glucose to internal CNa binding site false false out true Rate for release of Sodium from internal Na bound cotransporter false false out true Rate for internal unbound cotransporter binding site to external unbound cotransporter binding site reaction step false false out true Rate for sodium ion release from internal Na bound cotransporter. false false out true Sodium concentration independent rate for sodium ion binding to external unbound cotransporter. false false in true Relationship between membrane voltage and F*R/T. false false in true Sodium-glucose coupling coefficent false false in true Na valence false false in true Empty transporter valence <math xmlns="http://www.w3.org/1998/Math/MathML"> <apply> <eq /> <ci>ks_12</ci> <apply> <times /> <ci>k0_12</ci> <apply> <exp /> <apply> <divide /> <apply> <times /> <apply> <minus /> <ci>n</ci> </apply> <ci>alpha_p</ci> <ci>mu</ci> </apply> <cn xmlns:cellml="http://www.cellml.org/cellml/1.0#" cellml:units="dimensionless">2</cn> </apply> </apply> </apply> </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 /> <apply> <times /> <ci>n</ci> <ci>z_Na</ci> <ci>alpha_p</ci> <ci>mu</ci> </apply> <cn xmlns:cellml="http://www.cellml.org/cellml/1.0#" 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 /> <apply> <plus /> <ci>z_c</ci> <ci>n</ci> </apply> </apply> <ci>delta</ci> <ci>mu</ci> </apply> <cn xmlns:cellml="http://www.cellml.org/cellml/1.0#" 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> </apply> <ci>delta</ci> <ci>mu</ci> </apply> <cn xmlns:cellml="http://www.cellml.org/cellml/1.0#" 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 xmlns:cellml="http://www.cellml.org/cellml/1.0#" 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 xmlns:cellml="http://www.cellml.org/cellml/1.0#" cellml:units="dimensionless">2</cn> </apply> </apply> </apply> </apply> <apply> <eq /> <ci>k_56</ci> <apply> <times /> <ci>k0_56</ci> <apply> <exp /> <apply> <divide /> <apply> <times /> <apply> <minus /> <ci>n</ci> </apply> <ci>z_Na</ci> <ci>alpha_pp</ci> <ci>mu</ci> </apply> <cn xmlns:cellml="http://www.cellml.org/cellml/1.0#" cellml:units="dimensionless">2</cn> </apply> </apply> </apply> </apply> <apply> <eq /> <ci>k_65</ci> <apply> <times /> <ci>k0_65</ci> <apply> <power /> <ci>Na_i</ci> <ci>n</ci> </apply> <apply> <exp /> <apply> <divide /> <apply> <times /> <ci>n</ci> <ci>z_Na</ci> <ci>alpha_pp</ci> <ci>mu</ci> </apply> <cn xmlns:cellml="http://www.cellml.org/cellml/1.0#" cellml:units="dimensionless">2</cn> </apply> </apply> </apply> </apply> <apply> <eq /> <ci>k_61</ci> <apply> <times /> <ci>k0_61</ci> <apply> <exp /> <apply> <divide /> <apply> <times /> <ci>z_c</ci> <ci>delta</ci> <ci>mu</ci> </apply> <cn xmlns:cellml="http://www.cellml.org/cellml/1.0#" cellml:units="dimensionless">2</cn> </apply> </apply> </apply> </apply> <apply> <eq /> <ci>k_16</ci> <apply> <times /> <ci>k0_16</ci> <apply> <exp /> <apply> <divide /> <apply> <times /> <apply> <minus /> <ci>z_c</ci> </apply> <ci>delta</ci> <ci>mu</ci> </apply> <cn xmlns:cellml="http://www.cellml.org/cellml/1.0#" cellml:units="dimensionless">2</cn> </apply> </apply> </apply> </apply> <apply> <eq /> <ci>k_52</ci> <apply> <divide /> <apply> <times /> <ci>k0_12</ci> <ci>k_25</ci> <ci>k0_56</ci> <ci>k0_61</ci> </apply> <apply> <times /> <ci>k0_21</ci> <ci>k0_16</ci> <ci>k0_65</ci> </apply> </apply> </apply> <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> </apply> <apply> <times /> <ci>k_43</ci> <ci>k_32</ci> <ci>k_25</ci> </apply> </apply> </apply> </math>