Rendering of the source text

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				<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>
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			<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.
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       <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>
				
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								<title>model diagram</title>
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					<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>
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                 <ci>k_25</ci>
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                 <ci>k_34</ci>
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                 <ci>k_32</ci>
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                 <ci>k_32</ci>
                 <ci>k_45</ci>
                 <ci>k_56</ci>
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                 <ci>k_25</ci>
                 <ci>k_32</ci>
                 <ci>k_43</ci>
                 <ci>k_56</ci>
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                 <ci>k_43</ci>
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                 <ci>k_21</ci>
                 <ci>k_32</ci>
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                 <ci>k_56</ci>
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                 <times/>
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                 <ci>k_21</ci>
                 <ci>k_32</ci>
                 <ci>k_43</ci>
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                 <times/>
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                 <ci>k_21</ci>
                 <ci>k_34</ci>
                 <ci>k_45</ci>
                 <ci>k_52</ci>
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                 <ci>k_43</ci>
                 <ci>k_52</ci>
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           </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>
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        <apply>
           <eq/>
           <ci>C1</ci>
           <apply>
              <divide/>
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                 <times/>
                 <ci>C_T</ci>
                 <ci>C1_sum</ci>
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              <ci>C_sum</ci>
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        <apply>
           <eq/>
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              <divide/>
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                 <times/>
                 <ci>C_T</ci>
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              <ci>C_sum</ci>
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        <apply>
           <eq/>
           <ci>C3</ci>
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              <divide/>
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                 <times/>
                 <ci>C_T</ci>
                 <ci>C3_sum</ci>
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              <ci>C_sum</ci>
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        <apply>
           <eq/>
           <ci>C4</ci>
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              <divide/>
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                 <times/>
                 <ci>C_T</ci>
                 <ci>C4_sum</ci>
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              <ci>C_sum</ci>
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        <apply>
           <eq/>
           <ci>C5</ci>
           <apply>
              <divide/>
              <apply>
                 <times/>
                 <ci>C_T</ci>
                 <ci>C5_sum</ci>
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              <ci>C_sum</ci>
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        <apply>
           <eq/>
           <ci>C6</ci>
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              <divide/>
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                 <times/>
                 <ci>C_T</ci>
                 <ci>C6_sum</ci>
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     <variable name="C_5" units="umol" public_interface="in"/>
     <variable name="C_6" units="umol" public_interface="in"/>
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     <variable name="delta" units="dimensionless" public_interface="in"/>
     <variable name="alpha_p" units="dimensionless" public_interface="in"/>
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     <variable name="z_Na" units="dimensionless" public_interface="in"/>
     <variable name="F" units="C_per_mol" public_interface="in"/>
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     <variable name="I_NaGl_SS" units="uA" public_interface="out"/>
     <variable name="J_Na" units="umol_per_second" public_interface="out"/>
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                    <times/>
                    <ci>n</ci>
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                       <minus/>
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                          <times/>
                          <ci>k_12</ci>
                          <ci>C_1</ci>
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                          <times/>
                          <ci>k_21</ci>
                          <ci>C_2</ci>
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                       <minus/>
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                          <times/>
                          <ci>k_16</ci>
                          <ci>C_1</ci>
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                       <apply>
                          <times/>
                          <ci>k_61</ci>
                          <ci>C_6</ci>
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                    <times/>
                    <ci>n</ci>
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                       <minus/>
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                          <times/>
                          <ci>k_56</ci>
                          <ci>C_5</ci>
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                          <times/>
                          <ci>k_65</ci>
                          <ci>C_6</ci>
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              <times/>
              <apply>
                 <minus/>
                 <ci>F</ci>
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              <apply>
                 <plus/>
                 <apply>
                    <times/>
                    <ci>z_c</ci>
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                       <minus/>
                       <apply>
                          <times/>
                          <ci>k_16</ci>
                          <ci>C1</ci>
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                       <apply>
                          <times/>
                          <ci>k_61</ci>
                          <ci>C6</ci>
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                    <times/>
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                          <times/>
                          <ci>z_Na</ci>
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                    <apply>
                       <minus/>
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                          <times/>
                          <ci>k_25</ci>
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                          <times/>
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                          <ci>C5</ci>
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                          <times/>
                          <ci>z_Na</ci>
                          <ci>n</ci>
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                          <times/>
                          <ci>k_34</ci>
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                       <apply>
                          <times/>
                          <ci>k_43</ci>
                          <ci>C4</ci>
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        <apply>
           <eq/>
           <ci>J_Na</ci>
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              <plus/>
              <apply>
                 <minus/>
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                    <times/>
                    <ci>k_25</ci>
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                 <apply>
                    <times/>
                    <ci>k_52</ci>
                    <ci>C5</ci>
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                    <times/>
                    <ci>k_34</ci>
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                 <apply>
                    <times/>
                    <ci>k_43</ci>
                    <ci>C4</ci>
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     <variable name="chi" units="M" public_interface="out"/>
     <variable name="alpha" units="M2" public_interface="out"/>
     <variable name="beta" units="M" public_interface="out"/>
     <variable name="gamma" units="M2_per_second" public_interface="out"/>
     <variable name="phi" units="M_per_second" public_interface="out"/>
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     <variable name="k_21" units="per_second" public_interface="in"/>
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     <variable name="k_16" units="per_second" public_interface="in"/>
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     <variable name="k0_23" units="per_M_per_second" public_interface="in"/>
     <variable name="C_T" units="umol" public_interface="in"/>
     <variable name="F" units="C_per_mol" public_interface="in"/>
     <variable name="Na_o" units="M" public_interface="in"/>
     <variable name="glucose_o" units="M" public_interface="in"/>
     <variable name="Imax_Na" units="uA" public_interface="out"/>
     <variable name="Imax_gluc" units="uA" public_interface="out"/>
     <variable name="Khalf_Na" units="M" public_interface="out"/>
     <variable name="Khalf_gluc" units="M" public_interface="out"/>
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           <eq/>
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              <plus/>
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                 <times/>
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                 <times/>
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                 <ci>k_45</ci>
                 <ci>k_65</ci>
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                 <ci>k_43</ci>
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              <times/>
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                 <plus/>
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                    <times/>
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                    <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 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 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>
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                    <times/>
                    <ci>k_16</ci>
                    <ci>k_21</ci>
                    <ci>k_34</ci>
                    <ci>k_45</ci>
                    <ci>k_56</ci>
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                 <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 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 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 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 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 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>
  </component>
  <connection xmlns="http://www.cellml.org/cellml/1.0#">
     <map_components component_1="rate_constants" component_2="ion_concentrations"/>
     <map_variables variable_1="Na_i" variable_2="Na_i"/>
     <map_variables variable_1="Na_o" variable_2="Na_o"/>
     <map_variables variable_1="glucose_i" variable_2="glucose_i"/>
     <map_variables variable_1="glucose_o" variable_2="glucose_o"/>
  </connection>
  <connection xmlns="http://www.cellml.org/cellml/1.0#">
     <map_components component_1="rate_constants" component_2="kinetic_equations"/>
     <map_variables variable_1="k_12" variable_2="k_12"/>
     <map_variables variable_1="k_21" variable_2="k_21"/>
     <map_variables variable_1="k_23" variable_2="k_23"/>
     <map_variables variable_1="k_32" variable_2="k_32"/>
     <map_variables variable_1="k_25" variable_2="k_25"/>
     <map_variables variable_1="k_52" variable_2="k_52"/>
     <map_variables variable_1="k_34" variable_2="k_34"/>
     <map_variables variable_1="k_43" variable_2="k_43"/>
     <map_variables variable_1="k_45" variable_2="k_45"/>
     <map_variables variable_1="k_54" variable_2="k_54"/>
     <map_variables variable_1="k_56" variable_2="k_56"/>
     <map_variables variable_1="k_65" variable_2="k_65"/>
     <map_variables variable_1="k_61" variable_2="k_61"/>
     <map_variables variable_1="k_16" variable_2="k_16"/>
  </connection>
  <connection xmlns="http://www.cellml.org/cellml/1.0#">
     <map_components component_1="rate_constants" component_2="king_altman_states"/>
     <map_variables variable_1="k_12" variable_2="k_12"/>
     <map_variables variable_1="k_21" variable_2="k_21"/>
     <map_variables variable_1="k_23" variable_2="k_23"/>
     <map_variables variable_1="k_32" variable_2="k_32"/>
     <map_variables variable_1="k_25" variable_2="k_25"/>
     <map_variables variable_1="k_52" variable_2="k_52"/>
     <map_variables variable_1="k_34" variable_2="k_34"/>
     <map_variables variable_1="k_43" variable_2="k_43"/>
     <map_variables variable_1="k_45" variable_2="k_45"/>
     <map_variables variable_1="k_54" variable_2="k_54"/>
     <map_variables variable_1="k_56" variable_2="k_56"/>
     <map_variables variable_1="k_65" variable_2="k_65"/>
     <map_variables variable_1="k_61" variable_2="k_61"/>
     <map_variables variable_1="k_16" variable_2="k_16"/>
  </connection>
  <connection xmlns="http://www.cellml.org/cellml/1.0#">
     <map_components component_1="rate_constants" component_2="NBC_current"/>
     <map_variables variable_1="k_12" variable_2="k_12"/>
     <map_variables variable_1="k_21" variable_2="k_21"/>
     <map_variables variable_1="k_56" variable_2="k_56"/>
     <map_variables variable_1="k_65" variable_2="k_65"/>
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     <dc:title>
       The Mackenzie et al 1996 quantitative model of the Na/glucose cotransporter (SGLT2).
     </dc:title>

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         This is the CellML description of the Mackenzie et al 1996 quantitative model of the Na/glucose cotransporter (SGLT2).
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          Biophysical Characteristics of the Pig Kidney Na/Glucose Cotransporter SGLT2 Reveal a Common Mechanism for SGLT1 and SGLT2
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           <dc:title>Journal of Biological Chemistry</dc:title>
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</model>
Source
Derived from workspace Mackenzie, Loo, Panayotova-Heiermann and Wright, 1996 at changeset b0a32e5099ae.
Collaboration
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License
The terms of use/license for this work is unspecified.