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BIOMD0000000633 - Bulik2016 - Regulation of hepatic glucose metabolism

 

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Reference Publication
Publication ID: 26935066
Bulik S, Holzhütter HG, Berndt N.
The relative importance of kinetic mechanisms and variable enzyme abundances for the regulation of hepatic glucose metabolism--insights from mathematical modeling.
BMC Biol. 2016 Mar; 14: 15
Charité - Universitätsmedizin Berlin, Institute of Biochemistry, Computational Systems Biochemistry Group, Charitéplatz 1, 10117, Berlin, Germany. sascha.bulik@charite.de.  [more]
Model
Original Model: BIOMD0000000633.origin
Submitter: Vijayalakshmi Chelliah
Submission ID: MODEL1603030000
Submission Date: 03 Mar 2016 13:18:42 UTC
Last Modification Date: 25 Apr 2017 16:19:44 UTC
Creation Date: 21 Apr 2017 14:55:58 UTC
Encoders:  Ryan Gutenkunst
   Alyssa Fortier
   Yingli Hillis
   Fuqiang Tan
   Brianna Uhall
   Gary Tyree
set #1
bqbiol:hasProperty Gene Ontology regulation of glucose metabolic process
Gene Ontology protein phosphorylation
Gene Ontology transmembrane transport
Mathematical Modelling Ontology constraint-based model
bqbiol:hasTaxon Taxonomy Rattus norvegicus
Notes
Bulik2016 - Regulation of hepatic glucose metabolism

This model is described in the article:

Bulik S, Holzhütter HG, Berndt N.
BMC Biol. 2016 Mar; 14: 15

Abstract:

Adaptation of the cellular metabolism to varying external conditions is brought about by regulated changes in the activity of enzymes and transporters. Hormone-dependent reversible enzyme phosphorylation and concentration changes of reactants and allosteric effectors are the major types of rapid kinetic enzyme regulation, whereas on longer time scales changes in protein abundance may also become operative. Here, we used a comprehensive mathematical model of the hepatic glucose metabolism of rat hepatocytes to decipher the relative importance of different regulatory modes and their mutual interdependencies in the hepatic control of plasma glucose homeostasis.Model simulations reveal significant differences in the capability of liver metabolism to counteract variations of plasma glucose in different physiological settings (starvation, ad libitum nutrient supply, diabetes). Changes in enzyme abundances adjust the metabolic output to the anticipated physiological demand but may turn into a regulatory disadvantage if sudden unexpected changes of the external conditions occur. Allosteric and hormonal control of enzyme activities allow the liver to assume a broad range of metabolic states and may even fully reverse flux changes resulting from changes of enzyme abundances alone. Metabolic control analysis reveals that control of the hepatic glucose metabolism is mainly exerted by enzymes alone, which are differently controlled by alterations in enzyme abundance, reversible phosphorylation, and allosteric effects.In hepatic glucose metabolism, regulation of enzyme activities by changes of reactants, allosteric effects, and reversible phosphorylation is equally important as changes in protein abundance of key regulatory enzymes.

To the extent possible under law, all copyright and related or neighbouring rights to this encoded model have been dedicated to the public domain worldwide. Please refer to CC0 Public Domain Dedication for more information.

Model
Publication ID: 26935066 Submission Date: 03 Mar 2016 13:18:42 UTC Last Modification Date: 25 Apr 2017 16:19:44 UTC Creation Date: 21 Apr 2017 14:55:58 UTC
Mathematical expressions
Reactions
ALD EN FBP1 GAPDH
GK GLCT_ER GLUT2 GP
G6P_ER GPI G1PI G6PT_ER
GS LACT LDH MALT
MDH MDH_mito NDK_GTP NDK_UTP
NDK_GTP_mito PC PEPCK PEPCK_mito
PEPT PFK1 PFK2 FBP2
PGK PGM PK PyrMalT
PyrT TPI UGT  
Rules
Assignment Rule (variable: ins) Assignment Rule (variable: glucagon) Assignment Rule (variable: gamma) Assignment Rule (variable: gamma_fbp1)
Assignment Rule (variable: gamma_gp) Assignment Rule (variable: gamma_gs) Assignment Rule (variable: gamma_pfk2) Assignment Rule (variable: gamma_fbp2)
Assignment Rule (variable: gamma_pk) Assignment Rule (variable: v_ALD) Assignment Rule (variable: v_EN) Assignment Rule (variable: v_fbp1_native)
Assignment Rule (variable: v_fbp1_phospho) Assignment Rule (variable: v_FBP1) Assignment Rule (variable: v_GAPDH) Assignment Rule (variable: vmax_gk)
Assignment Rule (variable: v_GK) Assignment Rule (variable: v_GLUT2) Assignment Rule (variable: v_GLCT_ER) Assignment Rule (variable: vmax_gp)
Assignment Rule (variable: vmax_gp_native) Assignment Rule (variable: km_glc1p_gp_native) Assignment Rule (variable: vmax_gp_phospho) Assignment Rule (variable: v_gp_native)
Assignment Rule (variable: v_gp_phospho) Assignment Rule (variable: v_GP) Assignment Rule (variable: v_G6P_ER) Assignment Rule (variable: v_GPI)
Assignment Rule (variable: v_G1PI) Assignment Rule (variable: v_G6PT_ER) Assignment Rule (variable: vmax_gs) Assignment Rule (variable: km_udpglc_gs_native)
Assignment Rule (variable: km_udpglc_gs_phospho) Assignment Rule (variable: v_gs_native) Assignment Rule (variable: v_gs_phospho) Assignment Rule (variable: v_GS)
Assignment Rule (variable: v_LACT) Assignment Rule (variable: v_LDH) Assignment Rule (variable: v_MALT) Assignment Rule (variable: v_MDH)
Assignment Rule (variable: v_MDH_mito) Assignment Rule (variable: v_NDK_GTP) Assignment Rule (variable: v_NDK_UTP) Assignment Rule (variable: v_NDK_GTP_mito)
Assignment Rule (variable: v_PC) Assignment Rule (variable: v_PEPCK) Assignment Rule (variable: v_PEPCK_mito) Assignment Rule (variable: keq_pept)
Assignment Rule (variable: v_PEPT) Assignment Rule (variable: km_atp_pfk1) Assignment Rule (variable: ki_atp_pfk1) Assignment Rule (variable: km_fru6p_pfk1)
Assignment Rule (variable: n_fru6p_pfk1) Assignment Rule (variable: v_PFK1) Assignment Rule (variable: v_pfk2_native) Assignment Rule (variable: v_pfk2_phospho)
Assignment Rule (variable: v_PFK2) Assignment Rule (variable: v_fbp2_native) Assignment Rule (variable: v_fbp2_phospho) Assignment Rule (variable: v_FBP2)
Assignment Rule (variable: v_PGK) Assignment Rule (variable: v_PGM) Assignment Rule (variable: v_pk_native) Assignment Rule (variable: v_pk_phospho)
Assignment Rule (variable: v_PK) Assignment Rule (variable: v_PyrMalT) Assignment Rule (variable: v_PyrT) Assignment Rule (variable: v_TPI)
Assignment Rule (variable: v_UGT)      
Physical entities
Compartments Species
ext glc_ext lac_ext  
cyto glc glc6p glc1p
udpglc glyglc fru6p
fru16bp fru26bp gap
dhap bpg13 pg3
pg2 pep pyr
oa lac mal
cit h atp
adp amp utp
udp gtp gdp
pp p p_er
co2 nad nadh
     
er glc_er glc6p_er  
mito co2_mito oa_mito pep_mito
pyr_mito mal_mito p_mito
h_mito atp_mito adp_mito
gtp_mito gdp_mito nadh_mito
nad_mito    
Global parameters
Vmm ONE O5 M1
O1 F R T
vmax_ald keq_ald km_fru16bp_ald km_gap_ald
km_dhap_ald vmax_en keq_en km_pg2_en
km_pep_en vmax_fbp1 km_fru16bp_fbp1_native ki_fru26bp_fbp1
n_fbp1 ki_amp_fbp1 n_amp_fbp1 km_fru16bp_fbp1_phospho
vmax_gapdh keq_gapdh km_nad_gapdh km_gap_gapdh
km_p_gapdh km_nadh_gapdh km_bpg13_gapdh v0_gk
n_gk ka_glc_gk f_gk ki_fru6p_gk
n2_gk km_glc_gk km_atp_gk vmax_glct_er
km_glc_glct_er km_glc_er_glct vmax_glut2 km_glc_ext_glut2
km_glc_glut2 v0_gp store keq_gp
km_glyglc_gp_native km_p_gp_native ka_amp_gp_native ka_amp_glc1p_gp
k0_glc1p_gp km_glyglc_gp_phospho km_p_gp_phospho km_glc1p_gp_phospho
ka_amp_gp_phospho vmax_g6p_er km_glc6p_er_g6p_er vmax_gpi
keq_gpi km_glc6p_gpi km_fru6p_gpi vmax_g1pi
keq_g1pi km_glc6p_g1pi km_glc1p_g1pi vmax_g6pt_er
km_glc6p_g6pt_er km_glc6p_er_g6pt_er v0_gs kb_udpglc_gs_native
k0_udpglc_gs_native ka_glc6p_gs_native kb_udpglc_gs_phospho k0_udpglc_gs_phospho
ka_glc6p_gs_phospho vmax_lact km_lac_lact km_lac_ext_lact
vmax_ldh keq_ldh km_lac_ldh km_nad_ldh
km_pyr_ldh km_nadh_ldh km_p_mito_malt km_mal_malt
km_p_malt km_mal_mito_malt vmax_malt km_oa_mdh
km_nadh_mdh km_nad_mdh km_mal_mdh keq_mdh
vmax_mdh km_oa_mito_mdh_mito km_nadh_mito_mdh_mito km_nad_mito_mdh_mito
km_mal_mito_mdh_mito keq_mdh_mito vmax_mdh_mito vmax_ndk_gtp
keq_ndk km_atp_ndk km_gdp_ndk km_adp_ndk
km_gtp_ndk vmax_ndk_utp km_udp_ndk km_utp_ndk
vmax_ndk_gtp_mito km_atp_mito_ndk km_gdp_mito_ndk km_adp_mito_ndk
km_gtp_mito_ndk vmax_PC keq_pc km_atp_mito_pc
km_pyr_mito_pc km_co2_mito_pc vmax_pepck keq_pepck
km_oa_pepck km_gtp_pepck km_pep_pepck km_gdp_pepck
km_co2_pepck vmax_pepck_mito keq_pepck_mito km_oa_mito_pepck_mito
km_gtp_mito_pepck_mito km_pep_mito_pepck_mito km_gdp_mito_pepck_mito km_co2_mito_pepck_mito
vmax_pept km_pep_pept km_pep_mito_pept vmax_pfk1
ni_pfk1 ka_fru26bp_pfk1_km_atp k0_atp_pfk1_km_atp ki0_atp_pfk1
f_fru26bp_pfk1_ki_atp ka_fru26bp_pfk1_ki_atp k0_fru6p_pfk1_km_fru6p ki_atp_pfk1_km_fru6p
ki_cit_pfk1_km_fru6p f_amp_pfk1_km_fru6p n_amp_pfk1_km_fru6p ka_amp_pfk1_km_fru6p
f_p_pfk1_km_fru6p ka_p_pfk1_km_fru6p f_fru26bp_pfk1_km_fru6p n_fru26bp_pfk1_km_fru6p
ka_fru26bp_pfk1_km_fru6p n0_pfk1_n_fru6p n_atp_pfk1_n_fru6p ki_atp_pfk1_n_fru6p
f_amp_pfk1_n_fru6p n_amp_pfk1_n_fru6p ka_amp_pfk1_n_fru6p f_cit_pfk1_n_fru6p
n_cit_pfk1_n_fru6p ki_cit_pfk1_n_fru6p f_p_pfk1_n_fru6p n_p_pfk1_n_fru6p
ka_p_pfk1_n_fru6p f_fru26bp_pfk1_n_fru6p n_fru26bp_pfk1_n_fru6p ka_fru26bp_pfk1_n_fru6p
vmax_pfk2 n_fru6p_pfk2_native km_fru6p_pfk2_native km_atp_pfk2_native
n0_pfk2 ki_pep_pfk2 n_fru6p_pfk2_phospho km_fru6p_pfk2_phospho
km_atp_pfk2_phospho vmax_fbp2 km_fru26bp_fbp2_native ki_fru6p_fbp2_native
f_fbp2_phospho km_fru26bp_fbp2_phospho ki_fru6p_fbp2_phospho vmax_pgk
keq_pgk km_adp_pgk km_bpg13_pgk km_atp_pgk
km_pg3_pgk vmax_pgm keq_pgm km_pg3_pgm
km_pg2_pgm vmax_pk km_pep_pk_native ki_atp_pk_native
ka_fru16bp_pk_native km_adp_pk_native km_pep_pk_phospho ki_atp_pk_phospho
ka_fru16bp_pk_phospho km_adp_pk_phospho n_pk_phospho vmax_pyrmalt
km_mal_mito_pyrmalt km_pyr_pyrmalt km_mal_pyrmalt km_pyr_mito_pyrmalt
vmax_pyrt km_pyr_pyrt km_pyr_mito_pyrt vmax_tpi
keq_tpi km_dhap_tpi km_gap_tpi vmax_ugt
keq_ugt km_utp_ugt km_glc1p_ugt km_udpglc_ugt
km_pp_ugt ins0 n_ins km_glc_ext_ins
glucagon0 n_glucagon km_glc_ext_glucagon glucagonb
n_ins_gamma km_ins_gamma n_glucagon_gamma km_glucagon_gamma
ins glucagon gamma gamma_fbp1
gamma_gp gamma_gs gamma_pfk2 gamma_fbp2
gamma_pk v_ALD v_EN v_fbp1_native
v_fbp1_phospho v_FBP1 v_GAPDH vmax_gk
v_GK v_GLUT2 v_GLCT_ER vmax_gp
vmax_gp_native km_glc1p_gp_native vmax_gp_phospho v_gp_native
v_gp_phospho v_GP v_G6P_ER v_GPI
v_G1PI v_G6PT_ER vmax_gs km_udpglc_gs_native
km_udpglc_gs_phospho v_gs_native v_gs_phospho v_GS
v_LACT v_LDH v_MALT v_MDH
v_MDH_mito v_NDK_GTP v_NDK_UTP v_NDK_GTP_mito
v_PC v_PEPCK v_PEPCK_mito keq_pept
v_PEPT km_atp_pfk1 ki_atp_pfk1 km_fru6p_pfk1
n_fru6p_pfk1 v_PFK1 v_pfk2_native v_pfk2_phospho
v_PFK2 v_fbp2_native v_fbp2_phospho v_FBP2
v_PGK v_PGM v_pk_native v_pk_phospho
v_PK v_PyrMalT v_PyrT v_TPI
v_UGT      
Reactions (35)
 
 ALD [fru16bp] ↔ [gap] + [dhap];  
 
 EN [pg2] ↔ [pep];  
 
 FBP1 [fru16bp] ↔ [fru6p];  
 
 GAPDH [gap] ↔ [bpg13];  
 
 GK [glc] ↔ [glc6p];  
 
 GLCT_ER [glc] ↔ [glc_er];  
 
 GLUT2  ↔ [glc];  
 
 GP [glyglc] ↔ [glc1p];  
 
 G6P_ER [glc6p_er] ↔ [glc_er];  
 
 GPI [glc6p] ↔ [fru6p];  
 
 G1PI [glc1p] ↔ [glc6p];  
 
 G6PT_ER [glc6p] ↔ [glc6p_er];  
 
 GS [udpglc] ↔ [glyglc] + [udp];  
 
 LACT  ↔ [lac];  
 
 LDH [pyr] ↔ [lac];  
 
 MALT 6.0 × [mal_mito] ↔ [mal];  
 
 MDH [mal] ↔ [oa];  
 
 MDH_mito [mal_mito] ↔ [oa_mito];  
 
 NDK_GTP [gdp] ↔ [gtp];  
 
 NDK_UTP [udp] ↔ [utp];  
 
 NDK_GTP_mito [gdp_mito] ↔ [gtp_mito];  
 
 PC [pyr_mito] ↔ [oa_mito];  
 
 PEPCK [oa] + [gtp] ↔ [pep] + [gdp];  
 
 PEPCK_mito [oa_mito] + [gtp_mito] ↔ [pep_mito] + [gdp_mito];  
 
 PEPT 6.0 × [pep_mito] ↔ [pep];  
 
 PFK1 [fru6p] ↔ [fru16bp];  
 
 PFK2 [fru6p] ↔ [fru26bp];  
 
 FBP2 [fru26bp] ↔ [fru6p];  
 
 PGK [bpg13] ↔ [pg3];  
 
 PGM [pg3] ↔ [pg2];  
 
 PK [pep] ↔ [pyr];  
 
 PyrMalT 6.0 × [mal_mito] + [pyr] ↔ 6.0 × [pyr_mito] + [mal];  
 
 PyrT [pyr] ↔ 6.0 × [pyr_mito];  
 
 TPI [dhap] ↔ [gap];  
 
 UGT [utp] + [glc1p] ↔ [udpglc];  
 
Rules (69)
 
 Assignment Rule (name: ins) ins = ins0*glc_ext^n_ins/(glc_ext^n_ins+km_glc_ext_ins^n_ins)
 
 Assignment Rule (name: glucagon) glucagon = glucagon0*(ONE-glc_ext^n_glucagon/(glc_ext^n_glucagon+km_glc_ext_glucagon^n_glucagon))+glucagonb
 
 Assignment Rule (name: gamma) gamma = O5*(ONE+(glucagon^n_glucagon_gamma/(glucagon^n_glucagon_gamma+km_glucagon_gamma^n_glucagon_gamma)-ins^n_ins_gamma/(ins^n_ins_gamma+km_ins_gamma^n_ins_gamma)))
 
 Assignment Rule (name: gamma_fbp1) gamma_fbp1 = gamma
 
 Assignment Rule (name: gamma_gp) gamma_gp = gamma
 
 Assignment Rule (name: gamma_gs) gamma_gs = gamma
 
 Assignment Rule (name: gamma_pfk2) gamma_pfk2 = gamma
 
 Assignment Rule (name: gamma_fbp2) gamma_fbp2 = gamma
 
 Assignment Rule (name: gamma_pk) gamma_pk = gamma
 
 Assignment Rule (name: v_ALD) v_ALD = vmax_ald*(fru16bp-gap*dhap/keq_ald)/((ONE+fru16bp/km_fru16bp_ald+(ONE+gap/km_gap_ald)*(ONE+dhap/km_dhap_ald))-ONE)
 
 Assignment Rule (name: v_EN) v_EN = vmax_en*(pg2-pep/keq_en)/(ONE+pg2/km_pg2_en+pep/km_pep_en)
 
 Assignment Rule (name: v_fbp1_native) v_fbp1_native = fru16bp/(fru16bp+km_fru16bp_fbp1_native)/((ONE+(fru26bp/ki_fru26bp_fbp1)^n_fbp1)*(ONE+(amp/ki_amp_fbp1)^n_amp_fbp1))
 
 Assignment Rule (name: v_fbp1_phospho) v_fbp1_phospho = fru16bp/(fru16bp+km_fru16bp_fbp1_phospho)/((ONE+(fru26bp/ki_fru26bp_fbp1)^n_fbp1)*(ONE+(amp/ki_amp_fbp1)^n_amp_fbp1))
 
 Assignment Rule (name: v_FBP1) v_FBP1 = vmax_fbp1*((ONE-gamma_fbp1)*v_fbp1_native+gamma_fbp1*v_fbp1_phospho)
 
 Assignment Rule (name: v_GAPDH) v_GAPDH = vmax_gapdh*(nad*gap*p-bpg13*nadh/keq_gapdh)/(((ONE+nad/km_nad_gapdh)*(ONE+gap/km_gap_gapdh)*(ONE+p/km_p_gapdh)+(ONE+nadh/km_nadh_gapdh)*(ONE+bpg13/km_bpg13_gapdh))-ONE)
 
 Assignment Rule (name: vmax_gk) vmax_gk = v0_gk*glc^n2_gk/(glc^n2_gk+ka_glc_gk^n2_gk)*(ONE-f_gk*fru6p/(fru6p+ki_fru6p_gk))
 
 Assignment Rule (name: v_GK) v_GK = vmax_gk*atp/(atp+km_atp_gk)*glc^n_gk/(glc^n_gk+km_glc_gk^n_gk)
 
 Assignment Rule (name: v_GLUT2) v_GLUT2 = vmax_glut2*(glc_ext-glc)/(ONE+glc_ext/km_glc_ext_glut2+glc/km_glc_glut2)
 
 Assignment Rule (name: v_GLCT_ER) v_GLCT_ER = vmax_glct_er*(glc-glc_er)/(ONE+glc/km_glc_glct_er+glc_er/km_glc_er_glct)
 
 Assignment Rule (name: vmax_gp) vmax_gp = v0_gp*glyglc/store
 
 Assignment Rule (name: vmax_gp_native) vmax_gp_native = amp/(amp+ka_amp_gp_native)/(km_glyglc_gp_native*km_p_gp_native)
 
 Assignment Rule (name: km_glc1p_gp_native) km_glc1p_gp_native = k0_glc1p_gp*(ONE-amp/(amp+ka_amp_glc1p_gp))
 
 Assignment Rule (name: vmax_gp_phospho) vmax_gp_phospho = amp/(amp+ka_amp_gp_phospho)/(km_glyglc_gp_phospho*km_p_gp_phospho)
 
 Assignment Rule (name: v_gp_native) v_gp_native = vmax_gp_native*(glyglc*p-glc1p/keq_gp)/(((ONE+glyglc/km_glyglc_gp_native)*(ONE+p/km_p_gp_native)+ONE+glc1p/km_glc1p_gp_native)-ONE)
 
 Assignment Rule (name: v_gp_phospho) v_gp_phospho = vmax_gp_phospho*(glyglc*p-glc1p/keq_gp)/(((ONE+glyglc/km_glyglc_gp_phospho)*(ONE+p/km_p_gp_phospho)+ONE+glc1p/km_glc1p_gp_phospho)-ONE)
 
 Assignment Rule (name: v_GP) v_GP = vmax_gp*((ONE-gamma_gp)*v_gp_native+gamma_gp*v_gp_phospho)
 
 Assignment Rule (name: v_G6P_ER) v_G6P_ER = vmax_g6p_er*glc6p_er/(glc6p_er+km_glc6p_er_g6p_er)
 
 Assignment Rule (name: v_GPI) v_GPI = vmax_gpi*(glc6p-fru6p/keq_gpi)/(ONE+glc6p/km_glc6p_gpi+fru6p/km_fru6p_gpi)
 
 Assignment Rule (name: v_G1PI) v_G1PI = vmax_g1pi*(glc1p-glc6p/keq_g1pi)/(ONE+glc6p/km_glc6p_g1pi+glc1p/km_glc1p_g1pi)
 
 Assignment Rule (name: v_G6PT_ER) v_G6PT_ER = vmax_g6pt_er*(glc6p-glc6p_er)/(ONE+glc6p/km_glc6p_g6pt_er+glc6p_er/km_glc6p_er_g6pt_er)
 
 Assignment Rule (name: vmax_gs) vmax_gs = v0_gs*(store-glyglc)/((store-glyglc)+store*O1)
 
 Assignment Rule (name: km_udpglc_gs_native) km_udpglc_gs_native = kb_udpglc_gs_native+k0_udpglc_gs_native*(ONE-glc6p/(glc6p+ka_glc6p_gs_native))
 
 Assignment Rule (name: km_udpglc_gs_phospho) km_udpglc_gs_phospho = kb_udpglc_gs_phospho+k0_udpglc_gs_phospho*(ONE-glc6p/(glc6p+ka_glc6p_gs_phospho))
 
 Assignment Rule (name: v_gs_native) v_gs_native = udpglc/(udpglc+km_udpglc_gs_native)
 
 Assignment Rule (name: v_gs_phospho) v_gs_phospho = udpglc/(udpglc+km_udpglc_gs_phospho)
 
 Assignment Rule (name: v_GS) v_GS = vmax_gs*((ONE-gamma_gs)*v_gs_native+gamma_gs*v_gs_phospho)
 
 Assignment Rule (name: v_LACT) v_LACT = vmax_lact*(lac_ext-lac)/(ONE+lac/km_lac_lact+lac_ext/km_lac_ext_lact)
 
 Assignment Rule (name: v_LDH) v_LDH = vmax_ldh*(pyr*nadh-lac*nad/keq_ldh)/(((ONE+nadh/km_nadh_ldh)*(ONE+pyr/km_pyr_ldh)+(ONE+nad/km_nad_ldh)*(ONE+lac/km_lac_ldh))-ONE)
 
 Assignment Rule (name: v_MALT) v_MALT = vmax_malt*(mal_mito*p-mal*p_mito)/(((ONE+mal_mito/km_mal_mito_malt)*(ONE+p/km_p_malt)+(ONE+mal/km_mal_malt)*(ONE+p_mito/km_p_mito_malt))-ONE)
 
 Assignment Rule (name: v_MDH) v_MDH = vmax_mdh*(mal*nad-oa*nadh/keq_mdh)/(((ONE+mal/km_mal_mdh)*(ONE+nad/km_nad_mdh)+(ONE+nadh/km_nadh_mdh)*(ONE+oa/km_oa_mdh))-ONE)
 
 Assignment Rule (name: v_MDH_mito) v_MDH_mito = vmax_mdh_mito*(mal_mito*nad_mito-oa_mito*nadh_mito/keq_mdh_mito)/(((ONE+mal_mito/km_mal_mito_mdh_mito)*(ONE+nad_mito/km_nad_mito_mdh_mito)+(ONE+nadh_mito/km_nadh_mito_mdh_mito)*(ONE+oa_mito/km_oa_mito_mdh_mito))-ONE)
 
 Assignment Rule (name: v_NDK_GTP) v_NDK_GTP = vmax_ndk_gtp*(atp*gdp-gtp*adp/keq_ndk)/(((ONE+atp/km_atp_ndk)*(ONE+gdp/km_gdp_ndk)+(ONE+adp/km_adp_ndk)*(ONE+gtp/km_gtp_ndk))-ONE)
 
 Assignment Rule (name: v_NDK_UTP) v_NDK_UTP = vmax_ndk_utp*(atp*udp-utp*adp/keq_ndk)/(((ONE+atp/km_atp_ndk)*(ONE+udp/km_udp_ndk)+(ONE+adp/km_adp_ndk)*(ONE+utp/km_utp_ndk))-ONE)
 
 Assignment Rule (name: v_NDK_GTP_mito) v_NDK_GTP_mito = vmax_ndk_gtp_mito*(atp_mito*gdp_mito-gtp_mito*adp_mito/keq_ndk)/(((ONE+atp_mito/km_atp_mito_ndk)*(ONE+gdp_mito/km_gdp_mito_ndk)+(ONE+adp_mito/km_adp_mito_ndk)*(ONE+gtp_mito/km_gtp_mito_ndk))-ONE)
 
 Assignment Rule (name: v_PC) v_PC = vmax_PC*(atp_mito*pyr_mito*co2_mito-oa_mito*adp_mito*p_mito/keq_pc)/((atp_mito+km_atp_mito_pc)*(pyr_mito+km_pyr_mito_pc)*(co2_mito+km_co2_mito_pc))
 
 Assignment Rule (name: v_PEPCK) v_PEPCK = vmax_pepck*(oa*gtp-pep*gdp*co2/keq_pepck)/(((ONE+oa/km_oa_pepck)*(ONE+gtp/km_gtp_pepck)+(ONE+pep/km_pep_pepck)*(ONE+gdp/km_gdp_pepck)*(ONE+co2/km_co2_pepck))-ONE)
 
 Assignment Rule (name: v_PEPCK_mito) v_PEPCK_mito = vmax_pepck_mito*(oa_mito*gtp_mito-pep_mito*gdp_mito*co2_mito/keq_pepck_mito)/(((ONE+oa_mito/km_oa_mito_pepck_mito)*(ONE+gtp_mito/km_gtp_mito_pepck_mito)+(ONE+pep_mito/km_pep_mito_pepck_mito)*(ONE+gdp_mito/km_gdp_mito_pepck_mito)*(ONE+co2_mito/km_co2_mito_pepck_mito))-ONE)
 
 Assignment Rule (name: keq_pept) keq_pept = exp(M1*Vmm*F/(R*T))
 
 Assignment Rule (name: v_PEPT) v_PEPT = vmax_pept*(pep_mito-pep/keq_pept)/(ONE+pep/km_pep_pept+pep_mito/km_pep_mito_pept)
 
 Assignment Rule (name: km_atp_pfk1) km_atp_pfk1 = k0_atp_pfk1_km_atp*fru26bp/(fru26bp+ka_fru26bp_pfk1_km_atp)
 
 Assignment Rule (name: ki_atp_pfk1) ki_atp_pfk1 = ki0_atp_pfk1*(ONE+f_fru26bp_pfk1_ki_atp*fru26bp/(fru26bp+ka_fru26bp_pfk1_ki_atp))
 
 Assignment Rule (name: km_fru6p_pfk1) km_fru6p_pfk1 = k0_fru6p_pfk1_km_fru6p*(ONE+atp/ki_atp_pfk1_km_fru6p)*(ONE+cit/ki_cit_pfk1_km_fru6p)*(ONE-f_amp_pfk1_km_fru6p*amp^n_amp_pfk1_km_fru6p/(amp^n_amp_pfk1_km_fru6p+ka_amp_pfk1_km_fru6p^n_amp_pfk1_km_fru6p))*(ONE-f_p_pfk1_km_fru6p*p/(p+ka_p_pfk1_km_fru6p))*(ONE-f_fru26bp_pfk1_km_fru6p*fru26bp^n_fru26bp_pfk1_km_fru6p/(fru26bp^n_fru26bp_pfk1_km_fru6p+ka_fru26bp_pfk1_km_fru6p^n_fru26bp_pfk1_km_fru6p))
 
 Assignment Rule (name: n_fru6p_pfk1) n_fru6p_pfk1 = (n0_pfk1_n_fru6p+atp^n_atp_pfk1_n_fru6p/(atp^n_atp_pfk1_n_fru6p+ki_atp_pfk1_n_fru6p^n_atp_pfk1_n_fru6p))*(ONE-f_amp_pfk1_n_fru6p*amp^n_amp_pfk1_n_fru6p/(amp^n_amp_pfk1_n_fru6p+ka_amp_pfk1_n_fru6p^n_amp_pfk1_n_fru6p))*(ONE+f_cit_pfk1_n_fru6p*cit^n_cit_pfk1_n_fru6p/(cit^n_cit_pfk1_n_fru6p+ki_cit_pfk1_n_fru6p^n_cit_pfk1_n_fru6p))*(ONE-f_p_pfk1_n_fru6p*p^n_p_pfk1_n_fru6p/(p^n_p_pfk1_n_fru6p+ka_p_pfk1_n_fru6p^n_p_pfk1_n_fru6p))*(ONE-f_fru26bp_pfk1_n_fru6p*fru26bp^n_fru26bp_pfk1_n_fru6p/(fru26bp^n_fru26bp_pfk1_n_fru6p+ka_fru26bp_pfk1_n_fru6p^n_fru26bp_pfk1_n_fru6p))
 
 Assignment Rule (name: v_PFK1) v_PFK1 = vmax_pfk1*atp/(atp+km_atp_pfk1)*(ONE-atp^ni_pfk1/(atp^ni_pfk1+ki_atp_pfk1^ni_pfk1))*fru6p^n_fru6p_pfk1/(fru6p^n_fru6p_pfk1+km_fru6p_pfk1^n_fru6p_pfk1)
 
 Assignment Rule (name: v_pfk2_native) v_pfk2_native = fru6p^n_fru6p_pfk2_native/(fru6p^n_fru6p_pfk2_native+km_fru6p_pfk2_native^n_fru6p_pfk2_native)*atp/(atp+km_atp_pfk2_native)*(ONE-n0_pfk2*pep/(pep+ki_pep_pfk2))
 
 Assignment Rule (name: v_pfk2_phospho) v_pfk2_phospho = fru6p^n_fru6p_pfk2_phospho/(fru6p^n_fru6p_pfk2_phospho+km_fru6p_pfk2_phospho^n_fru6p_pfk2_phospho)*atp/(atp+km_atp_pfk2_phospho)*(ONE-n0_pfk2*pep/(pep+ki_pep_pfk2))
 
 Assignment Rule (name: v_PFK2) v_PFK2 = vmax_pfk2*((ONE-gamma_pfk2)*v_pfk2_native+gamma_pfk2*v_pfk2_phospho)
 
 Assignment Rule (name: v_fbp2_native) v_fbp2_native = fru26bp/(fru26bp+km_fru26bp_fbp2_native)/(ONE+fru6p/ki_fru6p_fbp2_native)
 
 Assignment Rule (name: v_fbp2_phospho) v_fbp2_phospho = f_fbp2_phospho*fru26bp/(fru26bp+km_fru26bp_fbp2_phospho)/(ONE+fru6p/ki_fru6p_fbp2_phospho)
 
 Assignment Rule (name: v_FBP2) v_FBP2 = vmax_fbp2*((ONE-gamma_fbp2)*v_fbp2_native+gamma_fbp2*v_fbp2_phospho)
 
 Assignment Rule (name: v_PGK) v_PGK = vmax_pgk*(adp*bpg13-atp*pg3/keq_pgk)/(((ONE+adp/km_adp_pgk)*(ONE+bpg13/km_bpg13_pgk)+(ONE+atp/km_atp_pgk)*(ONE+pg3/km_pg3_pgk))-ONE)
 
 Assignment Rule (name: v_PGM) v_PGM = vmax_pgm*(pg3-pg2/keq_pgm)/(ONE+pg3/km_pg3_pgm+pg2/km_pg2_pgm)
 
 Assignment Rule (name: v_pk_native) v_pk_native = pep/(pep+km_pep_pk_native*(ONE+atp/ki_atp_pk_native)*(ONE-fru16bp/(fru16bp+ka_fru16bp_pk_native)))*adp/(adp+km_adp_pk_native)
 
 Assignment Rule (name: v_pk_phospho) v_pk_phospho = pep^n_pk_phospho/(pep^n_pk_phospho+(km_pep_pk_phospho*(ONE+atp/ki_atp_pk_phospho)*(ONE-fru16bp/(fru16bp+ka_fru16bp_pk_phospho)))^n_pk_phospho)*adp/(adp+km_adp_pk_phospho)
 
 Assignment Rule (name: v_PK) v_PK = vmax_pk*((ONE-gamma_pk)*v_pk_native+gamma_pk*v_pk_phospho)
 
 Assignment Rule (name: v_PyrMalT) v_PyrMalT = vmax_pyrmalt*(mal_mito*pyr-mal*pyr_mito)/(((ONE+mal_mito/km_mal_mito_pyrmalt)*(ONE+pyr/km_pyr_pyrmalt)+(ONE+mal/km_mal_pyrmalt)*(ONE+pyr_mito/km_pyr_mito_pyrmalt))-ONE)
 
 Assignment Rule (name: v_PyrT) v_PyrT = vmax_pyrt*(pyr*h-pyr_mito*h_mito)/(ONE+pyr/km_pyr_pyrt+pyr_mito/km_pyr_mito_pyrt)
 
 Assignment Rule (name: v_TPI) v_TPI = vmax_tpi*(dhap-gap/keq_tpi)/(ONE+dhap/km_dhap_tpi+gap/km_gap_tpi)
 
 Assignment Rule (name: v_UGT) v_UGT = vmax_ugt*(utp*glc1p-udpglc*pp/keq_ugt)/(((ONE+utp/km_utp_ugt)*(ONE+glc1p/km_glc1p_ugt)+(ONE+udpglc/km_udpglc_ugt)*(ONE+pp/km_pp_ugt))-ONE)
 
  Spatial dimensions: 3.0  Compartment size: 1.0
 
 glc_ext
Compartment: ext
Initial concentration: 4.0
 
 lac_ext
Compartment: ext
Initial concentration: 1.0
 
  Spatial dimensions: 3.0  Compartment size: 1.0
 
 glc
Compartment: cyto
Initial concentration: 4.92256221964406
 
 glc6p
Compartment: cyto
Initial concentration: 0.189929897759633
 
 glc1p
Compartment: cyto
Initial concentration: 0.0117240677629403
 
 udpglc
Compartment: cyto
Initial concentration: 0.34072421289254
 
 glyglc
Compartment: cyto
Initial concentration: 17.6332590346802
 
 fru6p
Compartment: cyto
Initial concentration: 0.0569790458002266
 
 fru16bp
Compartment: cyto
Initial concentration: 8.08171827419162E-4
 
 fru26bp
Compartment: cyto
Initial concentration: 5.40214088345945E-4
 
 gap
Compartment: cyto
Initial concentration: 0.00190705466540883
 
 dhap
Compartment: cyto
Initial concentration: 0.0419590925497022
 
 bpg13
Compartment: cyto
Initial concentration: 0.00108805669971103
 
 pg3
Compartment: cyto
Initial concentration: 0.306393473858216
 
 pg2
Compartment: cyto
Initial concentration: 0.029376172923967
 
 pep
Compartment: cyto
Initial concentration: 0.0499395072312233
 
 pyr
Compartment: cyto
Initial concentration: 0.0852149179429942
 
 oa
Compartment: cyto
Initial concentration: 0.0243963256092358
 
 lac
Compartment: cyto
Initial concentration: 0.678771756200856
 
 mal
Compartment: cyto
Initial concentration: 0.719656614702017
 
 cit
Compartment: cyto
Initial concentration: 0.75
 
 h
Compartment: cyto
Initial concentration: 1.0E-4
 
 atp
Compartment: cyto
Initial concentration: 3.25
 
 adp
Compartment: cyto
Initial concentration: 0.5
 
 amp
Compartment: cyto
Initial concentration: 0.16
 
 utp
Compartment: cyto
Initial concentration: 0.744701216226319
 
 udp
Compartment: cyto
Initial concentration: 0.114570584540414
 
 gtp
Compartment: cyto
Initial concentration: 0.693339386314472
 
 gdp
Compartment: cyto
Initial concentration: 0.106667613680444
 
 pp
Compartment: cyto
Initial concentration: 0.008
 
 p
Compartment: cyto
Initial concentration: 5.0
 
 p_er
Compartment: cyto
Initial concentration: 5.0
 
 co2
Compartment: cyto
Initial concentration: 5.0
 
 nad
Compartment: cyto
Initial concentration: 1.13
 
 nadh
Compartment: cyto
Initial concentration: 0.001
 
  Spatial dimensions: 3.0  Compartment size: 1.0
 
 glc_er
Compartment: er
Initial concentration: 4.92256224638738
 
 glc6p_er
Compartment: er
Initial concentration: 0.189929893608775
 
  Spatial dimensions: 3.0  Compartment size: 1.0
 
 co2_mito
Compartment: mito
Initial concentration: 5.0
 
 oa_mito
Compartment: mito
Initial concentration: 4.64731194122211E-4
 
 pep_mito
Compartment: mito
Initial concentration: 1.11191930877581E-4
 
 pyr_mito
Compartment: mito
Initial concentration: 0.132228238255726
 
 mal_mito
Compartment: mito
Initial concentration: 1.30359361718805
 
 p_mito
Compartment: mito
Initial concentration: 8.0
 
 h_mito
Compartment: mito
Initial concentration: 6.0E-5
 
 atp_mito
Compartment: mito
Initial concentration: 17.5
 
 adp_mito
Compartment: mito
Initial concentration: 7.5
 
 gtp_mito
Compartment: mito
Initial concentration: 0.559995709478379
 
 gdp_mito
Compartment: mito
Initial concentration: 0.240004290518223
 
 nadh_mito
Compartment: mito
Initial concentration: 0.004
 
 nad_mito
Compartment: mito
Initial concentration: 0.046
 
Global Parameters (305)
 
   Vmm
Value: -0.155   (Units: V)
Constant
 
   ONE
Value: 1.0   (Units: dimensionless)
Constant
 
   O5
Value: 0.5   (Units: dimensionless)
Constant
 
   M1
Value: -1.0   (Units: dimensionless)
Constant
 
   O1
Value: 0.1   (Units: dimensionless)
Constant
 
   F
Value: 96490.0   (Units: J_per_V_per_mol)
Constant
 
   R
Value: 8.3   (Units: J_per_K_per_mol)
Constant
 
   T
Value: 293.0   (Units: K)
Constant
 
   vmax_ald
Value: 7.78E8   (Units: per_h)
Constant
 
   keq_ald
Value: 0.099   (Units: mM)
Constant
 
   km_fru16bp_ald
Value: 0.004   (Units: mM)
Constant
 
   km_gap_ald
Value: 0.48   (Units: mM)
Constant
 
   km_dhap_ald
Value: 0.38   (Units: mM)
Constant
 
   vmax_en
Value: 1.94E10   (Units: per_h)
Constant
 
   keq_en
Value: 1.7   (Units: dimensionless)
Constant
 
   km_pg2_en
Value: 0.14   (Units: mM)
Constant
 
   km_pep_en
Value: 0.31   (Units: mM)
Constant
 
   vmax_fbp1
Value: 29200.0   (Units: mM_per_h)
Constant
 
   km_fru16bp_fbp1_native
Value: 0.0029   (Units: mM)
Constant
 
   ki_fru26bp_fbp1
Value: 0.00113   (Units: mM)
Constant
 
   n_fbp1
Value: 1.26   (Units: dimensionless)
Constant
 
   ki_amp_fbp1
Value: 0.023   (Units: mM)
Constant
 
   n_amp_fbp1
Value: 2.43   (Units: dimensionless)
Constant
 
   km_fru16bp_fbp1_phospho
Value: 0.0019   (Units: mM)
Constant
 
   vmax_gapdh
Value: 2.92E8   (Units: per_h_per_mMmM)
Constant
 
   keq_gapdh
Value: 1.0E-4   (Units: per_mM)
Constant
 
   km_nad_gapdh
Value: 0.01   (Units: mM)
Constant
 
   km_gap_gapdh
Value: 0.035
Constant
 
   km_p_gapdh
Value: 3.8   (Units: mM)
Constant
 
   km_nadh_gapdh
Value: 0.006   (Units: mM)
Constant
 
   km_bpg13_gapdh
Value: 0.01   (Units: mM)
Constant
 
   v0_gk
Value: 10500.0   (Units: mM_per_h)
Constant
 
   n_gk
Value: 1.5   (Units: dimensionless)
Constant
 
   ka_glc_gk
Value: 15.9   (Units: mM)
Constant
 
   f_gk
Value: 0.75   (Units: dimensionless)
Constant
 
   ki_fru6p_gk
Value: 0.005   (Units: mM)
Constant
 
   n2_gk
Value: 3.7   (Units: dimensionless)
Constant
 
   km_glc_gk
Value: 9.0   (Units: mM)
Constant
 
   km_atp_gk
Value: 0.55   (Units: mM)
Constant
 
   vmax_glct_er
Value: 1.94E10   (Units: per_h)
Constant
 
   km_glc_glct_er
Value: 1.37   (Units: mM)
Constant
 
   km_glc_er_glct
Value: 1.22   (Units: mM)
Constant
 
   vmax_glut2
Value: 90.9   (Units: per_h)
Constant
 
   km_glc_ext_glut2
Value: 17.3   (Units: mM)
Constant
 
   km_glc_glut2
Value: 17.3   (Units: mM)
Constant
 
   v0_gp
Value: 129.0   (Units: mM_per_h)
Constant
 
   store
Value: 300.0   (Units: mM)
Constant
 
   keq_gp
Value: 0.21   (Units: per_mM)
Constant
 
   km_glyglc_gp_native
Value: 2.5   (Units: mM)
Constant
 
   km_p_gp_native
Value: 500.0   (Units: mM)
Constant
 
   ka_amp_gp_native
Value: 0.36   (Units: mM)
Constant
 
   ka_amp_glc1p_gp
Value: 0.5   (Units: mM)
Constant
 
   k0_glc1p_gp
Value: 250.0   (Units: mM)
Constant
 
   km_glyglc_gp_phospho
Value: 1.8   (Units: mM)
Constant
 
   km_p_gp_phospho
Value: 2.1   (Units: mM)
Constant
 
   km_glc1p_gp_phospho
Value: 0.7   (Units: mM)
Constant
 
   ka_amp_gp_phospho
Value: 0.017   (Units: mM)
Constant
 
   vmax_g6p_er
Value: 457.0   (Units: mM_per_h)
Constant
 
   km_glc6p_er_g6p_er
Value: 1.84   (Units: mM)
Constant
 
   vmax_gpi
Value: 1.07E9   (Units: per_h)
Constant
 
   keq_gpi
Value: 0.3   (Units: dimensionless)
Constant
 
   km_glc6p_gpi
Value: 0.055   (Units: mM)
Constant
 
   km_fru6p_gpi
Value: 0.12   (Units: mM)
Constant
 
   vmax_g1pi
Value: 7.78E7   (Units: per_h)
Constant
 
   keq_g1pi
Value: 16.2   (Units: dimensionless)
Constant
 
   km_glc6p_g1pi
Value: 0.045   (Units: mM)
Constant
 
   km_glc1p_g1pi
Value: 0.67   (Units: mM)
Constant
 
   vmax_g6pt_er
Value: 1.94E10   (Units: per_h)
Constant
 
   km_glc6p_g6pt_er
Value: 1.12   (Units: mM)
Constant
 
   km_glc6p_er_g6pt_er
Value: 1.12   (Units: mM)
Constant
 
   v0_gs
Value: 116.0   (Units: mM_per_h)
Constant
 
   kb_udpglc_gs_native
Value: 0.2   (Units: mM)
Constant
 
   k0_udpglc_gs_native
Value: 1.4   (Units: mM)
Constant
 
   ka_glc6p_gs_native
Value: 0.007   (Units: mM)
Constant
 
   kb_udpglc_gs_phospho
Value: 0.3   (Units: mM)
Constant
 
   k0_udpglc_gs_phospho
Value: 32.0   (Units: mM)
Constant
 
   ka_glc6p_gs_phospho
Value: 0.09   (Units: mM)
Constant
 
   vmax_lact
Value: 583.0   (Units: per_h)
Constant
 
   km_lac_lact
Value: 2.42   (Units: mM)
Constant
 
   km_lac_ext_lact
Value: 2.42   (Units: mM)
Constant
 
   vmax_ldh
Value: 1.56E11   (Units: per_h_per_mM)
Constant
 
   keq_ldh
Value: 9000.0   (Units: dimensionless)
Constant
 
   km_lac_ldh
Value: 36.0   (Units: mM)
Constant
 
   km_nad_ldh
Value: 0.11   (Units: mM)
Constant
 
   km_pyr_ldh
Value: 0.15   (Units: mM)
Constant
 
   km_nadh_ldh
Value: 0.015   (Units: mM)
Constant
 
   km_p_mito_malt
Value: 1.41   (Units: mM)
Constant
 
   km_mal_malt
Value: 0.49   (Units: mM)
Constant
 
   km_p_malt
Value: 1.41   (Units: mM)
Constant
 
   km_mal_mito_malt
Value: 0.49   (Units: mM)
Constant
 
   vmax_malt
Value: 1940.0   (Units: per_h_per_mM)
Constant
 
   km_oa_mdh
Value: 0.088   (Units: mM)
Constant
 
   km_nadh_mdh
Value: 0.026   (Units: mM)
Constant
 
   km_nad_mdh
Value: 0.114   (Units: mM)
Constant
 
   km_mal_mdh
Value: 1.1   (Units: mM)
Constant
 
   keq_mdh
Value: 3.0E-5   (Units: dimensionless)
Constant
 
   vmax_mdh
Value: 1.94E9   (Units: per_h_per_mM)
Constant
 
   km_oa_mito_mdh_mito
Value: 0.017   (Units: mM)
Constant
 
   km_nadh_mito_mdh_mito
Value: 0.044   (Units: mM)
Constant
 
   km_nad_mito_mdh_mito
Value: 0.06   (Units: mM)
Constant
 
   km_mal_mito_mdh_mito
Value: 0.33   (Units: mM)
Constant
 
   keq_mdh_mito
Value: 3.1E-5   (Units: dimensionless)
Constant
 
   vmax_mdh_mito
Value: 6.97E11   (Units: per_h_per_mM)
Constant
 
   vmax_ndk_gtp
Value: 1.94E11   (Units: per_h_per_mM)
Constant
 
   keq_ndk
Value: 1.0   (Units: dimensionless)
Constant
 
   km_atp_ndk
Value: 1.33   (Units: mM)
Constant
 
   km_gdp_ndk
Value: 0.031   (Units: mM)
Constant
 
   km_adp_ndk
Value: 0.042   (Units: mM)
Constant
 
   km_gtp_ndk
Value: 0.15   (Units: mM)
Constant
 
   vmax_ndk_utp
Value: 1.94E7   (Units: per_h_per_mM)
Constant
 
   km_udp_ndk
Value: 0.19   (Units: mM)
Constant
 
   km_utp_ndk
Value: 16.0   (Units: mM)
Constant
 
   vmax_ndk_gtp_mito
Value: 1.94E7   (Units: per_h_per_mM)
Constant
 
   km_atp_mito_ndk
Value: 1.66   (Units: mM)
Constant
 
   km_gdp_mito_ndk
Value: 0.036   (Units: mM)
Constant
 
   km_adp_mito_ndk
Value: 0.073   (Units: mM)
Constant
 
   km_gtp_mito_ndk
Value: 0.15   (Units: mM)
Constant
 
   vmax_PC
Value: 3590.0   (Units: mM_per_h)
Constant
 
   keq_pc
Value: 6.55   (Units: dimensionless)
Constant
 
   km_atp_mito_pc
Value: 0.14   (Units: mM)
Constant
 
   km_pyr_mito_pc
Value: 0.33   (Units: mM)
Constant
 
   km_co2_mito_pc
Value: 4.2   (Units: mM)
Constant
 
   vmax_pepck
Value: 1000000.0   (Units: per_h_per_mM)
Constant
 
   keq_pepck
Value: 110.0   (Units: mM)
Constant
 
   km_oa_pepck
Value: 0.024   (Units: mM)
Constant
 
   km_gtp_pepck
Value: 0.021   (Units: mM)
Constant
 
   km_pep_pepck
Value: 0.4   (Units: mM)
Constant
 
   km_gdp_pepck
Value: 0.02   (Units: mM)
Constant
 
   km_co2_pepck
Value: 1.194   (Units: mM)
Constant
 
   vmax_pepck_mito
Value: 1410000.0   (Units: per_h_per_mM)
Constant
 
   keq_pepck_mito
Value: 160.0   (Units: mM)
Constant
 
   km_oa_mito_pepck_mito
Value: 0.0085   (Units: mM)
Constant
 
   km_gtp_mito_pepck_mito
Value: 0.022   (Units: mM)
Constant
 
   km_pep_mito_pepck_mito
Value: 0.4   (Units: mM)
Constant
 
   km_gdp_mito_pepck_mito
Value: 0.02   (Units: mM)
Constant
 
   km_co2_mito_pepck_mito
Value: 1.06   (Units: mM)
Constant
 
   vmax_pept
Value: 194000.0   (Units: per_h)
Constant
 
   km_pep_pept
Value: 0.1   (Units: mM)
Constant
 
   km_pep_mito_pept
Value: 0.1   (Units: mM)
Constant
 
   vmax_pfk1
Value: 76800.0   (Units: mM_per_h)
Constant
 
   ni_pfk1
Value: 4.0   (Units: dimensionless)
Constant
 
   ka_fru26bp_pfk1_km_atp
Value: 0.0027   (Units: mM)
Constant
 
   k0_atp_pfk1_km_atp
Value: 0.2   (Units: mM)
Constant
 
   ki0_atp_pfk1
Value: 0.7   (Units: mM)
Constant
 
   f_fru26bp_pfk1_ki_atp
Value: 9.0   (Units: dimensionless)
Constant
 
   ka_fru26bp_pfk1_ki_atp
Value: 0.54   (Units: mM)
Constant
 
   k0_fru6p_pfk1_km_fru6p
Value: 1.14   (Units: mM)
Constant
 
   ki_atp_pfk1_km_fru6p
Value: 0.6   (Units: mM)
Constant
 
   ki_cit_pfk1_km_fru6p
Value: 3.27   (Units: mM)
Constant
 
   f_amp_pfk1_km_fru6p
Value: 0.77   (Units: dimensionless)
Constant
 
   n_amp_pfk1_km_fru6p
Value: 1.84   (Units: dimensionless)
Constant
 
   ka_amp_pfk1_km_fru6p
Value: 0.1   (Units: mM)
Constant
 
   f_p_pfk1_km_fru6p
Value: 0.85   (Units: dimensionless)
Constant
 
   ka_p_pfk1_km_fru6p
Value: 0.69   (Units: mM)
Constant
 
   f_fru26bp_pfk1_km_fru6p
Value: 0.92   (Units: dimensionless)
Constant
 
   n_fru26bp_pfk1_km_fru6p
Value: 1.2   (Units: dimensionless)
Constant
 
   ka_fru26bp_pfk1_km_fru6p
Value: 0.0045   (Units: mM)
Constant
 
   n0_pfk1_n_fru6p
Value: 3.67   (Units: dimensionless)
Constant
 
   n_atp_pfk1_n_fru6p
Value: 1.59   (Units: dimensionless)
Constant
 
   ki_atp_pfk1_n_fru6p
Value: 0.13   (Units: mM)
Constant
 
   f_amp_pfk1_n_fru6p
Value: 0.4   (Units: dimensionless)
Constant
 
   n_amp_pfk1_n_fru6p
Value: 2.22   (Units: dimensionless)
Constant
 
   ka_amp_pfk1_n_fru6p
Value: 0.086   (Units: mM)
Constant
 
   f_cit_pfk1_n_fru6p
Value: 0.1   (Units: dimensionless)
Constant
 
   n_cit_pfk1_n_fru6p
Value: 4.0   (Units: dimensionless)
Constant
 
   ki_cit_pfk1_n_fru6p
Value: 0.18   (Units: mM)
Constant
 
   f_p_pfk1_n_fru6p
Value: 0.28   (Units: dimensionless)
Constant
 
   n_p_pfk1_n_fru6p
Value: 4.0   (Units: dimensionless)
Constant
 
   ka_p_pfk1_n_fru6p
Value: 0.53   (Units: mM)
Constant
 
   f_fru26bp_pfk1_n_fru6p
Value: 0.37   (Units: dimensionless)
Constant
 
   n_fru26bp_pfk1_n_fru6p
Value: 4.0   (Units: dimensionless)
Constant
 
   ka_fru26bp_pfk1_n_fru6p
Value: 0.0021   (Units: mM)
Constant
 
   vmax_pfk2
Value: 151.0   (Units: mM_per_h)
Constant
 
   n_fru6p_pfk2_native
Value: 1.3   (Units: dimensionless)
Constant
 
   km_fru6p_pfk2_native
Value: 0.015   (Units: mM)
Constant
 
   km_atp_pfk2_native
Value: 0.25   (Units: mM)
Constant
 
   n0_pfk2
Value: 0.85   (Units: dimensionless)
Constant
 
   ki_pep_pfk2
Value: 0.25   (Units: mM)
Constant
 
   n_fru6p_pfk2_phospho
Value: 2.0   (Units: dimensionless)
Constant
 
   km_fru6p_pfk2_phospho
Value: 0.05   (Units: mM)
Constant
 
   km_atp_pfk2_phospho
Value: 0.5   (Units: mM)
Constant
 
   vmax_fbp2
Value: 549.0   (Units: mM_per_h)
Constant
 
   km_fru26bp_fbp2_native
Value: 0.01   (Units: mM)
Constant
 
   ki_fru6p_fbp2_native
Value: 0.0035   (Units: mM)
Constant
 
   f_fbp2_phospho
Value: 1.0   (Units: dimensionless)
Constant
 
   km_fru26bp_fbp2_phospho
Value: 5.0E-4   (Units: mM)
Constant
 
   ki_fru6p_fbp2_phospho
Value: 0.01   (Units: mM)
Constant
 
   vmax_pgk
Value: 1.94E10   (Units: per_h_per_mM)
Constant
 
   keq_pgk
Value: 1830.0   (Units: dimensionless)
Constant
 
   km_adp_pgk
Value: 0.35   (Units: mM)
Constant
 
   km_bpg13_pgk
Value: 0.0022   (Units: mM)
Constant
 
   km_atp_pgk
Value: 0.24   (Units: mM)
Constant
 
   km_pg3_pgk
Value: 1.65   (Units: mM)
Constant
 
   vmax_pgm
Value: 1.94E10   (Units: per_h)
Constant
 
   keq_pgm
Value: 0.096   (Units: dimensionless)
Constant
 
   km_pg3_pgm
Value: 0.52   (Units: mM)
Constant
 
   km_pg2_pgm
Value: 0.24   (Units: mM)
Constant
 
   vmax_pk
Value: 12800.0   (Units: mM_per_h)
Constant
 
   km_pep_pk_native
Value: 0.13   (Units: mM)
Constant
 
   ki_atp_pk_native
Value: 1.0   (Units: mM)
Constant
 
   ka_fru16bp_pk_native
Value: 0.0078   (Units: mM)
Constant
 
   km_adp_pk_native
Value: 0.25   (Units: mM)
Constant
 
   km_pep_pk_phospho
Value: 5.8   (Units: mM)
Constant
 
   ki_atp_pk_phospho
Value: 0.32   (Units: mM)
Constant
 
   ka_fru16bp_pk_phospho
Value: 0.0095   (Units: mM)
Constant
 
   km_adp_pk_phospho
Value: 0.33   (Units: mM)
Constant
 
   n_pk_phospho
Value: 2.9   (Units: dimensionless)
Constant
 
   vmax_pyrmalt
Value: 19400.0   (Units: per_h_per_mM)
Constant
 
   km_mal_mito_pyrmalt
Value: 0.7   (Units: mM)
Constant
 
   km_pyr_pyrmalt
Value: 0.84   (Units: mM)
Constant
 
   km_mal_pyrmalt
Value: 0.7   (Units: mM)
Constant
 
   km_pyr_mito_pyrmalt
Value: 0.84   (Units: mM)
Constant
 
   vmax_pyrt
Value: 1.94E8   (Units: per_h_per_mM)
Constant
 
   km_pyr_pyrt
Value: 0.15   (Units: mM)
Constant
 
   km_pyr_mito_pyrt
Value: 0.15   (Units: mM)
Constant
 
   vmax_tpi
Value: 1.94E8   (Units: per_h)
Constant
 
   keq_tpi
Value: 0.04545   (Units: dimensionless)
Constant
 
   km_dhap_tpi
Value: 0.59   (Units: mM)
Constant
 
   km_gap_tpi
Value: 0.415   (Units: mM)
Constant
 
   vmax_ugt
Value: 7.78E9   (Units: per_h_per_mM)
Constant
 
   keq_ugt
Value: 0.3122   (Units: dimensionless)
Constant
 
   km_utp_ugt
Value: 0.2   (Units: mM)
Constant
 
   km_glc1p_ugt
Value: 0.055   (Units: mM)
Constant
 
   km_udpglc_ugt
Value: 0.06   (Units: mM)
Constant
 
   km_pp_ugt
Value: 0.084   (Units: mM)
Constant
 
   ins0
Value: 3.1E-6   (Units: mM)
Constant
 
   n_ins
Value: 5.7   (Units: dimensionless)
Constant
 
   km_glc_ext_ins
Value: 7.7   (Units: mM)
Constant
 
   glucagon0
Value: 5.06E-7   (Units: mM)
Constant
 
   n_glucagon
Value: 5.65   (Units: dimensionless)
Constant
 
   km_glc_ext_glucagon
Value: 4.7   (Units: mM)
Constant
 
   glucagonb
Value: 4.0E-8   (Units: mM)
Constant
 
   n_ins_gamma
Value: 1.75   (Units: dimensionless)
Constant
 
   km_ins_gamma
Value: 7.0E-7   (Units: mM)
Constant
 
   n_glucagon_gamma
Value: 3.6   (Units: dimensionless)
Constant
 
   km_glucagon_gamma
Value: 8.0E-8   (Units: mM)
Constant
 
   ins
Value: NaN   (Units: mM)
 
   glucagon
Value: NaN   (Units: mM)
 
   gamma
Value: NaN   (Units: dimensionless)
 
   gamma_fbp1
Value: NaN   (Units: dimensionless)
 
   gamma_gp
Value: NaN   (Units: dimensionless)
 
   gamma_gs
Value: NaN   (Units: dimensionless)
 
   gamma_pfk2
Value: NaN   (Units: dimensionless)
 
   gamma_fbp2
Value: NaN   (Units: dimensionless)
 
   gamma_pk
Value: NaN   (Units: dimensionless)
 
   v_ALD
Value: NaN   (Units: mM_per_h)
 
   v_EN
Value: NaN   (Units: mM_per_h)
 
   v_fbp1_native
Value: NaN   (Units: dimensionless)
 
   v_fbp1_phospho
Value: NaN   (Units: dimensionless)
 
   v_FBP1
Value: NaN   (Units: mM_per_h)
 
   v_GAPDH
Value: NaN   (Units: mM_per_h)
 
   vmax_gk
Value: NaN   (Units: mM_per_h)
 
   v_GK
Value: NaN   (Units: dimensionless)
 
   v_GLUT2
Value: NaN   (Units: mM_per_h)
 
   v_GLCT_ER
Value: NaN   (Units: mM_per_h)
 
   vmax_gp
Value: NaN   (Units: mM_per_h)
 
   vmax_gp_native
Value: NaN   (Units: per_mMmM)
 
   km_glc1p_gp_native
Value: NaN   (Units: mM)
 
   vmax_gp_phospho
Value: NaN   (Units: per_mMmM)
 
   v_gp_native
Value: NaN   (Units: dimensionless)
 
   v_gp_phospho
Value: NaN   (Units: dimensionless)
 
   v_GP
Value: NaN   (Units: mM_per_h)
 
   v_G6P_ER
Value: NaN   (Units: mM_per_h)
 
   v_GPI
Value: NaN   (Units: mM_per_h)
 
   v_G1PI
Value: NaN   (Units: mM_per_h)
 
   v_G6PT_ER
Value: NaN   (Units: mM_per_h)
 
   vmax_gs
Value: NaN   (Units: mM_per_h)
 
   km_udpglc_gs_native
Value: NaN   (Units: mM)
 
   km_udpglc_gs_phospho
Value: NaN   (Units: mM)
 
   v_gs_native
Value: NaN   (Units: dimensionless)
 
   v_gs_phospho
Value: NaN   (Units: dimensionless)
 
   v_GS
Value: NaN   (Units: mM_per_h)
 
   v_LACT
Value: NaN   (Units: mM_per_h)
 
   v_LDH
Value: NaN   (Units: mM_per_h)
 
   v_MALT
Value: NaN   (Units: mM_per_h)
 
   v_MDH
Value: NaN   (Units: mM_per_h)
 
   v_MDH_mito
Value: NaN   (Units: mM_per_h)
 
   v_NDK_GTP
Value: NaN   (Units: mM_per_h)
 
   v_NDK_UTP
Value: NaN   (Units: mM_per_h)
 
   v_NDK_GTP_mito
Value: NaN   (Units: mM_per_h)
 
   v_PC
Value: NaN   (Units: mM_per_h)
 
   v_PEPCK
Value: NaN   (Units: mM_per_h)
 
   v_PEPCK_mito
Value: NaN   (Units: mM_per_h)
 
   keq_pept
Value: NaN   (Units: dimensionless)
 
   v_PEPT
Value: NaN   (Units: mM_per_h)
 
   km_atp_pfk1
Value: NaN   (Units: mM)
 
   ki_atp_pfk1
Value: NaN   (Units: mM)
 
   km_fru6p_pfk1
Value: NaN   (Units: mM)
 
   n_fru6p_pfk1
Value: NaN   (Units: dimensionless)
 
   v_PFK1
Value: NaN   (Units: mM_per_h)
 
   v_pfk2_native
Value: NaN   (Units: mM_per_h)
 
   v_pfk2_phospho
Value: NaN   (Units: mM_per_h)
 
   v_PFK2
Value: NaN   (Units: mM_per_h)
 
   v_fbp2_native
Value: NaN   (Units: mM_per_h)
 
   v_fbp2_phospho
Value: NaN   (Units: mM_per_h)
 
   v_FBP2
Value: NaN   (Units: mM_per_h)
 
   v_PGK
Value: NaN   (Units: mM_per_h)
 
   v_PGM
Value: NaN   (Units: mM_per_h)
 
   v_pk_native
Value: NaN   (Units: dimensionless)
 
   v_pk_phospho
Value: NaN   (Units: dimensionless)
 
   v_PK
Value: NaN   (Units: mM_per_h)
 
   v_PyrMalT
Value: NaN   (Units: mM_per_h)
 
   v_PyrT
Value: NaN   (Units: mM_per_h)
 
   v_TPI
Value: NaN   (Units: mM_per_h)
 
   v_UGT
Value: NaN   (Units: mM_per_h)
 
Representative curation result(s)
Representative curation result(s) of BIOMD0000000633

Curator's comment: (updated: 21 Apr 2017 16:45:48 BST)

Figure 6 from the paper was reproduced using SloppyCell. The model appears to be correct, and reproduces the figure qualitatively with small quantitative differences in the Glc exchange rate levels. Python script used to generate the figure can be found in the additional files.

Additional file(s)
  • Python_Script_to_generate_figure_6:
    Python script used to generate figure 6 from the paper.
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