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BIOMD0000000183 - Stefan2008 - calmodulin allostery

 

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Reference Publication
Publication ID: 18669651
Stefan MI, Edelstein SJ, Le Novère N.
An allosteric model of calmodulin explains differential activation of PP2B and CaMKII.
Proc. Natl. Acad. Sci. U.S.A. 2008 Aug; 105(31): 10768-10773
European Molecular Biology Laboratory-European Bioinformatics Institute, Wellcome Trust Genome Campus, Hinxton CB10 1SD, United Kingdom.  [more]
Model
Original Model: BIOMD0000000183.origin
Submitter: Melanie Stefan
Submission ID: MODEL9885984404
Submission Date: 14 Aug 2008 16:08:52 UTC
Last Modification Date: 04 Jun 2014 11:22:28 UTC
Creation Date: 15 Jul 2008 14:18:02 UTC
Encoders:  Melanie Stefan
   Lukas Endler
set #1
bqbiol:hasTaxon Taxonomy Mammalia
set #2
bqbiol:isVersionOf Gene Ontology detection of calcium ion
Gene Ontology positive regulation of synaptic plasticity
Gene Ontology protein serine/threonine phosphatase activity
Gene Ontology calmodulin-dependent protein kinase activity
Gene Ontology calmodulin binding
set #3
bqbiol:hasProperty Mathematical Modelling Ontology MAMO_0000046
set #4
bqbiol:isPartOf KEGG Pathway Long-term potentiation
Reactome REACT_9053.2
Notes
Stefan2008 - calmodulin allostery

An allosteric model for calmodulin activation, in which binding to calcium facilitates the transition between a low-affinity [tense (T)] and a high-affinity [relaxed (R)] state.

This model is described in the article:

Stefan MI, Edelstein SJ, Le Novère N
Proc. Natl. Acad. Sci. U.S.A. 2008 Aug; 105(31): 10768-10773

Abstract:

Calmodulin plays a vital role in mediating bidirectional synaptic plasticity by activating either calcium/calmodulin-dependent protein kinase II (CaMKII) or protein phosphatase 2B (PP2B) at different calcium concentrations. We propose an allosteric model for calmodulin activation, in which binding to calcium facilitates the transition between a low-affinity [tense (T)] and a high-affinity [relaxed (R)] state. The four calcium-binding sites are assumed to be nonidentical. The model is consistent with previously reported experimental data for calcium binding to calmodulin. It also accounts for known properties of calmodulin that have been difficult to model so far, including the activity of nonsaturated forms of calmodulin (we predict the existence of open conformations in the absence of calcium), an increase in calcium affinity once calmodulin is bound to a target, and the differential activation of CaMKII and PP2B depending on calcium concentration.

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: 18669651 Submission Date: 14 Aug 2008 16:08:52 UTC Last Modification Date: 04 Jun 2014 11:22:28 UTC Creation Date: 15 Jul 2008 14:18:02 UTC
Mathematical expressions
Reactions
Ca binding to camR site A Ca binding to camR site B Ca binding to camR site C Ca binding to camR site D
Ca dissociating from camR_ca1_A site A Ca dissociating from camR_ca1_B site B Ca dissociating from camR_ca1_C site C Ca dissociating from camR_ca1_D site D
Ca binding to camR_ca1_A site B Ca binding to camR_ca1_A site C Ca binding to camR_ca1_A site D Ca binding to camR_ca1_B site A
Ca binding to camR_ca1_B site C Ca binding to camR_ca1_B site D Ca binding to camR_ca1_C site A Ca binding to camR_ca1_C site B
Ca binding to camR_ca1_C site D Ca binding to camR_ca1_D site A Ca binding to camR_ca1_D site B Ca binding to camR_ca1_D site C
Ca dissociating from camR_ca2_AB site B Ca dissociating from camR_ca2_AC site C Ca dissociating from camR_ca2_AD site D Ca dissociating from camR_ca2_AB site A
Ca dissociating from camR_ca2_BC site C Ca dissociating from camR_ca2_BD site D Ca dissociating from camR_ca2_AC site A Ca dissociating from camR_ca2_BC site B
Ca dissociating from camR_ca2_CD site D Ca dissociating from camR_ca2_AD site A Ca dissociating from camR_ca2_BD site B Ca dissociating from camR_ca2_CD site C
Ca binding to camR_ca2_AB site C Ca binding to camR_ca2_AB site D Ca binding to camR_ca2_AC site B Ca binding to camR_ca2_AC site D
Ca binding to camR_ca2_AD site B Ca binding to camR_ca2_AD site C Ca binding to camR_ca2_BC site A Ca binding to camR_ca2_BC site D
Ca binding to camR_ca2_BD site A Ca binding to camR_ca2_BD site C Ca binding to camR_ca2_CD site A Ca binding to camR_ca2_CD site B
Ca dissociating from camR_ca3_ABC site A Ca dissociating from camR_ca3_ABC site B Ca dissociating from camR_ca3_ABC site C Ca dissociating from camR_ca3_ABD site A
Ca dissociating from camR_ca3_ABD site B Ca dissociating from camR_ca3_ABD site D Ca dissociating from camR_ca3_ACD site A Ca dissociating from camR_ca3_ACD site C
Ca dissociating from camR_ca3_ACD site D Ca dissociating from camR_ca3_BCD site B Ca dissociating from camR_ca3_BCD site C Ca dissociating from camR_ca3_BCD site D
Ca binding to camR_ca3_ABC site D Ca binding to camR_ca3_ABD site C Ca binding to camR_ca3_ACD site B Ca binding to camR_ca3_BCD site A
Ca dissociating from camR_ca4_ABCD site D Ca dissociating from camR_ca4_ABCD site C Ca dissociating from camR_ca4_ABCD site B Ca dissociating from camR_ca4_ABCD site A
Ca binding to camT site A Ca binding to camT site B Ca binding to camT site C Ca binding to camT site D
Ca dissociating from camT_ca1_A site A Ca dissociating from camT_ca1_B site B Ca dissociating from camT_ca1_C site C Ca dissociating from camT_ca1_D site D
Ca binding to camT_ca1_A site B Ca binding to camT_ca1_A site C Ca binding to camT_ca1_A site D Ca binding to camT_ca1_B site A
Ca binding to camT_ca1_B site C Ca binding to camT_ca1_B site D Ca binding to camT_ca1_C site A Ca binding to camT_ca1_C site B
Ca binding to camT_ca1_C site D Ca binding to camT_ca1_D site A Ca binding to camT_ca1_D site B Ca binding to camT_ca1_D site C
Ca dissociating from camT_ca2_AB site A Ca dissociating from camT_ca2_AB site B Ca dissociating from camT_ca2_AC site A Ca dissociating from camT_ca2_AC site C
Ca dissociating from camT_ca2_AD site A Ca dissociating from camT_ca2_AD site D Ca dissociating from camT_ca2_BC site B Ca dissociating from camT_ca2_BC site C
Ca dissociating from camT_ca2_BD site B Ca dissociating from camT_ca2_BD site D Ca dissociating from camT_ca2_CD site C Ca dissociating from camT_ca2_CD site D
Ca binding to camT_ca2_AB site C Ca binding to camT_ca2_AB site D Ca binding to camT_ca2_AC site B Ca binding to camT_ca2_AC site D
Ca binding to camT_ca2_AD site B Ca binding to camT_ca2_AD site C Ca binding to camT_ca2_BC site A Ca binding to camT_ca2_BC site D
Ca binding to camT_ca2_BD site A Ca binding to camT_ca2_BD site C Ca binding to camT_ca2_CD site A Ca binding to camT_ca2_CD site B
Ca dissociating from camT_ca3_ABC site B Ca dissociating from camT_ca3_ABC site A Ca dissociating from camT_ca3_ABD site D Ca dissociating from camT_ca3_ABD site B
Ca dissociating from camT_ca3_ABD site A Ca dissociating from camT_ca3_ACD site D Ca dissociating from camT_ca3_ACD site C Ca dissociating from camT_ca3_ACD site A
Ca dissociating from camT_ca3_BCD site D Ca dissociating from camT_ca3_BCD site C Ca dissociating from camT_ca3_BCD site B Ca binding to camT_ca3_ABC site D
Ca binding to camT_ca3_ABD site C Ca binding to camT_ca3_ACD site B Ca binding to camT_ca3_BCD site A Ca dissociating from camT_ca4_ABCD site D
Ca dissociating from camT_ca4_ABCD site C Ca dissociating from camT_ca4_ABCD site B Ca dissociating from camT_ca4_ABCD site A Transition camR to camT
Transition camT to camR Transition camR_ca1_A to camT_ca1_A Transition camR_ca1_B to camT_ca1_B Transition camR_ca1_C to camT_ca1_C
Transition camR_ca1_D to camT_ca1_D Transition camT_ca1_A to camR_ca1_A Transition camT_ca1_B to camR_ca1_B Transition camT_ca1_C to camR_ca1_C
Transition camT_ca1_D to camR_ca1_D Transition camR_ca2_AB to camT_ca2_AB Transition camR_ca2_AC to camT_ca2_AC Transition camR_ca2_AD to camT_ca2_AD
Transition camR_ca2_BC to camT_ca2_BC Transition camR_ca2_BD to camT_ca2_BD Transition camR_ca2_CD to camT_ca2_CD Transition camT_ca2_AB to camR_ca2_AB
Transition camT_ca2_AC to camR_ca2_AC Transition camT_ca2_AD to camR_ca2_AD Transition camT_ca2_BC to camR_ca2_BC Transition camT_ca2_BD to camR_ca2_BD
Transition camT_ca2_CD to camR_ca2_CD Transition camR_ca3_ABC to camT_ca3_ABC Transition camR_ca3_ABD to camT_ca3_ABD Transition camR_ca3_ACD to camT_ca3_ACD
Transition camR_ca3_BCD to camT_ca3_BCD Transition camT_ca3_ABC to camR_ca3_ABC Transition camT_ca3_ABD to camR_ca3_ABD Transition camT_ca3_ACD to camR_ca3_ACD
Transition camT_ca3_BCD to camR_ca3_BCD Transition camR_ca4_ABCD to camT_ca4_ABCD Transition camT_ca4_ABCD to camR_ca4_ABCD CaMKII binding to camR
CaMKII binding to camR_ca1_A CaMKII binding to camR_ca1_B CaMKII binding to camR_ca1_C CaMKII binding to camR_ca1_D
CaMKII binding to camR_ca2_AB CaMKII binding to camR_ca2_AC CaMKII binding to camR_ca2_AD CaMKII binding to camR_ca2_BC
CaMKII binding to camR_ca2_BD CaMKII binding to camR_ca2_CD CaMKII binding to camR_ca3_ABC CaMKII binding to camR_ca3_ABD
CaMKII binding to camR_ca3_ACD CaMKII binding to camR_ca3_BCD CaMKII binding to camR_ca4_ABCD CaMKII dissociation from camR
CaMKII dissociation from camR_ca1_A CaMKII dissociation from camR_ca1_B CaMKII dissociation from camR_ca1_C CaMKII dissociation from camR_ca1_D
CaMKII dissociation from camR_ca2_AB CaMKII dissociation from camR_ca2_AC CaMKII dissociation from camR_ca2_AD CaMKII dissociation from camR_ca2_BC
CaMKII dissociation from camR_ca2_BD CaMKII dissociation from camR_ca2_CD CaMKII dissociation from camR_ca3_ABC CaMKII dissociation from camR_ca3_ABD
CaMKII dissociation from camR_ca3_ACD CaMKII dissociation from camR_ca3_BCD CaMKII dissociation from camR_ca4_ABCD PP2B binding to camR
PP2B binding to camR_ca1_A PP2B binding to camR_ca1_B PP2B binding to camR_ca1_C PP2B binding to camR_ca1_D
PP2B binding to camR_ca2_AB PP2B binding to camR_ca2_AC PP2B binding to camR_ca2_AD PP2B binding to camR_ca2_BC
PP2B binding to camR_ca2_BD PP2B binding to camR_ca2_CD PP2B binding to camR_ca3_ABC PP2B binding to camR_ca3_ABD
PP2B binding to camR_ca3_ACD PP2B binding to camR_ca3_BCD PP2B binding to camR_ca4_ABCD PP2B dissociation from camR
PP2B dissociation from camR_ca1_A PP2B dissociation from camR_ca1_B PP2B dissociation from camR_ca1_C PP2B dissociation from camR_ca1_D
PP2B dissociation from camR_ca2_AB PP2B dissociation from camR_ca2_AC PP2B dissociation from camR_ca2_AD PP2B dissociation from camR_ca2_BC
PP2B dissociation from camR_ca2_BD PP2B dissociation from camR_ca2_CD PP2B dissociation from camR_ca3_ABC PP2B dissociation from camR_ca3_ABD
PP2B dissociation from camR_ca3_ACD PP2B dissociation from camR_ca3_BCD PP2B dissociation from camR_ca4_ABCD Ca binding to camR_CaMKII site A
Ca binding to camR_CaMKII site B Ca binding to camR_CaMKII site C Ca binding to camR_CaMKII site D Ca dissociation from camR_ca1_CaMKII site A
Ca dissociation from camR_ca1_CaMKII site C Ca dissociation from camR_ca1_CaMKII site D Ca binding to camR_ca1_A_CaMKII site B Ca binding to camR_ca1_A_CaMKII site C
Ca binding to camR_ca1_A_CaMKII site D Ca binding to camR_ca1_B_CaMKII site A Ca binding to camR_ca1_B_CaMKII site C Ca binding to camR_ca1_B_CaMKII site D
Ca binding to camR_ca1_C_CaMKII site A Ca binding to camR_ca1_C_CaMKII site B Ca binding to camR_ca1_C_CaMKII site D Ca binding to camR_ca1_D_CaMKII site A
Ca binding to camR_ca1_D_CaMKII site B Ca binding to camR_ca1_D_CaMKII site C Ca dissociation from camR_ca2_AB_CaMKII site A Ca dissociation from camR_ca2_AB_CaMKII site B
Ca dissociation from camR_ca2_AC_CaMKII site A Ca dissociation from camR_ca2_AC_CaMKII site C Ca dissociation from camR_ca2_AD_CaMKII site A Ca dissociation from camR_ca2_AD_CaMKII site D
Ca dissociation from camR_ca2_BC_CaMKII site B Ca dissociation from camR_ca2_BC_CaMKII site C Ca dissociation from camR_ca2_BD_CaMKII site B Ca dissociation from camR_ca2_BD_CaMKII site D
Ca dissociation from camR_ca2_CD_CaMKII site C Ca dissociation from camR_ca2_CD_CaMKII site D Ca binding to camR_ca2_AB_CaMKII site C Ca binding to camR_ca2_AB_CaMKII site D
Ca binding to camR_ca2_AC_CaMKII site B Ca binding to camR_ca2_AC_CaMKII site D Ca binding to camR_ca2_AD_CaMKII site B Ca binding to camR_ca2_AD_CaMKII site C
Ca binding to camR_ca2_BC_CaMKII site A Ca binding to camR_ca2_BC_CaMKII site D Ca binding to camR_ca2_BD_CaMKII site A Ca binding to camR_ca2_BD_CaMKII site C
Ca binding to camR_ca2_CD_CaMKII site A Ca binding to camR_ca2_CD_CaMKII site B Ca dissociation from camR_ca3_ABC_CaMKII site C Ca dissociation from camR_ca3_ABC_CaMKII site B
Ca dissociation from camR_ca3_ABC_CaMKII site A Ca dissociation from camR_ca3_ABD_CaMKII site D Ca dissociation from camR_ca3_ABD_CaMKII site B Ca dissociation from camR_ca3_ABD_CaMKII site A
Ca dissociation from camR_ca3_ACD_CaMKII site D Ca dissociation from camR_ca3_ACD_CaMKII site C Ca dissociation from camR_ca3_ACD_CaMKII site A Ca dissociation from camR_ca3_BCD_CaMKII site D
Ca dissociation from camR_ca3_BCD_CaMKII site C Ca dissociation from camR_ca3_BCD_CaMKII site B Ca binding to camR_ca3_BCD_CaMKII site A Ca binding to camR_ca3_ACD_CaMKII site B
Ca binding to camR_ca3_ABD_CaMKII site C Ca binding to camR_ca3_ABC_CaMKII site D Ca dissociation from camR_ca4_ABCD_CaMKII site A Ca dissociation from camR_ca4_ABCD_CaMKII site B
Ca dissociation from camR_ca4_ABCD_CaMKII site C Ca dissociation from camR_ca4_ABCD_CaMKII site D Ca binding to camR_PP2B site A Ca binding to camR_PP2B site B
Ca binding to camR_PP2B site C Ca binding to camR_PP2B site D Ca dissociation from camR_ca1_A_PP2B site A Ca dissociation from camR_ca1_B_PP2B site B
Ca dissociation from camR_ca1_C_PP2B site C Ca dissociation from camR_ca1_D_PP2B site D Ca binding to camR_ca1_A_PP2B site B Ca binding to camR_ca1_A_PP2B site C
Ca binding to camR_ca1_A_PP2B site D Ca binding to camR_ca1_B_PP2B site A Ca binding to camR_ca1_B_PP2B site C Ca binding to camR_ca1_B_PP2B site D
Ca binding to camR_ca1_C_PP2B site A Ca binding to camR_ca1_C_PP2B site B Ca binding to camR_ca1_C_PP2B site D Ca binding to camR_ca1_D_PP2B site A
Ca binding to camR_ca1_D_PP2B site B Ca binding to camR_ca1_D_PP2B site C Ca dissociating from camR_ca2_AB_PP2B site A Ca dissociating from camR_ca2_AB_PP2B site B
Ca dissociating from camR_ca2_AC_PP2B site A Ca dissociating from camR_ca2_AC_PP2B site C Ca dissociating from camR_ca2_AD_PP2B site A Ca dissociating from camR_ca2_AD_PP2B site D
Ca dissociating from camR_ca2_BC_PP2B site B Ca dissociating from camR_ca2_BC_PP2B site C Ca dissociating from camR_ca2_BD_PP2B site B Ca dissociating from camR_ca2_BD_PP2B site D
Ca dissociating from camR_ca2_CD_PP2B site C Ca dissociating from camR_ca2_CD_PP2B site D Ca binding to camR_ca2_AB_PP2B site C Ca binding to camR_ca2_AB_PP2B site D
Ca binding to camR_ca2_AC_PP2B site B Ca binding to camR_ca2_AC_PP2B site D Ca binding to camR_ca2_AD_PP2B site B Ca binding to camR_ca2_AD_PP2B site C
Ca binding to camR_ca2_BC_PP2B site A Ca binding to camR_ca2_BC_PP2B site D Ca binding to camR_ca2_BD_PP2B site A Ca binding to camR_ca2_BD_PP2B site C
Ca binding to camR_ca2_CD_PP2B site A Ca binding to camR_ca2_CD_PP2B site B Ca dissociation from camR_ca3_ABC_PP2B site A Ca dissociation from camR_ca3_ABC_PP2B site B
Ca dissociation from camR_ca3_ABC_PP2B site C Ca dissociation from camR_ca3_ABD_PP2B site A Ca dissociation from camR_ca3_ABD_PP2B site B Ca dissociation from camR_ca3_ABD_PP2B site D
Ca dissociation from camR_ca3_ACD_PP2B site A Ca dissociation from camR_ca3_ACD_PP2B site C Ca dissociation from camR_ca3_ACD_PP2B site D Ca dissociation from camR_ca3_BCD_PP2B site B
Ca dissociation from camR_ca3_BCD_PP2B site C Ca dissociation from camR_ca3_BCD_PP2B site D Ca binding to camR_ca3_ABC_PP2B site D Ca binding to camR_ca3_ABD_PP2B site C
Ca binding to camR_ca3_ACD_PP2B site B Ca binding to camR_ca3_BCD_PP2B site A Ca dissociating from camR_ca4_ABCD_PP2B site A Ca dissociating from camR_ca4_ABCD_PP2B site B
Ca dissociating from camR_ca4_ABCD_PP2B site C Ca dissociating from camR_ca4_ABCD_PP2B site D Ca dissociation from camR_ca1_CaMKII site B Ca dissociating from camT_ca3_ABC site C
Rules
Assignment Rule (variable: koffRA) Assignment Rule (variable: free_camT_ca3_total) Assignment Rule (variable: koffRB) Assignment Rule (variable: koffRC)
Assignment Rule (variable: koffRD) Assignment Rule (variable: koffTA) Assignment Rule (variable: koffTB) Assignment Rule (variable: koffTC)
Assignment Rule (variable: koffTD) Assignment Rule (variable: kTR) Assignment Rule (variable: free_camR_ca3_total) Assignment Rule (variable: free_cam_ca3_total)
Assignment Rule (variable: cam_ca4_total) Assignment Rule (variable: CaMKII_camR_ca1) Assignment Rule (variable: CaMKII_camR_ca3) Assignment Rule (variable: CaMKII_camR_ca2)
Assignment Rule (variable: total_CaMKII_bound) Assignment Rule (variable: total CaMKII) Assignment Rule (variable: CaMKIIbar) Assignment Rule (variable: PP2B_camR_ca1)
Assignment Rule (variable: PP2B_camR_ca2) Assignment Rule (variable: PP2B_camR_ca3) Assignment Rule (variable: total_PP2B_bound) Assignment Rule (variable: camR_unbound)
Assignment Rule (variable: total PP2B) Assignment Rule (variable: PP2Bbar) Assignment Rule (variable: total camT) Assignment Rule (variable: total camR)
Assignment Rule (variable: cam_ca3_total) Assignment Rule (variable: Rbar) Assignment Rule (variable: free_camR_ca2_total) Assignment Rule (variable: free_camT_ca2_total)
Assignment Rule (variable: free_cam_ca2_total) Assignment Rule (variable: cam_ca2_total) Assignment Rule (variable: free_cam_ca1_total) Assignment Rule (variable: cam_ca1_total)
Assignment Rule (variable: cam_ca0_total) Assignment Rule (variable: cam_total) Assignment Rule (variable: moles_bound_ca_per_moles_cam) Assignment Rule (variable: alpha)
Assignment Rule (variable: beta) Assignment Rule (variable: gamma) Assignment Rule (variable: delta) Assignment Rule (variable: epsilon)
Assignment Rule (variable: ybar) Assignment Rule (variable: ybar/(1-ybar))    
Physical entities
Compartments Species
Spine camR ca camR_ca1_A
camR_ca1_B camR_ca1_C camR_ca1_D
camR_ca2_AB camR_ca2_AC camR_ca2_AD
camR_ca2_BC camR_ca2_BD camR_ca2_CD
camR_ca3_ABC camR_ca3_ABD camR_ca3_ACD
camR_ca3_BCD camR_ca4_ABCD camT
camT_ca1_A camT_ca1_B camT_ca1_C
camT_ca1_D camT_ca2_AB camT_ca2_AC
camT_ca2_AD camT_ca2_BC camT_ca2_BD
camT_ca2_CD camT_ca3_ABC camT_ca3_ABD
camT_ca3_ACD camT_ca3_BCD camT_ca4_ABCD
CaMKII camR_CaMKII camR_ca1_A_CaMKII
camR_ca1_B_CaMKII camR_ca1_C_CaMKII camR_ca1_D_CaMKII
camR_ca2_AB_CaMKII camR_ca2_AC_CaMKII camR_ca2_AD_CaMKII
camR_ca2_BC_CaMKII camR_ca2_BD_CaMKII camR_ca2_CD_CaMKII
camR_ca3_ABC_CaMKII camR_ca3_ABD_CaMKII camR_ca3_ACD_CaMKII
camR_ca3_BCD_CaMKII camR_ca4_ABCD_CaMKII PP2B
camR_PP2B camR_ca1_A_PP2B camR_ca1_B_PP2B
camR_ca1_C_PP2B camR_ca1_D_PP2B camR_ca2_AB_PP2B
camR_ca2_AC_PP2B camR_ca2_AD_PP2B camR_ca2_BC_PP2B
camR_ca2_BD_PP2B camR_ca2_CD_PP2B camR_ca3_ABC_PP2B
camR_ca3_ABD_PP2B camR_ca3_ACD_PP2B camR_ca3_BCD_PP2B
camR_ca4_ABCD_PP2B    
Global parameters
kon koffRA koffRB koffRC
koffRD koffTA koffTB koffTC
koffTD kRT kTR cA
cB cC cD konCaMKII
koffCaMKII konPP2B koffPP2B free_camR_ca3_total
free_camT_ca3_total free_cam_ca3_total cam_ca4_total CaMKII_camR_ca1
CaMKII_camR_ca2 CaMKII_camR_ca3 total_CaMKII_bound total CaMKII
CaMKIIbar PP2B_camR_ca1 PP2B_camR_ca2 PP2B_camR_ca3
total_PP2B_bound total PP2B PP2Bbar camR_unbound
total camR total camT Rbar cam_ca3_total
free_camR_ca2_total free_camT_ca2_total free_cam_ca2_total cam_ca2_total
free_cam_ca1_total cam_ca1_total cam_ca0_total cam_total
moles_bound_ca_per_moles_cam L KRA KRB
KRC KRD alpha beta
gamma delta epsilon ybar
ybar/(1-ybar)      
Reactions (352)
 
 Ca binding to camR site A [camR] + [ca] → [camR_ca1_A];  
 
 Ca binding to camR site B [camR] + [ca] → [camR_ca1_B];  
 
 Ca binding to camR site C [camR] + [ca] → [camR_ca1_C];  
 
 Ca binding to camR site D [camR] + [ca] → [camR_ca1_D];  
 
 Ca dissociating from camR_ca1_A site A [camR_ca1_A] → [camR] + [ca];  
 
 Ca dissociating from camR_ca1_B site B [camR_ca1_B] → [camR] + [ca];  
 
 Ca dissociating from camR_ca1_C site C [camR_ca1_C] → [camR] + [ca];  
 
 Ca dissociating from camR_ca1_D site D [camR_ca1_D] → [camR] + [ca];  
 
 Ca binding to camR_ca1_A site B [camR_ca1_A] + [ca] → [camR_ca2_AB];  
 
 Ca binding to camR_ca1_A site C [camR_ca1_A] + [ca] → [camR_ca2_AC];  
 
 Ca binding to camR_ca1_A site D [camR_ca1_A] + [ca] → [camR_ca2_AD];  
 
 Ca binding to camR_ca1_B site A [camR_ca1_B] + [ca] → [camR_ca2_AB];  
 
 Ca binding to camR_ca1_B site C [camR_ca1_B] + [ca] → [camR_ca2_BC];  
 
 Ca binding to camR_ca1_B site D [camR_ca1_B] + [ca] → [camR_ca2_BD];  
 
 Ca binding to camR_ca1_C site A [camR_ca1_C] + [ca] → [camR_ca2_AC];  
 
 Ca binding to camR_ca1_C site B [camR_ca1_C] + [ca] → [camR_ca2_BC];  
 
 Ca binding to camR_ca1_C site D [camR_ca1_C] + [ca] → [camR_ca2_CD];  
 
 Ca binding to camR_ca1_D site A [camR_ca1_D] + [ca] → [camR_ca2_AD];  
 
 Ca binding to camR_ca1_D site B [camR_ca1_D] + [ca] → [camR_ca2_BD];  
 
 Ca binding to camR_ca1_D site C [camR_ca1_D] + [ca] → [camR_ca2_CD];  
 
 Ca dissociating from camR_ca2_AB site B [camR_ca2_AB] → [camR_ca1_A] + [ca];  
 
 Ca dissociating from camR_ca2_AC site C [camR_ca2_AC] → [camR_ca1_A] + [ca];  
 
 Ca dissociating from camR_ca2_AD site D [camR_ca2_AD] → [camR_ca1_A] + [ca];  
 
 Ca dissociating from camR_ca2_AB site A [camR_ca2_AB] → [camR_ca1_B] + [ca];  
 
 Ca dissociating from camR_ca2_BC site C [camR_ca2_BC] → [camR_ca1_B] + [ca];  
 
 Ca dissociating from camR_ca2_BD site D [camR_ca2_BD] → [camR_ca1_B] + [ca];  
 
 Ca dissociating from camR_ca2_AC site A [camR_ca2_AC] → [camR_ca1_C] + [ca];  
 
 Ca dissociating from camR_ca2_BC site B [camR_ca2_BC] → [camR_ca1_C] + [ca];  
 
 Ca dissociating from camR_ca2_CD site D [camR_ca2_CD] → [camR_ca1_C] + [ca];  
 
 Ca dissociating from camR_ca2_AD site A [camR_ca2_AD] → [camR_ca1_D] + [ca];  
 
 Ca dissociating from camR_ca2_BD site B [camR_ca2_BD] → [camR_ca1_D] + [ca];  
 
 Ca dissociating from camR_ca2_CD site C [camR_ca2_CD] → [camR_ca1_D] + [ca];  
 
 Ca binding to camR_ca2_AB site C [camR_ca2_AB] + [ca] → [camR_ca3_ABC];  
 
 Ca binding to camR_ca2_AB site D [camR_ca2_AB] + [ca] → [camR_ca3_ABD];  
 
 Ca binding to camR_ca2_AC site B [camR_ca2_AC] + [ca] → [camR_ca3_ABC];  
 
 Ca binding to camR_ca2_AC site D [camR_ca2_AC] + [ca] → [camR_ca3_ACD];  
 
 Ca binding to camR_ca2_AD site B [camR_ca2_AD] + [ca] → [camR_ca3_ABD];  
 
 Ca binding to camR_ca2_AD site C [camR_ca2_AD] + [ca] → [camR_ca3_ACD];  
 
 Ca binding to camR_ca2_BC site A [camR_ca2_BC] + [ca] → [camR_ca3_ABC];  
 
 Ca binding to camR_ca2_BC site D [camR_ca2_BC] + [ca] → [camR_ca3_BCD];  
 
 Ca binding to camR_ca2_BD site A [camR_ca2_BD] + [ca] → [camR_ca3_ABD];  
 
 Ca binding to camR_ca2_BD site C [camR_ca2_BD] + [ca] → [camR_ca3_BCD];  
 
 Ca binding to camR_ca2_CD site A [camR_ca2_CD] + [ca] → [camR_ca3_ACD];  
 
 Ca binding to camR_ca2_CD site B [camR_ca2_CD] + [ca] → [camR_ca3_BCD];  
 
 Ca dissociating from camR_ca3_ABC site A [camR_ca3_ABC] → [camR_ca2_BC] + [ca];  
 
 Ca dissociating from camR_ca3_ABC site B [camR_ca3_ABC] → [camR_ca2_AC] + [ca];  
 
 Ca dissociating from camR_ca3_ABC site C [camR_ca3_ABC] → [camR_ca2_AB] + [ca];  
 
 Ca dissociating from camR_ca3_ABD site A [camR_ca3_ABD] → [camR_ca2_BD] + [ca];  
 
 Ca dissociating from camR_ca3_ABD site B [camR_ca3_ABD] → [camR_ca2_AD] + [ca];  
 
 Ca dissociating from camR_ca3_ABD site D [camR_ca3_ABD] → [camR_ca2_AB] + [ca];  
 
 Ca dissociating from camR_ca3_ACD site A [camR_ca3_ACD] → [camR_ca2_CD] + [ca];  
 
 Ca dissociating from camR_ca3_ACD site C [camR_ca3_ACD] → [camR_ca2_AD] + [ca];  
 
 Ca dissociating from camR_ca3_ACD site D [camR_ca3_ACD] → [camR_ca2_AC] + [ca];  
 
 Ca dissociating from camR_ca3_BCD site B [camR_ca3_BCD] → [camR_ca2_CD] + [ca];  
 
 Ca dissociating from camR_ca3_BCD site C [camR_ca3_BCD] → [camR_ca2_BD] + [ca];  
 
 Ca dissociating from camR_ca3_BCD site D [camR_ca3_BCD] → [camR_ca2_BC] + [ca];  
 
 Ca binding to camR_ca3_ABC site D [camR_ca3_ABC] + [ca] → [camR_ca4_ABCD];  
 
 Ca binding to camR_ca3_ABD site C [camR_ca3_ABD] + [ca] → [camR_ca4_ABCD];  
 
 Ca binding to camR_ca3_ACD site B [camR_ca3_ACD] + [ca] → [camR_ca4_ABCD];  
 
 Ca binding to camR_ca3_BCD site A [camR_ca3_BCD] + [ca] → [camR_ca4_ABCD];  
 
 Ca dissociating from camR_ca4_ABCD site D [camR_ca4_ABCD] → [camR_ca3_ABC] + [ca];  
 
 Ca dissociating from camR_ca4_ABCD site C [camR_ca4_ABCD] → [camR_ca3_ABD] + [ca];  
 
 Ca dissociating from camR_ca4_ABCD site B [camR_ca4_ABCD] → [camR_ca3_ACD] + [ca];  
 
 Ca dissociating from camR_ca4_ABCD site A [camR_ca4_ABCD] → [camR_ca3_BCD] + [ca];  
 
 Ca binding to camT site A [camT] + [ca] → [camT_ca1_A];  
 
 Ca binding to camT site B [camT] + [ca] → [camT_ca1_B];  
 
 Ca binding to camT site C [camT] + [ca] → [camT_ca1_C];  
 
 Ca binding to camT site D [camT] + [ca] → [camT_ca1_D];  
 
 Ca dissociating from camT_ca1_A site A [camT_ca1_A] → [camT] + [ca];  
 
 Ca dissociating from camT_ca1_B site B [camT_ca1_B] → [camT] + [ca];  
 
 Ca dissociating from camT_ca1_C site C [camT_ca1_C] → [camT] + [ca];  
 
 Ca dissociating from camT_ca1_D site D [camT_ca1_D] → [camT] + [ca];  
 
 Ca binding to camT_ca1_A site B [camT_ca1_A] + [ca] → [camT_ca2_AB];  
 
 Ca binding to camT_ca1_A site C [camT_ca1_A] + [ca] → [camT_ca2_AC];  
 
 Ca binding to camT_ca1_A site D [camT_ca1_A] + [ca] → [camT_ca2_AD];  
 
 Ca binding to camT_ca1_B site A [camT_ca1_B] + [ca] → [camT_ca2_AB];  
 
 Ca binding to camT_ca1_B site C [camT_ca1_B] + [ca] → [camT_ca2_BC];  
 
 Ca binding to camT_ca1_B site D [camT_ca1_B] + [ca] → [camT_ca2_BD];  
 
 Ca binding to camT_ca1_C site A [camT_ca1_C] + [ca] → [camT_ca2_AC];  
 
 Ca binding to camT_ca1_C site B [camT_ca1_C] + [ca] → [camT_ca2_BC];  
 
 Ca binding to camT_ca1_C site D [camT_ca1_C] + [ca] → [camT_ca2_CD];  
 
 Ca binding to camT_ca1_D site A [camT_ca1_D] + [ca] → [camT_ca2_AD];  
 
 Ca binding to camT_ca1_D site B [camT_ca1_D] + [ca] → [camT_ca2_BD];  
 
 Ca binding to camT_ca1_D site C [camT_ca1_D] + [ca] → [camT_ca2_CD];  
 
 Ca dissociating from camT_ca2_AB site A [camT_ca2_AB] → [camT_ca1_B] + [ca];  
 
 Ca dissociating from camT_ca2_AB site B [camT_ca2_AB] → [camT_ca1_A] + [ca];  
 
 Ca dissociating from camT_ca2_AC site A [camT_ca2_AC] → [camT_ca1_C] + [ca];  
 
 Ca dissociating from camT_ca2_AC site C [camT_ca2_AC] → [camT_ca1_A] + [ca];  
 
 Ca dissociating from camT_ca2_AD site A [camT_ca2_AD] → [camT_ca1_D] + [ca];  
 
 Ca dissociating from camT_ca2_AD site D [camT_ca2_AD] → [camT_ca1_A] + [ca];  
 
 Ca dissociating from camT_ca2_BC site B [camT_ca2_BC] → [camT_ca1_C] + [ca];  
 
 Ca dissociating from camT_ca2_BC site C [camT_ca2_BC] → [camT_ca1_B] + [ca];  
 
 Ca dissociating from camT_ca2_BD site B [camT_ca2_BD] → [camT_ca1_D] + [ca];  
 
 Ca dissociating from camT_ca2_BD site D [camT_ca2_BD] → [camT_ca1_B] + [ca];  
 
 Ca dissociating from camT_ca2_CD site C [camT_ca2_CD] → [camT_ca1_D] + [ca];  
 
 Ca dissociating from camT_ca2_CD site D [camT_ca2_CD] → [camT_ca1_C] + [ca];  
 
 Ca binding to camT_ca2_AB site C [camT_ca2_AB] + [ca] → [camT_ca3_ABC];  
 
 Ca binding to camT_ca2_AB site D [camT_ca2_AB] + [ca] → [camT_ca3_ABD];  
 
 Ca binding to camT_ca2_AC site B [camT_ca2_AC] + [ca] → [camT_ca3_ABC];  
 
 Ca binding to camT_ca2_AC site D [camT_ca2_AC] + [ca] → [camT_ca3_ACD];  
 
 Ca binding to camT_ca2_AD site B [camT_ca2_AD] + [ca] → [camT_ca3_ABD];  
 
 Ca binding to camT_ca2_AD site C [camT_ca2_AD] + [ca] → [camT_ca3_ACD];  
 
 Ca binding to camT_ca2_BC site A [camT_ca2_BC] + [ca] → [camT_ca3_ABC];  
 
 Ca binding to camT_ca2_BC site D [camT_ca2_BC] + [ca] → [camT_ca3_BCD];  
 
 Ca binding to camT_ca2_BD site A [camT_ca2_BD] + [ca] → [camT_ca3_ABD];  
 
 Ca binding to camT_ca2_BD site C [camT_ca2_BD] + [ca] → [camT_ca3_BCD];  
 
 Ca binding to camT_ca2_CD site A [camT_ca2_CD] + [ca] → [camT_ca3_ACD];  
 
 Ca binding to camT_ca2_CD site B [camT_ca2_CD] + [ca] → [camT_ca3_BCD];  
 
 Ca dissociating from camT_ca3_ABC site B [camT_ca3_ABC] → [camT_ca2_AC] + [ca];  
 
 Ca dissociating from camT_ca3_ABC site A [camT_ca3_ABC] → [camT_ca2_BC] + [ca];  
 
 Ca dissociating from camT_ca3_ABD site D [camT_ca3_ABD] → [camT_ca2_AB] + [ca];  
 
 Ca dissociating from camT_ca3_ABD site B [camT_ca3_ABD] → [camT_ca2_AD] + [ca];  
 
 Ca dissociating from camT_ca3_ABD site A [camT_ca3_ABD] → [camT_ca2_BD] + [ca];  
 
 Ca dissociating from camT_ca3_ACD site D [camT_ca3_ACD] → [camT_ca2_AC] + [ca];  
 
 Ca dissociating from camT_ca3_ACD site C [camT_ca3_ACD] → [camT_ca2_AD] + [ca];  
 
 Ca dissociating from camT_ca3_ACD site A [camT_ca3_ACD] → [camT_ca2_CD] + [ca];  
 
 Ca dissociating from camT_ca3_BCD site D [camT_ca3_BCD] → [camT_ca2_BC] + [ca];  
 
 Ca dissociating from camT_ca3_BCD site C [camT_ca3_BCD] → [camT_ca2_BD] + [ca];  
 
 Ca dissociating from camT_ca3_BCD site B [camT_ca3_BCD] → [camT_ca2_CD] + [ca];  
 
 Ca binding to camT_ca3_ABC site D [camT_ca3_ABC] + [ca] → [camT_ca4_ABCD];  
 
 Ca binding to camT_ca3_ABD site C [camT_ca3_ABD] + [ca] → [camT_ca4_ABCD];  
 
 Ca binding to camT_ca3_ACD site B [camT_ca3_ACD] + [ca] → [camT_ca4_ABCD];  
 
 Ca binding to camT_ca3_BCD site A [camT_ca3_BCD] + [ca] → [camT_ca4_ABCD];  
 
 Ca dissociating from camT_ca4_ABCD site D [camT_ca4_ABCD] → [camT_ca3_ABC] + [ca];  
 
 Ca dissociating from camT_ca4_ABCD site C [camT_ca4_ABCD] → [camT_ca3_ABD] + [ca];  
 
 Ca dissociating from camT_ca4_ABCD site B [camT_ca4_ABCD] → [camT_ca3_ACD] + [ca];  
 
 Ca dissociating from camT_ca4_ABCD site A [camT_ca4_ABCD] → [camT_ca3_BCD] + [ca];  
 
 Transition camR to camT [camR] → [camT];  
 
 Transition camT to camR [camT] → [camR];  
 
 Transition camR_ca1_A to camT_ca1_A [camR_ca1_A] → [camT_ca1_A];  
 
 Transition camR_ca1_B to camT_ca1_B [camR_ca1_B] → [camT_ca1_B];  
 
 Transition camR_ca1_C to camT_ca1_C [camR_ca1_C] → [camT_ca1_C];  
 
 Transition camR_ca1_D to camT_ca1_D [camR_ca1_D] → [camT_ca1_D];  
 
 Transition camT_ca1_A to camR_ca1_A [camT_ca1_A] → [camR_ca1_A];  
 
 Transition camT_ca1_B to camR_ca1_B [camT_ca1_B] → [camR_ca1_B];  
 
 Transition camT_ca1_C to camR_ca1_C [camT_ca1_C] → [camR_ca1_C];  
 
 Transition camT_ca1_D to camR_ca1_D [camT_ca1_D] → [camR_ca1_D];  
 
 Transition camR_ca2_AB to camT_ca2_AB [camR_ca2_AB] → [camT_ca2_AB];  
 
 Transition camR_ca2_AC to camT_ca2_AC [camR_ca2_AC] → [camT_ca2_AC];  
 
 Transition camR_ca2_AD to camT_ca2_AD [camR_ca2_AD] → [camT_ca2_AD];  
 
 Transition camR_ca2_BC to camT_ca2_BC [camR_ca2_BC] → [camT_ca2_BC];  
 
 Transition camR_ca2_BD to camT_ca2_BD [camR_ca2_BD] → [camT_ca2_BD];  
 
 Transition camR_ca2_CD to camT_ca2_CD [camR_ca2_CD] → [camT_ca2_CD];  
 
 Transition camT_ca2_AB to camR_ca2_AB [camT_ca2_AB] → [camR_ca2_AB];  
 
 Transition camT_ca2_AC to camR_ca2_AC [camT_ca2_AC] → [camR_ca2_AC];  
 
 Transition camT_ca2_AD to camR_ca2_AD [camT_ca2_AD] → [camR_ca2_AD];  
 
 Transition camT_ca2_BC to camR_ca2_BC [camT_ca2_BC] → [camR_ca2_BC];  
 
 Transition camT_ca2_BD to camR_ca2_BD [camT_ca2_BD] → [camR_ca2_BD];  
 
 Transition camT_ca2_CD to camR_ca2_CD [camT_ca2_CD] → [camR_ca2_CD];  
 
 Transition camR_ca3_ABC to camT_ca3_ABC [camR_ca3_ABC] → [camT_ca3_ABC];  
 
 Transition camR_ca3_ABD to camT_ca3_ABD [camR_ca3_ABD] → [camT_ca3_ABD];  
 
 Transition camR_ca3_ACD to camT_ca3_ACD [camR_ca3_ACD] → [camT_ca3_ACD];  
 
 Transition camR_ca3_BCD to camT_ca3_BCD [camR_ca3_BCD] → [camT_ca3_BCD];  
 
 Transition camT_ca3_ABC to camR_ca3_ABC [camT_ca3_ABC] → [camR_ca3_ABC];  
 
 Transition camT_ca3_ABD to camR_ca3_ABD [camT_ca3_ABD] → [camR_ca3_ABD];  
 
 Transition camT_ca3_ACD to camR_ca3_ACD [camT_ca3_ACD] → [camR_ca3_ACD];  
 
 Transition camT_ca3_BCD to camR_ca3_BCD [camT_ca3_BCD] → [camR_ca3_BCD];  
 
 Transition camR_ca4_ABCD to camT_ca4_ABCD [camR_ca4_ABCD] → [camT_ca4_ABCD];  
 
 Transition camT_ca4_ABCD to camR_ca4_ABCD [camT_ca4_ABCD] → [camR_ca4_ABCD];  
 
 CaMKII binding to camR [camR] + [CaMKII] → [camR_CaMKII];  
 
 CaMKII binding to camR_ca1_A [camR_ca1_A] + [CaMKII] → [camR_ca1_A_CaMKII];  
 
 CaMKII binding to camR_ca1_B [camR_ca1_B] + [CaMKII] → [camR_ca1_B_CaMKII];  
 
 CaMKII binding to camR_ca1_C [camR_ca1_C] + [CaMKII] → [camR_ca1_C_CaMKII];  
 
 CaMKII binding to camR_ca1_D [camR_ca1_D] + [CaMKII] → [camR_ca1_D_CaMKII];  
 
 CaMKII binding to camR_ca2_AB [camR_ca2_AB] + [CaMKII] → [camR_ca2_AB_CaMKII];  
 
 CaMKII binding to camR_ca2_AC [camR_ca2_AC] + [CaMKII] → [camR_ca2_AC_CaMKII];  
 
 CaMKII binding to camR_ca2_AD [camR_ca2_AD] + [CaMKII] → [camR_ca2_AD_CaMKII];  
 
 CaMKII binding to camR_ca2_BC [camR_ca2_BC] + [CaMKII] → [camR_ca2_BC_CaMKII];  
 
 CaMKII binding to camR_ca2_BD [camR_ca2_BD] + [CaMKII] → [camR_ca2_BD_CaMKII];  
 
 CaMKII binding to camR_ca2_CD [camR_ca2_CD] + [CaMKII] → [camR_ca2_CD_CaMKII];  
 
 CaMKII binding to camR_ca3_ABC [camR_ca3_ABC] + [CaMKII] → [camR_ca3_ABC_CaMKII];  
 
 CaMKII binding to camR_ca3_ABD [camR_ca3_ABD] + [CaMKII] → [camR_ca3_ABD_CaMKII];  
 
 CaMKII binding to camR_ca3_ACD [camR_ca3_ACD] + [CaMKII] → [camR_ca3_ACD_CaMKII];  
 
 CaMKII binding to camR_ca3_BCD [camR_ca3_BCD] + [CaMKII] → [camR_ca3_BCD_CaMKII];  
 
 CaMKII binding to camR_ca4_ABCD [camR_ca4_ABCD] + [CaMKII] → [camR_ca4_ABCD_CaMKII];  
 
 CaMKII dissociation from camR [camR_CaMKII] → [camR] + [CaMKII];  
 
 CaMKII dissociation from camR_ca1_A [camR_ca1_A_CaMKII] → [camR_ca1_A] + [CaMKII];  
 
 CaMKII dissociation from camR_ca1_B [camR_ca1_B_CaMKII] → [camR_ca1_B] + [CaMKII];  
 
 CaMKII dissociation from camR_ca1_C [camR_ca1_C_CaMKII] → [camR_ca1_C] + [CaMKII];  
 
 CaMKII dissociation from camR_ca1_D [camR_ca1_D_CaMKII] → [camR_ca1_D] + [CaMKII];  
 
 CaMKII dissociation from camR_ca2_AB [camR_ca2_AB_CaMKII] → [camR_ca2_AB] + [CaMKII];  
 
 CaMKII dissociation from camR_ca2_AC [camR_ca2_AC_CaMKII] → [camR_ca2_AC] + [CaMKII];  
 
 CaMKII dissociation from camR_ca2_AD [camR_ca2_AD_CaMKII] → [camR_ca2_AD] + [CaMKII];  
 
 CaMKII dissociation from camR_ca2_BC [camR_ca2_BC_CaMKII] → [camR_ca2_BC] + [CaMKII];  
 
 CaMKII dissociation from camR_ca2_BD [camR_ca2_BD_CaMKII] → [camR_ca2_BD] + [CaMKII];  
 
 CaMKII dissociation from camR_ca2_CD [camR_ca2_CD_CaMKII] → [camR_ca2_CD] + [CaMKII];  
 
 CaMKII dissociation from camR_ca3_ABC [camR_ca3_ABC_CaMKII] → [camR_ca3_ABC] + [CaMKII];  
 
 CaMKII dissociation from camR_ca3_ABD [camR_ca3_ABD_CaMKII] → [camR_ca3_ABD] + [CaMKII];  
 
 CaMKII dissociation from camR_ca3_ACD [camR_ca3_ACD_CaMKII] → [camR_ca3_ACD] + [CaMKII];  
 
 CaMKII dissociation from camR_ca3_BCD [camR_ca3_BCD_CaMKII] → [camR_ca3_BCD] + [CaMKII];  
 
 CaMKII dissociation from camR_ca4_ABCD [camR_ca4_ABCD_CaMKII] → [camR_ca4_ABCD] + [CaMKII];  
 
 PP2B binding to camR [camR] + [PP2B] → [camR_PP2B];  
 
 PP2B binding to camR_ca1_A [camR_ca1_A] + [PP2B] → [camR_ca1_A_PP2B];  
 
 PP2B binding to camR_ca1_B [camR_ca1_B] + [PP2B] → [camR_ca1_B_PP2B];  
 
 PP2B binding to camR_ca1_C [camR_ca1_C] + [PP2B] → [camR_ca1_C_PP2B];  
 
 PP2B binding to camR_ca1_D [camR_ca1_D] + [PP2B] → [camR_ca1_D_PP2B];  
 
 PP2B binding to camR_ca2_AB [camR_ca2_AB] + [PP2B] → [camR_ca2_AB_PP2B];  
 
 PP2B binding to camR_ca2_AC [camR_ca2_AC] + [PP2B] → [camR_ca2_AC_PP2B];  
 
 PP2B binding to camR_ca2_AD [camR_ca2_AD] + [PP2B] → [camR_ca2_AD_PP2B];  
 
 PP2B binding to camR_ca2_BC [camR_ca2_BC] + [PP2B] → [camR_ca2_BC_PP2B];  
 
 PP2B binding to camR_ca2_BD [camR_ca2_BD] + [PP2B] → [camR_ca2_BD_PP2B];  
 
 PP2B binding to camR_ca2_CD [camR_ca2_CD] + [PP2B] → [camR_ca2_CD_PP2B];  
 
 PP2B binding to camR_ca3_ABC [camR_ca3_ABC] + [PP2B] → [camR_ca3_ABC_PP2B];  
 
 PP2B binding to camR_ca3_ABD [camR_ca3_ABD] + [PP2B] → [camR_ca3_ABD_PP2B];  
 
 PP2B binding to camR_ca3_ACD [camR_ca3_ACD] + [PP2B] → [camR_ca3_ACD_PP2B];  
 
 PP2B binding to camR_ca3_BCD [camR_ca3_BCD] + [PP2B] → [camR_ca3_BCD_PP2B];  
 
 PP2B binding to camR_ca4_ABCD [camR_ca4_ABCD] + [PP2B] → [camR_ca4_ABCD_PP2B];  
 
 PP2B dissociation from camR [camR_PP2B] → [camR] + [PP2B];  
 
 PP2B dissociation from camR_ca1_A [camR_ca1_A_PP2B] → [camR_ca1_A] + [PP2B];  
 
 PP2B dissociation from camR_ca1_B [camR_ca1_B_PP2B] → [camR_ca1_B] + [PP2B];  
 
 PP2B dissociation from camR_ca1_C [camR_ca1_C_PP2B] → [camR_ca1_C] + [PP2B];  
 
 PP2B dissociation from camR_ca1_D [camR_ca1_D_PP2B] → [camR_ca1_D] + [PP2B];  
 
 PP2B dissociation from camR_ca2_AB [camR_ca2_AB_PP2B] → [camR_ca2_AB] + [PP2B];  
 
 PP2B dissociation from camR_ca2_AC [camR_ca2_AC_PP2B] → [camR_ca2_AC] + [PP2B];  
 
 PP2B dissociation from camR_ca2_AD [camR_ca2_AD_PP2B] → [camR_ca2_AD] + [PP2B];  
 
 PP2B dissociation from camR_ca2_BC [camR_ca2_BC_PP2B] → [camR_ca2_BC] + [PP2B];  
 
 PP2B dissociation from camR_ca2_BD [camR_ca2_BD_PP2B] → [camR_ca2_BD] + [PP2B];  
 
 PP2B dissociation from camR_ca2_CD [camR_ca2_CD_PP2B] → [camR_ca2_CD] + [PP2B];  
 
 PP2B dissociation from camR_ca3_ABC [camR_ca3_ABC_PP2B] → [camR_ca3_ABC] + [PP2B];  
 
 PP2B dissociation from camR_ca3_ABD [camR_ca3_ABD_PP2B] → [camR_ca3_ABD] + [PP2B];  
 
 PP2B dissociation from camR_ca3_ACD [camR_ca3_ACD_PP2B] → [camR_ca3_ACD] + [PP2B];  
 
 PP2B dissociation from camR_ca3_BCD [camR_ca3_BCD_PP2B] → [camR_ca3_BCD] + [PP2B];  
 
 PP2B dissociation from camR_ca4_ABCD [camR_ca4_ABCD_PP2B] → [camR_ca4_ABCD] + [PP2B];  
 
 Ca binding to camR_CaMKII site A [camR_CaMKII] + [ca] → [camR_ca1_A_CaMKII];  
 
 Ca binding to camR_CaMKII site B [camR_CaMKII] + [ca] → [camR_ca1_B_CaMKII];  
 
 Ca binding to camR_CaMKII site C [camR_CaMKII] + [ca] → [camR_ca1_C_CaMKII];  
 
 Ca binding to camR_CaMKII site D [camR_CaMKII] + [ca] → [camR_ca1_D_CaMKII];  
 
 Ca dissociation from camR_ca1_CaMKII site A [camR_ca1_A_CaMKII] → [camR_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca1_CaMKII site C [camR_ca1_C_CaMKII] → [camR_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca1_CaMKII site D [camR_ca1_D_CaMKII] → [camR_CaMKII] + [ca];  
 
 Ca binding to camR_ca1_A_CaMKII site B [camR_ca1_A_CaMKII] + [ca] → [camR_ca2_AB_CaMKII];  
 
 Ca binding to camR_ca1_A_CaMKII site C [camR_ca1_A_CaMKII] + [ca] → [camR_ca2_AC_CaMKII];  
 
 Ca binding to camR_ca1_A_CaMKII site D [camR_ca1_A_CaMKII] + [ca] → [camR_ca2_AD_CaMKII];  
 
 Ca binding to camR_ca1_B_CaMKII site A [camR_ca1_B_CaMKII] + [ca] → [camR_ca2_AB_CaMKII];  
 
 Ca binding to camR_ca1_B_CaMKII site C [camR_ca1_B_CaMKII] + [ca] → [camR_ca2_BC_CaMKII];  
 
 Ca binding to camR_ca1_B_CaMKII site D [camR_ca1_B_CaMKII] + [ca] → [camR_ca2_BD_CaMKII];  
 
 Ca binding to camR_ca1_C_CaMKII site A [camR_ca1_C_CaMKII] + [ca] → [camR_ca2_AC_CaMKII];  
 
 Ca binding to camR_ca1_C_CaMKII site B [camR_ca1_C_CaMKII] + [ca] → [camR_ca2_BC_CaMKII];  
 
 Ca binding to camR_ca1_C_CaMKII site D [camR_ca1_C_CaMKII] + [ca] → [camR_ca2_CD_CaMKII];  
 
 Ca binding to camR_ca1_D_CaMKII site A [camR_ca1_D_CaMKII] + [ca] → [camR_ca2_AD_CaMKII];  
 
 Ca binding to camR_ca1_D_CaMKII site B [camR_ca1_D_CaMKII] + [ca] → [camR_ca2_BD_CaMKII];  
 
 Ca binding to camR_ca1_D_CaMKII site C [camR_ca1_D_CaMKII] + [ca] → [camR_ca2_CD_CaMKII];  
 
 Ca dissociation from camR_ca2_AB_CaMKII site A [camR_ca2_AB_CaMKII] → [camR_ca1_B_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca2_AB_CaMKII site B [camR_ca2_AB_CaMKII] → [camR_ca1_A_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca2_AC_CaMKII site A [camR_ca2_AC_CaMKII] → [camR_ca1_C_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca2_AC_CaMKII site C [camR_ca2_AC_CaMKII] → [camR_ca1_A_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca2_AD_CaMKII site A [camR_ca2_AD_CaMKII] → [camR_ca1_D_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca2_AD_CaMKII site D [camR_ca2_AD_CaMKII] → [camR_ca1_A_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca2_BC_CaMKII site B [camR_ca2_BC_CaMKII] → [camR_ca1_C_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca2_BC_CaMKII site C [camR_ca2_BC_CaMKII] → [camR_ca1_B_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca2_BD_CaMKII site B [camR_ca2_BD_CaMKII] → [camR_ca1_D_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca2_BD_CaMKII site D [camR_ca2_BD_CaMKII] → [camR_ca1_B_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca2_CD_CaMKII site C [camR_ca2_CD_CaMKII] → [camR_ca1_D_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca2_CD_CaMKII site D [camR_ca2_CD_CaMKII] → [camR_ca1_C_CaMKII] + [ca];  
 
 Ca binding to camR_ca2_AB_CaMKII site C [camR_ca2_AB_CaMKII] + [ca] → [camR_ca3_ABC_CaMKII];  
 
 Ca binding to camR_ca2_AB_CaMKII site D [camR_ca2_AB_CaMKII] + [ca] → [camR_ca3_ABD_CaMKII];  
 
 Ca binding to camR_ca2_AC_CaMKII site B [camR_ca2_AC_CaMKII] + [ca] → [camR_ca3_ABC_CaMKII];  
 
 Ca binding to camR_ca2_AC_CaMKII site D [camR_ca2_AC_CaMKII] + [ca] → [camR_ca3_ACD_CaMKII];  
 
 Ca binding to camR_ca2_AD_CaMKII site B [camR_ca2_AD_CaMKII] + [ca] → [camR_ca3_ABD_CaMKII];  
 
 Ca binding to camR_ca2_AD_CaMKII site C [camR_ca2_AD_CaMKII] + [ca] → [camR_ca3_ACD_CaMKII];  
 
 Ca binding to camR_ca2_BC_CaMKII site A [camR_ca2_BC_CaMKII] + [ca] → [camR_ca3_ABC_CaMKII];  
 
 Ca binding to camR_ca2_BC_CaMKII site D [camR_ca2_BC_CaMKII] + [ca] → [camR_ca3_BCD_CaMKII];  
 
 Ca binding to camR_ca2_BD_CaMKII site A [camR_ca2_BD_CaMKII] + [ca] → [camR_ca3_ABD_CaMKII];  
 
 Ca binding to camR_ca2_BD_CaMKII site C [camR_ca2_BD_CaMKII] + [ca] → [camR_ca3_BCD_CaMKII];  
 
 Ca binding to camR_ca2_CD_CaMKII site A [camR_ca2_CD_CaMKII] + [ca] → [camR_ca3_ACD_CaMKII];  
 
 Ca binding to camR_ca2_CD_CaMKII site B [camR_ca2_CD_CaMKII] + [ca] → [camR_ca3_BCD_CaMKII];  
 
 Ca dissociation from camR_ca3_ABC_CaMKII site C [camR_ca3_ABC_CaMKII] → [camR_ca2_AB_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca3_ABC_CaMKII site B [camR_ca3_ABC_CaMKII] → [camR_ca2_AC_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca3_ABC_CaMKII site A [camR_ca3_ABC_CaMKII] → [camR_ca2_BC_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca3_ABD_CaMKII site D [camR_ca3_ABD_CaMKII] → [camR_ca2_AB_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca3_ABD_CaMKII site B [camR_ca3_ABD_CaMKII] → [camR_ca2_AD_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca3_ABD_CaMKII site A [camR_ca3_ABD_CaMKII] → [camR_ca2_BD_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca3_ACD_CaMKII site D [camR_ca3_ACD_CaMKII] → [camR_ca2_AC_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca3_ACD_CaMKII site C [camR_ca3_ACD_CaMKII] → [camR_ca2_AD_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca3_ACD_CaMKII site A [camR_ca3_ACD_CaMKII] → [camR_ca2_CD_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca3_BCD_CaMKII site D [camR_ca3_BCD_CaMKII] → [camR_ca2_BC_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca3_BCD_CaMKII site C [camR_ca3_BCD_CaMKII] → [camR_ca2_BD_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca3_BCD_CaMKII site B [camR_ca3_BCD_CaMKII] → [camR_ca2_CD_CaMKII] + [ca];  
 
 Ca binding to camR_ca3_BCD_CaMKII site A [camR_ca3_BCD_CaMKII] + [ca] → [camR_ca4_ABCD_CaMKII];  
 
 Ca binding to camR_ca3_ACD_CaMKII site B [camR_ca3_ACD_CaMKII] + [ca] → [camR_ca4_ABCD_CaMKII];  
 
 Ca binding to camR_ca3_ABD_CaMKII site C [camR_ca3_ABD_CaMKII] + [ca] → [camR_ca4_ABCD_CaMKII];  
 
 Ca binding to camR_ca3_ABC_CaMKII site D [camR_ca3_ABC_CaMKII] + [ca] → [camR_ca4_ABCD_CaMKII];  
 
 Ca dissociation from camR_ca4_ABCD_CaMKII site A [camR_ca4_ABCD_CaMKII] → [camR_ca3_BCD_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca4_ABCD_CaMKII site B [camR_ca4_ABCD_CaMKII] → [camR_ca3_ACD_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca4_ABCD_CaMKII site C [camR_ca4_ABCD_CaMKII] → [camR_ca3_ABD_CaMKII] + [ca];  
 
 Ca dissociation from camR_ca4_ABCD_CaMKII site D [camR_ca4_ABCD_CaMKII] → [camR_ca3_ABC_CaMKII] + [ca];  
 
 Ca binding to camR_PP2B site A [camR_PP2B] + [ca] → [camR_ca1_A_PP2B];  
 
 Ca binding to camR_PP2B site B [camR_PP2B] + [ca] → [camR_ca1_B_PP2B];  
 
 Ca binding to camR_PP2B site C [camR_PP2B] + [ca] → [camR_ca1_C_PP2B];  
 
 Ca binding to camR_PP2B site D [camR_PP2B] + [ca] → [camR_ca1_D_PP2B];  
 
 Ca dissociation from camR_ca1_A_PP2B site A [camR_ca1_A_PP2B] → [camR_PP2B] + [ca];  
 
 Ca dissociation from camR_ca1_B_PP2B site B [camR_ca1_B_PP2B] → [camR_PP2B] + [ca];  
 
 Ca dissociation from camR_ca1_C_PP2B site C [camR_ca1_C_PP2B] → [camR_PP2B] + [ca];  
 
 Ca dissociation from camR_ca1_D_PP2B site D [camR_ca1_D_PP2B] → [camR_PP2B] + [ca];  
 
 Ca binding to camR_ca1_A_PP2B site B [camR_ca1_A_PP2B] + [ca] → [camR_ca2_AB_PP2B];  
 
 Ca binding to camR_ca1_A_PP2B site C [camR_ca1_A_PP2B] + [ca] → [camR_ca2_AC_PP2B];  
 
 Ca binding to camR_ca1_A_PP2B site D [camR_ca1_A_PP2B] + [ca] → [camR_ca2_AD_PP2B];  
 
 Ca binding to camR_ca1_B_PP2B site A [camR_ca1_B_PP2B] + [ca] → [camR_ca2_AB_PP2B];  
 
 Ca binding to camR_ca1_B_PP2B site C [camR_ca1_B_PP2B] + [ca] → [camR_ca2_BC_PP2B];  
 
 Ca binding to camR_ca1_B_PP2B site D [camR_ca1_B_PP2B] + [ca] → [camR_ca2_BD_PP2B];  
 
 Ca binding to camR_ca1_C_PP2B site A [camR_ca1_C_PP2B] + [ca] → [camR_ca2_AC_PP2B];  
 
 Ca binding to camR_ca1_C_PP2B site B [camR_ca1_C_PP2B] + [ca] → [camR_ca2_BC_PP2B];  
 
 Ca binding to camR_ca1_C_PP2B site D [camR_ca1_C_PP2B] + [ca] → [camR_ca2_CD_PP2B];  
 
 Ca binding to camR_ca1_D_PP2B site A [camR_ca1_D_PP2B] + [ca] → [camR_ca2_AD_PP2B];  
 
 Ca binding to camR_ca1_D_PP2B site B [camR_ca1_D_PP2B] + [ca] → [camR_ca2_BD_PP2B];  
 
 Ca binding to camR_ca1_D_PP2B site C [camR_ca1_D_PP2B] + [ca] → [camR_ca2_CD_PP2B];  
 
 Ca dissociating from camR_ca2_AB_PP2B site A [camR_ca2_AB_PP2B] → [camR_ca1_B_PP2B] + [ca];  
 
 Ca dissociating from camR_ca2_AB_PP2B site B [camR_ca2_AB_PP2B] → [camR_ca1_A_PP2B] + [ca];  
 
 Ca dissociating from camR_ca2_AC_PP2B site A [camR_ca2_AC_PP2B] → [camR_ca1_C_PP2B] + [ca];  
 
 Ca dissociating from camR_ca2_AC_PP2B site C [camR_ca2_AC_PP2B] → [camR_ca1_A_PP2B] + [ca];  
 
 Ca dissociating from camR_ca2_AD_PP2B site A [camR_ca2_AD_PP2B] → [camR_ca1_D_PP2B] + [ca];  
 
 Ca dissociating from camR_ca2_AD_PP2B site D [camR_ca2_AD_PP2B] → [camR_ca1_A_PP2B] + [ca];  
 
 Ca dissociating from camR_ca2_BC_PP2B site B [camR_ca2_BC_PP2B] → [camR_ca1_C_PP2B] + [ca];  
 
 Ca dissociating from camR_ca2_BC_PP2B site C [camR_ca2_BC_PP2B] → [camR_ca1_B_PP2B] + [ca];  
 
 Ca dissociating from camR_ca2_BD_PP2B site B [camR_ca2_BD_PP2B] → [camR_ca1_D_PP2B] + [ca];  
 
 Ca dissociating from camR_ca2_BD_PP2B site D [camR_ca2_BD_PP2B] → [camR_ca1_B_PP2B] + [ca];  
 
 Ca dissociating from camR_ca2_CD_PP2B site C [camR_ca2_CD_PP2B] → [camR_ca1_D_PP2B] + [ca];  
 
 Ca dissociating from camR_ca2_CD_PP2B site D [camR_ca2_CD_PP2B] → [camR_ca1_C_PP2B] + [ca];  
 
 Ca binding to camR_ca2_AB_PP2B site C [camR_ca2_AB_PP2B] + [ca] → [camR_ca3_ABC_PP2B];  
 
 Ca binding to camR_ca2_AB_PP2B site D [camR_ca2_AB_PP2B] + [ca] → [camR_ca3_ABD_PP2B];  
 
 Ca binding to camR_ca2_AC_PP2B site B [camR_ca2_AC_PP2B] + [ca] → [camR_ca3_ABC_PP2B];  
 
 Ca binding to camR_ca2_AC_PP2B site D [camR_ca2_AC_PP2B] + [ca] → [camR_ca3_ACD_PP2B];  
 
 Ca binding to camR_ca2_AD_PP2B site B [camR_ca2_AD_PP2B] + [ca] → [camR_ca3_ABD_PP2B];  
 
 Ca binding to camR_ca2_AD_PP2B site C [camR_ca2_AD_PP2B] + [ca] → [camR_ca3_ACD_PP2B];  
 
 Ca binding to camR_ca2_BC_PP2B site A [camR_ca2_BC_PP2B] + [ca] → [camR_ca3_ABC_PP2B];  
 
 Ca binding to camR_ca2_BC_PP2B site D [camR_ca2_BC_PP2B] + [ca] → [camR_ca3_BCD_PP2B];  
 
 Ca binding to camR_ca2_BD_PP2B site A [camR_ca2_BD_PP2B] + [ca] → [camR_ca3_ABD_PP2B];  
 
 Ca binding to camR_ca2_BD_PP2B site C [camR_ca2_BD_PP2B] + [ca] → [camR_ca3_BCD_PP2B];  
 
 Ca binding to camR_ca2_CD_PP2B site A [camR_ca2_CD_PP2B] + [ca] → [camR_ca3_ACD_PP2B];  
 
 Ca binding to camR_ca2_CD_PP2B site B [camR_ca2_CD_PP2B] + [ca] → [camR_ca3_BCD_PP2B];  
 
 Ca dissociation from camR_ca3_ABC_PP2B site A [camR_ca3_ABC_PP2B] → [camR_ca2_BC_PP2B] + [ca];  
 
 Ca dissociation from camR_ca3_ABC_PP2B site B [camR_ca3_ABC_PP2B] → [camR_ca2_AC_PP2B] + [ca];  
 
 Ca dissociation from camR_ca3_ABC_PP2B site C [camR_ca3_ABC_PP2B] → [camR_ca2_AB_PP2B] + [ca];  
 
 Ca dissociation from camR_ca3_ABD_PP2B site A [camR_ca3_ABD_PP2B] → [camR_ca2_BD_PP2B] + [ca];  
 
 Ca dissociation from camR_ca3_ABD_PP2B site B [camR_ca3_ABD_PP2B] → [camR_ca2_AD_PP2B] + [ca];  
 
 Ca dissociation from camR_ca3_ABD_PP2B site D [camR_ca3_ABD_PP2B] → [camR_ca2_AB_PP2B] + [ca];  
 
 Ca dissociation from camR_ca3_ACD_PP2B site A [camR_ca3_ACD_PP2B] → [camR_ca2_CD_PP2B] + [ca];  
 
 Ca dissociation from camR_ca3_ACD_PP2B site C [camR_ca3_ACD_PP2B] → [camR_ca2_AD_PP2B] + [ca];  
 
 Ca dissociation from camR_ca3_ACD_PP2B site D [camR_ca3_ACD_PP2B] → [camR_ca2_AC_PP2B] + [ca];  
 
 Ca dissociation from camR_ca3_BCD_PP2B site B [camR_ca3_BCD_PP2B] → [camR_ca2_CD_PP2B] + [ca];  
 
 Ca dissociation from camR_ca3_BCD_PP2B site C [camR_ca3_BCD_PP2B] → [camR_ca2_BD_PP2B] + [ca];  
 
 Ca dissociation from camR_ca3_BCD_PP2B site D [camR_ca3_BCD_PP2B] → [camR_ca2_BC_PP2B] + [ca];  
 
 Ca binding to camR_ca3_ABC_PP2B site D [camR_ca3_ABC_PP2B] + [ca] → [camR_ca4_ABCD_PP2B];  
 
 Ca binding to camR_ca3_ABD_PP2B site C [camR_ca3_ABD_PP2B] + [ca] → [camR_ca4_ABCD_PP2B];  
 
 Ca binding to camR_ca3_ACD_PP2B site B [camR_ca3_ACD_PP2B] + [ca] → [camR_ca4_ABCD_PP2B];  
 
 Ca binding to camR_ca3_BCD_PP2B site A [camR_ca3_BCD_PP2B] + [ca] → [camR_ca4_ABCD_PP2B];  
 
 Ca dissociating from camR_ca4_ABCD_PP2B site A [camR_ca4_ABCD_PP2B] → [camR_ca3_BCD_PP2B] + [ca];  
 
 Ca dissociating from camR_ca4_ABCD_PP2B site B [camR_ca4_ABCD_PP2B] → [camR_ca3_ACD_PP2B] + [ca];  
 
 Ca dissociating from camR_ca4_ABCD_PP2B site C [camR_ca4_ABCD_PP2B] → [camR_ca3_ABD_PP2B] + [ca];  
 
 Ca dissociating from camR_ca4_ABCD_PP2B site D [camR_ca4_ABCD_PP2B] → [camR_ca3_ABC_PP2B] + [ca];  
 
 Ca dissociation from camR_ca1_CaMKII site B [camR_ca1_B_CaMKII] → [camR_CaMKII] + [ca];  
 
 Ca dissociating from camT_ca3_ABC site C [camT_ca3_ABC] → [camT_ca2_AB] + [ca];  
 
Rules (46)
 
 Assignment Rule (name: parameter_1) koffRA = parameter_50*parameter_0
 
 Assignment Rule (name: parameter_20) free_camT_ca3_total = species_28+species_29+species_30+species_31
 
 Assignment Rule (name: parameter_2) koffRB = parameter_51*parameter_0
 
 Assignment Rule (name: parameter_3) koffRC = parameter_52*parameter_0
 
 Assignment Rule (name: parameter_4) koffRD = parameter_53*parameter_0
 
 Assignment Rule (name: parameter_5) koffTA = parameter_1/parameter_11
 
 Assignment Rule (name: parameter_6) koffTB = parameter_2/parameter_12
 
 Assignment Rule (name: parameter_7) koffTC = parameter_3/parameter_13
 
 Assignment Rule (name: parameter_8) koffTD = parameter_4/parameter_14
 
 Assignment Rule (name: parameter_10) kTR = parameter_9/parameter_49
 
 Assignment Rule (name: parameter_19) free_camR_ca3_total = species_12+species_13+species_14+species_15
 
 Assignment Rule (name: parameter_21) free_cam_ca3_total = parameter_19+parameter_20
 
 Assignment Rule (name: parameter_22) cam_ca4_total = species_16+species_32+species_49+species_66
 
 Assignment Rule (name: parameter_23) CaMKII_camR_ca1 = species_35+species_36+species_37+species_38
 
 Assignment Rule (name: parameter_25) CaMKII_camR_ca3 = species_45+species_46+species_47+species_48
 
 Assignment Rule (name: parameter_24) CaMKII_camR_ca2 = species_39+species_40+species_41+species_42+species_43+species_44
 
 Assignment Rule (name: parameter_26) total_CaMKII_bound = species_34+parameter_23+parameter_24+parameter_25+species_49
 
 Assignment Rule (name: parameter_27) total CaMKII = parameter_26+species_33
 
 Assignment Rule (name: parameter_28) CaMKIIbar = parameter_26/parameter_27
 
 Assignment Rule (name: parameter_29) PP2B_camR_ca1 = species_52+species_53+species_54+species_55
 
 Assignment Rule (name: parameter_30) PP2B_camR_ca2 = species_56+species_57+species_58+species_59+species_60+species_61
 
 Assignment Rule (name: parameter_31) PP2B_camR_ca3 = species_62+species_63+species_64+species_65
 
 Assignment Rule (name: parameter_32) total_PP2B_bound = species_51+parameter_29+parameter_30+parameter_31+species_66
 
 Assignment Rule (name: parameter_35) camR_unbound = species_0+species_2+species_3+species_4+species_5+species_6+species_7+species_8+species_9+species_10+species_11+species_12+species_13+species_14+species_15+species_16
 
 Assignment Rule (name: parameter_33) total PP2B = parameter_32+species_50
 
 Assignment Rule (name: parameter_34) PP2Bbar = parameter_32/parameter_33
 
 Assignment Rule (name: parameter_37) total camT = species_17+species_18+species_19+species_20+species_21+species_22+species_23+species_24+species_25+species_26+species_27+species_28+species_29+species_30+species_31+species_32
 
 Assignment Rule (name: parameter_36) total camR = parameter_35+parameter_26+parameter_32
 
 Assignment Rule (name: parameter_39) cam_ca3_total = parameter_25+parameter_31+parameter_21
 
 Assignment Rule (name: parameter_38) Rbar = parameter_36/(parameter_36+parameter_37)
 
 Assignment Rule (name: parameter_40) free_camR_ca2_total = species_6+species_7+species_8+species_9+species_10+species_11
 
 Assignment Rule (name: parameter_41) free_camT_ca2_total = species_22+species_23+species_24+species_25+species_26+species_27
 
 Assignment Rule (name: parameter_42) free_cam_ca2_total = parameter_40+parameter_41
 
 Assignment Rule (name: parameter_43) cam_ca2_total = parameter_42+parameter_24+parameter_30
 
 Assignment Rule (name: parameter_44) free_cam_ca1_total = species_2+species_3+species_4+species_5+species_18+species_19+species_20+species_21
 
 Assignment Rule (name: parameter_45) cam_ca1_total = parameter_44+parameter_23+parameter_29
 
 Assignment Rule (name: parameter_46) cam_ca0_total = species_0+species_17+species_34+species_51
 
 Assignment Rule (name: parameter_47) cam_total = parameter_46+parameter_45+parameter_43+parameter_39+parameter_22
 
 Assignment Rule (name: parameter_48) moles_bound_ca_per_moles_cam = (4*parameter_22+3*parameter_39+2*parameter_43+parameter_45)/parameter_47
 
 Assignment Rule (name: parameter_54) alpha = species_16/(species_16+species_32)
 
 Assignment Rule (name: parameter_55) beta = parameter_19/parameter_21
 
 Assignment Rule (name: parameter_56) gamma = parameter_40/parameter_42
 
 Assignment Rule (name: parameter_57) delta = (species_2+species_3+species_4+species_5)/(species_2+species_3+species_4+species_5+species_18+species_19+species_20+species_21)
 
 Assignment Rule (name: parameter_58) epsilon = species_0/(species_0+species_17)
 
 Assignment Rule (name: parameter_59) ybar = parameter_48/4
 
 Assignment Rule (name: parameter_60) ybar/(1-ybar) = parameter_59/(1-parameter_59)
 
Functions (27)
 
 Transition camR_ca1_X to camT_ca1_X lambda(ka, b, species, species*ka*b^(1/2))
 
 function_4_Transition camT_ca1_A to camR_ca1_A lambda(parameter_10, parameter_11, species_18, species_18*parameter_10/parameter_11^(1/2))
 
 function_4_Transition camT_ca1_B to camR_ca1_B lambda(parameter_10, parameter_12, species_19, species_19*parameter_10/parameter_12^(1/2))
 
 function_4_Transition camR_ca2_AB to camT_ca2_AB lambda(parameter_11, parameter_12, parameter_9, species_6, species_6*parameter_9*(parameter_11*parameter_12)^(1/2))
 
 function_4_Transition camT_ca1_C to camR_ca1_C lambda(parameter_10, parameter_13, species_20, species_20*parameter_10/parameter_13^(1/2))
 
 function_4_Transition camR_ca2_AC to camT_ca2_AC lambda(parameter_11, parameter_13, parameter_9, species_7, species_7*parameter_9*(parameter_11*parameter_13)^(1/2))
 
 function_4_Transition camR_ca2_AD to camT_ca2_AD lambda(parameter_11, parameter_14, parameter_9, species_8, species_8*parameter_9*(parameter_11*parameter_14)^(1/2))
 
 function_4_Transition camT_ca1_D to camR_ca1_D lambda(parameter_10, parameter_14, species_21, species_21*parameter_10/parameter_14^(1/2))
 
 function_4_Transition camT_ca2_AD to camR_ca2_AD lambda(parameter_10, parameter_11, parameter_14, species_24, species_24*parameter_10/(parameter_11*parameter_14)^(1/2))
 
 function_4_Transition camT_ca2_BC to camR_ca2_BC lambda(parameter_10, parameter_12, parameter_13, species_25, species_25*parameter_10/(parameter_12*parameter_13)^(1/2))
 
 function_4_Transition camT_ca2_BD to camR_ca2_BD lambda(parameter_10, parameter_12, parameter_14, species_26, species_26*parameter_10/(parameter_12*parameter_14)^(1/2))
 
 function_4_Transition camT_ca2_CD to camR_ca2_CD lambda(parameter_10, parameter_13, parameter_14, species_27, species_27*parameter_10/(parameter_13*parameter_14)^(1/2))
 
 function_4_Transition camR_ca3_ABC to camT_ca3_ABC lambda(parameter_11, parameter_12, parameter_13, parameter_9, species_12, species_12*parameter_9*(parameter_11*parameter_12*parameter_13)^(1/2))
 
 function_4_Transition camR_ca3_ABD to camT_ca3_ABD lambda(parameter_11, parameter_12, parameter_14, parameter_9, species_13, species_13*parameter_9*(parameter_11*parameter_12*parameter_14)^(1/2))
 
 function_4_Transition camR_ca3_ACD to camT_ca3_ACD lambda(parameter_11, parameter_13, parameter_14, parameter_9, species_14, species_14*parameter_9*(parameter_11*parameter_13*parameter_14)^(1/2))
 
 function_4_Transition camR_ca3_BCD to camT_ca3_BCD lambda(parameter_12, parameter_13, parameter_14, parameter_9, species_15, species_15*parameter_9*(parameter_12*parameter_13*parameter_14)^(1/2))
 
 function_4_Transition camR_ca2_BC to camT_ca2_BC lambda(parameter_12, parameter_13, parameter_9, species_9, species_9*parameter_9*(parameter_12*parameter_13)^(1/2))
 
 function_4_Transition camR_ca2_BD to camT_ca2_BD lambda(parameter_12, parameter_14, parameter_9, species_10, species_10*parameter_9*(parameter_12*parameter_14)^(1/2))
 
 function_4_Transition camR_ca2_CD to camT_ca2_CD lambda(parameter_13, parameter_14, parameter_9, species_11, species_11*parameter_9*(parameter_13*parameter_14)^(1/2))
 
 function_4_Transition camT_ca2_AB to camR_ca2_AB lambda(parameter_10, parameter_11, parameter_12, species_22, species_22*parameter_10/(parameter_11*parameter_12)^(1/2))
 
 function_4_Transition camT_ca2_AC to camR_ca2_AC lambda(parameter_10, parameter_11, parameter_13, species_23, species_23*parameter_10/(parameter_11*parameter_13)^(1/2))
 
 function_4_Transition camT_ca3_ABC to camR_ca3_ABC lambda(parameter_10, parameter_11, parameter_12, parameter_13, species_28, species_28*parameter_10/(parameter_11*parameter_12*parameter_13)^(1/2))
 
 function_4_Transition camT_ca3_ABD to camR_ca3_ABD lambda(parameter_10, parameter_11, parameter_12, parameter_14, species_29, species_29*parameter_10/(parameter_11*parameter_12*parameter_14)^(1/2))
 
 function_4_Transition camT_ca3_ACD to camR_ca3_ACD lambda(parameter_10, parameter_11, parameter_13, parameter_14, species_30, species_30*parameter_10/(parameter_11*parameter_13*parameter_14)^(1/2))
 
 function_4_Transition camT_ca3_BCD to camR_ca3_BCD lambda(parameter_10, parameter_12, parameter_13, parameter_14, species_31, species_31*parameter_10/(parameter_12*parameter_13*parameter_14)^(1/2))
 
 function_4_Transition camR_ca4_ABCD to camT_ca4_ABCD lambda(parameter_11, parameter_12, parameter_13, parameter_14, parameter_9, species_16, species_16*parameter_9*(parameter_11*parameter_12*parameter_13*parameter_14)^(1/2))
 
 function_4_Transition camT_ca4_ABCD to camR_ca4_ABCD lambda(parameter_10, parameter_11, parameter_12, parameter_13, parameter_14, species_32, species_32*parameter_10/(parameter_11*parameter_12*parameter_13*parameter_14)^(1/2))
 
 Spine Spatial dimensions: 3.0  Compartment size: 1.0E-15
 
 camR
Compartment: Spine
Initial concentration: 9.7E-12
 
 ca
Compartment: Spine
Initial concentration: 1.0E-5
 
 camR_ca1_A
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca1_B
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca1_C
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca1_D
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca2_AB
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca2_AC
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca2_AD
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca2_BC
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca2_BD
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca2_CD
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca3_ABC
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca3_ABD
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca3_ACD
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca3_BCD
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca4_ABCD
Compartment: Spine
Initial concentration: 0.0
 
 camT
Compartment: Spine
Initial concentration: 2.0E-7
 
 camT_ca1_A
Compartment: Spine
Initial concentration: 0.0
 
 camT_ca1_B
Compartment: Spine
Initial concentration: 0.0
 
 camT_ca1_C
Compartment: Spine
Initial concentration: 0.0
 
 camT_ca1_D
Compartment: Spine
Initial concentration: 0.0
 
 camT_ca2_AB
Compartment: Spine
Initial concentration: 0.0
 
 camT_ca2_AC
Compartment: Spine
Initial concentration: 0.0
 
 camT_ca2_AD
Compartment: Spine
Initial concentration: 0.0
 
 camT_ca2_BC
Compartment: Spine
Initial concentration: 0.0
 
 camT_ca2_BD
Compartment: Spine
Initial concentration: 0.0
 
 camT_ca2_CD
Compartment: Spine
Initial concentration: 0.0
 
 camT_ca3_ABC
Compartment: Spine
Initial concentration: 0.0
 
 camT_ca3_ABD
Compartment: Spine
Initial concentration: 0.0
 
 camT_ca3_ACD
Compartment: Spine
Initial concentration: 0.0
 
 camT_ca3_BCD
Compartment: Spine
Initial concentration: 0.0
 
 camT_ca4_ABCD
Compartment: Spine
Initial concentration: 0.0
 
 CaMKII
Compartment: Spine
Initial concentration: 7.0E-5
 
 camR_CaMKII
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca1_A_CaMKII
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca1_B_CaMKII
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca1_C_CaMKII
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca1_D_CaMKII
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca2_AB_CaMKII
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca2_AC_CaMKII
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca2_AD_CaMKII
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca2_BC_CaMKII
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca2_BD_CaMKII
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca2_CD_CaMKII
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca3_ABC_CaMKII
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca3_ABD_CaMKII
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca3_ACD_CaMKII
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca3_BCD_CaMKII
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca4_ABCD_CaMKII
Compartment: Spine
Initial concentration: 0.0
 
 PP2B
Compartment: Spine
Initial concentration: 1.6E-6
 
 camR_PP2B
Compartment: Spine
Initial concentration: 0.0
 
 camR_ca1_A_PP2B
Compartment: Spine
Initial concentration: 0.0