BIOMD0000000429
type: SBMLModel
parameter: COPASI27
parameter: COPASI37
compartment: 170196
species: COPASI6
parameter: 170293
parameter: COPASI68
notes: Schaber2012 - Hog pathway in yeast The high osmolarity glycerol (HOG) pathway in the yeast Saccharomyces cerevisiae is one of the best-studied mitogen-activated protein kinase (MAPK) pathways and serves as a prototype signalling system for eukaryotes. This pathway is necessary and sufficient to adapt to high external osmolarity. A key component of this pathway is the stress-activated protein kinase (SAPK) Hog1, which is rapidly phosphorylated by the SAPK kinase Pbs2 upon hyper-osmotic shock, and which is the terminal kinase of two parallel signalling pathways, subsequently called the Sho1 branch and the Sln1 branch, respectively. Ensemble modelling (192 models) is used to study the yeast HOG pathway, a prototype for eukaryotic mitogen-activated kinase signalling systems. The best fit model (Model Nr.22: described here) provides new insights into the function of this system, some of which are then experimentally validated. This model is described in the article: Modelling reveals novel roles of two parallel signalling pathways and homeostatic feedbacks in yeast. Schaber J, Baltanas R, Bush A, Klipp E, Colman-Lerner A. Mol Syst Biol. 2012 Nov 13;8:622. Abstract: The high osmolarity glycerol (HOG) pathway in yeast serves as a prototype signalling system for eukaryotes. We used an unprecedented amount of data to parameterise 192 models capturing different hypotheses about molecular mechanisms underlying osmo-adaptation and selected a best approximating model. This model implied novel mechanisms regulating osmo-adaptation in yeast. The model suggested that (i) the main mechanism for osmo-adaptation is a fast and transient non-transcriptional Hog1-mediated activation of glycerol production, (ii) the transcriptional response serves to maintain an increased steady-state glycerol production with low steady-state Hog1 activity, and (iii) fast negative feedbacks of activated Hog1 on upstream signalling branches serves to stabilise adaptation response. The best approximating mo…
rule: 171001
rule: 171145
parameter: 170281
parameter: COPASI56
parameter: 170413
rule: 171061
parameter: COPASI61
rule: 171157
parameter: 170341
species: COPASI15
occurs in: http://identifiers.org/taxonomy/4932
parameter: 170401
name: mod. MA OR (irrev)
rule: 171073
parameter: COPASI55
parameter: 170269
parameter: COPASI65
reaction: COPASI100
parameter: COPASI60
reaction: COPASI89
parameter: 170437
rule: 171085
parameter: COPASI85
parameter: COPASI59
function def: COPASI105
function def: COPASI104
species: COPASI13
rule: 171097
parameter: 170425
compartment: COPASI3
parameter: COPASI21
parameter: COPASI79
parameter: COPASI64
reaction: COPASI88
parameter: COPASI84
function def: COPASI103
reaction: COPASI92
parameter: COPASI52
name: 2 param 2 S MA with inhibition (irrev)
parameter: COPASI25
function def: 170160
species: COPASI5
compartment: COPASI2
parameter: COPASI20
area units: 844749
parameter: COPASI72
species: COPASI9
parameter: COPASI45
name: 2 param mod. add. constant flux MM (irrev)
reaction: COPASI96
function def: COPASI108
rule: 170905
function def: 170136
reaction: COPASI91
parameter: COPASI51
parameter: COPASI24
parameter: COPASI76
units def: 170172
parameter: COPASI49
rule: 170917
parameter: COPASI71
parameter: COPASI44
function def: COPASI107
reaction: COPASI95
rule: 170929
rule: 170977
parameter: COPASI32
parameter: COPASI75
parameter: 170329
species: COPASI18
parameter: COPASI48
rule: 171121

COPASI27
type: Parameter
notes: Membrane rigidity (Schaber, et al., 2010)
constant: true
value: 14.3
name: eps

COPASI37
type: Parameter
notes: Initial total cellular osmolyte concentration.
constant: true
value: 502026.122077738
name: ci_0

170196
type: Compartment
is version of: GO:0005886
constant: true
size: 1.0
spatial dimensions: 3.0
name: M

COPASI6
type: Species
initial amount: 9.6760009944572
initial concentration: NaN
constant: false
is encoded by: S000004103
notes: MAP kinase
has only substance units: true
name: Hog1
is version of: P32485
in compartment: COPASI2
boundary condition: false

COPASI2
type: Compartment
notes: Osmolytically active volume, derived from a total cell volume of 50 fl and a solid base volume of 41%.
is version of: GO:0005623
constant: false
size: 29.5
spatial dimensions: 3.0
name: Vos

170293
type: Parameter
constant: true
value: 0.075474
name: v1_fb_k

COPASI68
type: Parameter
constant: true
value: 1800.0
name: Glyex_0

170281
type: Parameter
constant: true
value: 0.345701
name: v1_fb_h

COPASI56
type: Parameter
notes: 〖2160f〗_n2μM: molecule numbers from http://yeastgfp.yeastgenome.org/
constant: true
value: 0.12158807536209
name: Pbs2Total

170413
type: Parameter
constant: true
value: 0.628719
name: v16_6_h

COPASI61
type: Parameter
constant: true
value: 2.23
name: Hog1PPrelIniSho1

170341
type: Parameter
constant: true
value: 0.00226722
name: v4_k1

COPASI15
type: Species
in compartment: COPASI2
initial amount: 0.00342460653415448
has part: P40073
constant: false
has part: P08018
initial concentration: NaN
has only substance units: true
name: Sho1Pbs2P
notes: Active scaffold complex (Sho1 branch). Initial condition set such that a steady state of Hog1 phosphorylation is maintained.
boundary condition: false

170401
type: Parameter
constant: true
value: 0.0811033
name: v16_6_Ki

COPASI55
type: Parameter
notes: 〖6788f〗_n2μM: molecule numbers from http://yeastgfp.yeastgenome.org/
constant: true
value: 0.382101784980495
name: Hog1Total

170269
type: Parameter
constant: true
value: 0.00940584
name: v1_fb_Ki

COPASI65
type: Parameter
constant: true
value: 65.6342903668733
name: A_0

COPASI100
type: Reaction
reversible: false
modifier: 688591
is version of: GO:0006461
name: v3_fb
modifier: 688603
reactant: 171745
modifier: 688579
kinetic law: 171781
notes: Simple cell surface area dependent binding of Pbs2 to Sho1. The complex is supposed to be the active form. With transient inhibition by activated Hog1.
modifier: 171769
fast: false
reactant: 171733
product: 171757

COPASI14
type: Species
constant: false
boundary condition: false
in compartment: COPASI2
initial concentration: NaN
initial amount: 3.86572185643496
notes: One of the putative upstream sensors
is encoded by: S000000920
is version of: P40073
name: Sho1
has only substance units: true

170208
type: Species
is version of: P32485
constant: false
boundary condition: true
has only substance units: true
initial concentration: NaN
initial amount: 0.201092527399535
in compartment: COPASI2
name: Hog1PPActive

171745
type: SpeciesReference
stoichiometry: 1.0
species: COPASI14

COPASI8
type: Species
initial amount: 3.57956846222666
notes: MAP kinase kinase
has only substance units: true
initial concentration: NaN
name: Pbs2
constant: false
boundary condition: false
encodes: S000003664
in compartment: COPASI2

171733
type: SpeciesReference
stoichiometry: 1.0
species: COPASI8

171757
type: SpeciesReference
stoichiometry: 1.0
species: COPASI15

COPASI60
type: Parameter
constant: true
value: 2.23
name: Hog1PPrelIniSln1

COPASI89
type: Reaction
notes: Linear phosphorylation of Hog1 by either the scaffold complex (Sho1-branch) or activated Pbs2 (Sln1-branch).
modifier: 171301
modifier: 688375
modifier: 688363
fast: false
modifier: 688351
reversible: false
is version of: GO:0016310
reactant: 171265
product: 171277
kinetic law: 171313
name: v5
modifier: 171289

COPASI9
type: Species
boundary condition: false
in compartment: COPASI2
constant: false
initial concentration: NaN
initial amount: 0.00385515442085361
is version of: P08018
name: Pbs2P
has only substance units: true
notes: Activated MAP kinase kinase (Sln1 branch). Initial condition set such that a steady state of Hog1 phosphorylation is maintained.

171265
type: SpeciesReference
stoichiometry: 1.0
species: COPASI6

171277
type: SpeciesReference
stoichiometry: 1.0
species: COPASI13

COPASI13
type: Species
has only substance units: true
is version of: P32485
boundary condition: false
notes: Single phosphorylated MAP kinase. Initial condition set such that a steady state of Hog1 phosphorylation is maintained.
initial concentration: NaN
is version of: C00562
initial amount: 1.39490913506788
name: Hog1P
in compartment: COPASI2
constant: false

170437
type: Parameter
constant: true
value: 46.8363
name: v13_k2

COPASI85
type: Parameter
constant: true
value: 0.0
name: Inhibition

COPASI59
type: Parameter
constant: true
value: 48.0003902091319
name: kHog1phos2

170425
type: Parameter
constant: true
value: 680.818
name: v13_k1

COPASI3
type: Compartment
is version of: GO:0005576
constant: true
size: 50000.0
spatial dimensions: 3.0
name: Vex

COPASI21
type: Parameter
notes: Mole number
constant: true
value: 6.022E23
name: mol

COPASI79
type: Parameter
constant: false
value: 0.499999999999999
name: TurgorActivation

COPASI64
type: Parameter
notes: Initial osmotically active volume.
constant: true
value: 29.5
name: Vos_0

COPASI88
type: Reaction
kinetic law: 171253
reversible: false
product: 171229
modifier: 688339
name: v2
modifier: 688327
notes: Constitutive phosphatase dependent deactivation.
fast: false
is version of: GO:0016311
reactant: 171217
modifier: 171241

171229
type: SpeciesReference
stoichiometry: 1.0
species: COPASI8

COPASI10
type: Species
notes: Placeholder for phosphatases like Ppt1/2/3 and others. Initial value derived from measured number of Ptp3 molecules (769, http://yeastgfp.yeastgenome.org)
initial concentration: NaN
in compartment: COPASI2
has only substance units: true
constant: false
boundary condition: false
is part of: P40048
initial amount: 1.27698439056792
name: Phosphatase

171217
type: SpeciesReference
stoichiometry: 1.0
species: COPASI9

COPASI84
type: Parameter
constant: false
value: 1.0
name: Hog1Inhibition

COPASI92
type: Reaction
kinetic law: 171421
modifier: 688423
reactant: 171409
notes: Protein/enzyme degradation
is version of: GO:0030163
fast: false
reversible: false
name: v12

COPASI11
type: Species
name: Protein
in compartment: COPASI2
constant: false
notes: Placeholder for Hog1-dependent proteins, especially Gpd1. Initial value derived from measured number of Gpd1 molecules (807, http://yeastgfp.yeastgenome.org)
initial concentration: NaN
initial amount: 1.34008635004982
has only substance units: true
boundary condition: false
is version of: CHEBI:36080

171409
type: SpeciesReference
stoichiometry: 1.0
species: COPASI11

COPASI52
type: Parameter
constant: true
value: 0.128378897200866
name: Fps1ClosureRate

COPASI25
type: Parameter
notes: Hydraulic conductivity (estimate from data from (Eriksson, et al., 2007)).
constant: true
value: 0.013
name: Lp

COPASI5
type: Species
has only substance units: true
constant: false
is version of: CHEBI:17754
name: Glyin
boundary condition: false
initial amount: 5310000.0
in compartment: COPASI2
notes: Intracellular glycerol, approximated by assuming a measured value of 0.1 mM/OD in 1 ml sample (Klipp, et al., 2005) and assuming 18•106 cells per ml sample culture and an average osmotic cell volume of 29.5 fl, i.e. 1/18/29.5•108.
initial concentration: NaN

COPASI20
type: Parameter
notes: Temperature, corresponds to 30oC.
constant: true
value: 303.15
name: T

COPASI72
type: Parameter
constant: true
value: 0.00681669584405203
name: Hog1PP_0

COPASI45
type: Parameter
notes: Maximal Fps1 glycerol transport capacity.
constant: false
value: 4.28194136809108E-4
name: ks

COPASI96
type: Reaction
kinetic law: 171565
modifier: 688495
reactant: 171553
notes: mRNA degradation
is version of: GO:0006402
fast: false
reversible: false
name: v10

COPASI12
type: Species
in compartment: COPASI2
constant: false
is version of: CHEBI:33699
initial concentration: NaN
initial amount: 1.003
name: RNA
has only substance units: true
notes: Placeholder for transcribed genes. Initial value derived from data, i.e. initial percentage of maximum.
boundary condition: false

171553
type: SpeciesReference
stoichiometry: 1.0
species: COPASI12

COPASI91
type: Reaction
name: v11
product: 171373
fast: false
notes: Protein/enzyme production/synthesis
kinetic law: 171397
is version of: GO:0006412
modifier: 171385
modifier: 688411
reversible: false

171373
type: SpeciesReference
stoichiometry: 1.0
species: COPASI11

COPASI51
type: Parameter
constant: false
value: 242026.122077737
name: Turgor2Osm

COPASI24
type: Parameter
notes: Mixing time [s] of salt in the medium.
constant: true
value: 10.0
name: tm

COPASI76
type: Parameter
notes: 〖907f〗_n2μM: molecule numbers from http://yeastgfp.yeastgenome.org/
constant: true
value: 0.0510557334969518
name: Fps1TotalIni

170172
type: SBMLUnitsDef
unit: 170184
name: substance

170184
type: SBMLUnit
exponent: 1.0
scale: -6
multipler: 1.0
kind: MOLE

COPASI49
type: Parameter
constant: false
value: 19.9
name: FitProteinrel

COPASI71
type: Parameter
constant: true
value: 180000.0
name: Glyin_0

COPASI44
type: Parameter
constant: true
value: 1.78587
name: kHog1dephos

COPASI95
type: Reaction
modifier: 688483
notes: Gene transcription. Hog1 mediated transcription also involves other proteins that are potentially limiting a saturation kinetic is assumed.
kinetic law: 171541
product: 171517
modifier: 171529
is version of: GO:0006351
fast: false
name: v9
reversible: false

171517
type: SpeciesReference
stoichiometry: 1.0
species: COPASI12

COPASI32
type: Parameter
notes: Fraction of activated Hog1 molecule in the nucleous upon maximal activation.
constant: true
value: 0.8
name: maxHog1nucf

COPASI75
type: Parameter
notes: Linear activation function for the Sln1 branch.
constant: true
value: 1.0
name: Sln1BranchActive

170329
type: Parameter
constant: true
value: 0.506878
name: v9_Km

COPASI18
type: Species
has only substance units: true
in compartment: 170196
boundary condition: false
constant: false
initial concentration: NaN
name: Fps1P
is encoded by: S000003966
notes: Membrane bound closed form of aquaglyceroporin Fps1 (assumed to be independent from volume change) . Initially half of the channels is assumed to be closed.
initial amount: 0.0255278667484759
is version of: P23900

COPASI48
type: Parameter
constant: false
value: 2.23
name: FitHog1PPrel

COPASI99
type: Reaction
fast: false
notes: Constitutive dissociation of the scaffold complex.
product: 171697
reactant: 171685
product: 171709
reversible: false
is version of: GO:0043241
modifier: 688567
name: v4
kinetic law: 171721

171697
type: SpeciesReference
stoichiometry: 1.0
species: COPASI14

171685
type: SpeciesReference
stoichiometry: 1.0
species: COPASI15

171709
type: SpeciesReference
stoichiometry: 1.0
species: COPASI8

COPASI36
type: Parameter
notes: Factor converting number of molecules in M concentrations per cell.
constant: true
value: 5.62907756305974E-5
name: N2uM

170317
type: Parameter
constant: true
value: 18.1824
name: v9_k

COPASI67
type: Parameter
notes: Osmotic salt shock. Starts at time t_s and has a certain mixing time t_m.
constant: false
value: 258200.0
name: cen

COPASI31
type: Parameter
notes: Solid or minimal volume of the cell.
constant: true
value: 20.5
name: Vb

COPASI41
type: Parameter
constant: true
value: 0.4
name: NaCl

COPASI4
type: Compartment
notes: Total cell volume.
is version of: GO:0005623
constant: false
size: 50.0
spatial dimensions: 3.0
name: V

COPASI35
type: Parameter
constant: true
value: 2.23
name: Hog1PPrelIniwt

COPASI66
type: Parameter
constant: true
value: 600.0
name: ts

COPASI40
type: Parameter
notes: Linear activation function for the Sln1 branch with feedback.
constant: false
value: 7.10539561053171E-4
name: ActivationSln1_nlfb

COPASI29
type: Parameter
notes: Initial osmolarity of the medium (Schaber, et al., 2010)
constant: true
value: 260000.0
name: ce_0

170233
type: Parameter
constant: false
value: 0.500000000000001
name: TurgorDeactivation

COPASI54
type: Parameter
constant: false
value: 3.4
name: FitRNArel

ORG:
  orgname: OvGU
n:
  given: Joerg
  family: Schaber

170221
type: Parameter
constant: true
value: 1.0
name: Sho1BranchActive

COPASI28
type: Parameter
notes: Minimal cell volume (as fraction of total) (Schaber, et al., 2010)
constant: true
value: 0.41
name: minf

COPASI58
type: Parameter
notes: 〖2330f〗_n2μM: molecule numbers from http://yeastgfp.yeastgenome.org/
constant: true
value: 0.131157507219292
name: Sho1Total

170257
type: Parameter
constant: true
value: 0.607124
name: v2_k

COPASI63
type: Parameter
notes: Intracellular osmolytically active concentration.
constant: false
value: 502026.122077738
name: Osmin

COPASI17
type: Species
initial amount: 0.0255278667484759
in compartment: 170196
constant: false
is version of: P23900
has only substance units: true
notes: Membrane bound open form of aquaglyceroporin Fps1 (assumed to be independent from volume change). Initially half of the channels is assumed to be open.
boundary condition: false
initial concentration: NaN
is encoded by: S000003966
name: Fps1

COPASI87
type: Reaction
kinetic law: 171205
modifier: 688303
reversible: false
notes: Simple cell surface area dependent linear activation of Pbs2 through the Sln1 branch, with transient inhibition by activated Hog1.
name: v1_fb
reactant: 171169
modifier: 688315
product: 171181
fast: false
modifier: 171193
is version of: GO:0016310

171169
type: SpeciesReference
stoichiometry: 1.0
species: COPASI8

171181
type: SpeciesReference
stoichiometry: 1.0
species: COPASI9

COPASI57
type: Parameter
notes: RNA degradation rate.
constant: true
value: 7.09644965005112
name: RNA_deg_k_MM

COPASI83
type: Parameter
constant: true
value: 0.001
name: Hog1Activity

170245
type: Parameter
constant: true
value: 600.0
name: tinh

170449
type: Parameter
constant: true
value: 0.420741
name: v13_Km

COPASI102
type: Reaction
modifier: 688627
reversible: false
notes: Activated Hog1 inhibited channel opening, in conjunction with v15_5.
product: 171841
reactant: 171829
modifier: 688639
name: v16_6
modifier: 171853
is version of: GO:0016311
fast: false
kinetic law: 171865

171841
type: SpeciesReference
stoichiometry: 1.0
species: COPASI17

171829
type: SpeciesReference
stoichiometry: 1.0
species: COPASI18

COPASI62
type: Parameter
notes: Extracellular osmolytically active concentration.
constant: false
value: 260000.0
name: Osmex

COPASI82
type: Parameter
constant: true
value: 0.00529124000000002
name: v16_7_k

COPASI90
type: Reaction
fast: false
reactant: 171325
name: v6
notes: Constitutive phosphatase dependent de-phosphorylation.
is version of: GO:0016311
modifier: 688387
product: 171337
reversible: false
modifier: 688399
modifier: 171349
kinetic law: 171361

171325
type: SpeciesReference
stoichiometry: 1.0
species: COPASI13

171337
type: SpeciesReference
stoichiometry: 1.0
species: COPASI6

COPASI50
type: Parameter
constant: false
value: 17.0
name: FitGlyinrel

COPASI23
type: Parameter
notes: Factor converting concentrations in  to pressures in MPa.
constant: true
value: 1.0E-9
name: c2p

COPASI19
type: Parameter
notes: Gas constant.
constant: true
value: 8.314
name: R

COPASI101
type: Reaction
notes: Turgor dependent closure of Fps1, in conjunction with v16_5, v16_6, v16_7, v16_8.
kinetic law: 171817
modifier: 688615
reversible: false
fast: false
product: 171805
reactant: 171793
name: v15_5
is version of: GO:0016310

171805
type: SpeciesReference
stoichiometry: 1.0
species: COPASI18

171793
type: SpeciesReference
stoichiometry: 1.0
species: COPASI17

W3 CDTF: Fri Dec 14 14:24:40 GMT 2012

COPASI70
type: Parameter
constant: true
value: 0.034
name: RNA_0

COPASI86
type: Parameter
constant: true
value: 1.0
name: Fps1Delta

COPASI43
type: Parameter
constant: true
value: 42.6396538263077
name: kHog1phos1

COPASI39
type: Parameter
notes: Turgor pressure [MPa].
constant: false
value: 0.609999999999999
name: Turgor

COPASI94
type: Reaction
notes: Gradient driven glycerol flow out of the cell, where k_tr characterises the state of the aquaglyceroporin channel Fps1 and k_s the maximal transport capacity of the channel.
reactant: 171481
reversible: true
modifier: 688471
fast: false
name: v14
kinetic law: 171505
is version of: GO:0015793
product: 171493
modifier: 688459

171481
type: SpeciesReference
stoichiometry: 1.0
species: COPASI5

171493
type: SpeciesReference
stoichiometry: 1.0
species: COPASI16

COPASI16
type: Species
notes: Extracellular glycerol, assumed to be 100 times lower than 〖Gly〗_in.
initial amount: 9.0E7
is version of: CHEBI:17754
has only substance units: true
boundary condition: false
initial concentration: NaN
in compartment: COPASI3
name: Glyex
constant: false

170365
type: Parameter
constant: true
value: 2.0793
name: v3_fb_h

COPASI7
type: Species
in compartment: COPASI2
name: Hog1PP
notes: Double phosphorylated, i.e. active, MAP kinase. It was derived from data that 2.23 % of the maximal phosphorylation is the steady state activation. f_n is the fraction in the nucleus at maximal phohsporylation.
constant: false
has only substance units: true
is version of: C00562
boundary condition: false
initial amount: 0.201092527399535
is version of: P32485
initial concentration: NaN

ORG:
  orgname: EMBL-EBI
n:
  given: Vijayalakshmi
  family: Chelliah

COPASI22
type: Parameter
notes: Osmotic coefficient for salt.
constant: true
value: 0.93
name: phi

COPASI74
type: Parameter
notes: Linear activation function for the Sln1 branch with feedback.
constant: false
value: 0.0036065403549782
name: ActivationSho1_nlfb

COPASI47
type: Parameter
constant: false
value: 100.0
name: FitVrel

COPASI98
type: Reaction
modifier: 688543
is version of: GO:0016311
modifier: 688555
modifier: 171661
product: 171649
reversible: false
notes: Constitutive phosphatase dependent de-phosphorylation.
name: v8
reactant: 171637
kinetic law: 171673
fast: false

171649
type: SpeciesReference
stoichiometry: 1.0
species: COPASI13

171637
type: SpeciesReference
stoichiometry: 1.0
species: COPASI7

170353
type: Parameter
constant: true
value: 0.297524
name: v3_fb_Ki

COPASI42
type: Parameter
notes: Steady state activation of the Sln1 branch with feedback.
constant: true
value: -7.10539561053171E-4
name: ActivOffsetSln1_nlfb

COPASI93
type: Reaction
modifier: 171445
modifier: 688447
is version of: GO:0006114
notes: Glycerol production. As glycerol concentration also involves other proteins and cofactors that are potentially limiting a saturation kinetic is assumed. In addition, effective enzyme concentration can be enhanced as a function of activated Hog1.
reversible: false
product: 171433
modifier: 688435
modifier: 171457
kinetic law: 171469
fast: false
name: v13

171433
type: SpeciesReference
stoichiometry: 1.0
species: COPASI5

COPASI53
type: Parameter
notes: Protein degradation rate.
constant: true
value: 6.78688610600496E-5
name: Protein_deg_k

COPASI26
type: Parameter
notes: Initial turgor pressure (Schaber, et al., 2010)
constant: true
value: 0.61
name: P0

COPASI80
type: Parameter
constant: true
value: 0.00264562
name: v16_5_k

COPASI38
type: Parameter
notes: Initial non-permeable cellular osmolyte concentration.
constant: true
value: 322026.122077738
name: cin_0

COPASI69
type: Parameter
constant: true
value: 0.0454266559338921
name: Protein_0

COPASI73
type: Parameter
notes: Steady state activation of the Sln1 branch with feedback.
constant: true
value: -0.0036065403549782
name: ActivOffsetSho1_nlfb

COPASI81
type: Parameter
constant: true
value: 0.00320327093093651
name: v16_6_k

COPASI33
type: Parameter
notes: Non-turgid volume.
constant: true
value: 47.9119840840503
name: VP_0

COPASI78
type: Parameter
notes: Hill parameter of the turgor-activate Fps1 closure.
constant: true
value: 2.0
name: TurgorActiv_h

170389
type: Parameter
constant: true
value: 0.00529124
name: v15_5_k

COPASI30
type: Parameter
notes: Initial total cell volume.
constant: true
value: 50.0
name: V_0

COPASI46
type: Parameter
notes: Fps1 channel closure.
constant: false
value: 0.5
name: Fps1TransportCapacitty

COPASI97
type: Reaction
modifier: 171613
modifier: 688531
fast: false
modifier: 688507
notes: Linear phosphorylation of Hog1 by either the scaffold complex (Sho1-branch) or activated Pbs2 (Sln1-branch).
name: v7
modifier: 171601
reversible: false
is version of: GO:0016310
modifier: 688519
product: 171589
kinetic law: 171625
reactant: 171577

171589
type: SpeciesReference
stoichiometry: 1.0
species: COPASI7

171577
type: SpeciesReference
stoichiometry: 1.0
species: COPASI13

170377
type: Parameter
constant: true
value: 0.00459138
name: v3_fb_k

COPASI34
type: Parameter
notes: Total cell surface area.
constant: false
value: 65.6342903668733
name: Area

COPASI77
type: Parameter
constant: false
value: 0.0510557334969518
name: Fps1TotalTrans

170305
type: Parameter
constant: true
value: 9.06781E-5
name: v11_k

W3 CDTF: Thu Nov 22 18:31:29 GMT 2012