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arnold10

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arnold10

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A quantitative comparison of Calvin-Benson cycle models.

Anne Arnold
Zoran Nikoloski

Trends Plant Sci. 2011; 16 (12): 676-683

PubMed: 22001849
DOI: 10.1016/j.tplants.2011.09.004
ISSN: 1878-4372
URL: https://ncbi.nlm.nih.gov/pubmed/22001849

Abstract

The Calvin-Benson cycle (CBC) provides the precursors for biomass synthesis necessary for plant growth. The dynamic behavior and yield of the CBC depend on the environmental conditions and regulation of the cellular state. Accurate quantitative models hold the promise of identifying the key determinants of the tightly regulated CBC function and their effects on the responses in future climates. We provide an integrative analysis of the largest compendium of existing models for photosynthetic processes. Based on the proposed ranking, our framework facilitates the discovery of best-performing models with regard to metabolomics data and of candidates for metabolic engineering.

Unit definitions 2

Unit definitions have no effect on the numerical analysis of the model. It remains the responsibility of the modeler to ensure the internal numerical consistency of the model. If units are provided, however, the consistency of the model units will be checked.

Name	Definition
—	1.0 litre
—	1.0 mole

Compartments 2

Id	Name	Spatial dimensions	Size
chloroplast	chloroplast	3.0	1.0
cytosol	cytosol	3.0	1.0

Species 53

Id	Name	Initial quantity	Compartment	Fixed
ADP	ADP	0.00082	chloroplast (chloroplast)	✔
ADPG	ADPG	0.000006295	chloroplast (chloroplast)	✘
ADPc	ADPc	0.00064	cytosol (cytosol)	✔
ATP	ATP	0.00068	chloroplast (chloroplast)	✘
ATPc	ATPc	0.00036	cytosol (cytosol)	✔
CO2	CO2	0.00000436233478328384	chloroplast (chloroplast)	✔
DHAP	DHAP	0.000250091578940897	chloroplast (chloroplast)	✔
DHAPc	DHAPc	0.00115042126312813	cytosol (cytosol)	✔
E	E	0.0000092303030303025	chloroplast (chloroplast)	✔
E4P	E4P	0.00005	chloroplast (chloroplast)	✘
EOP	EOP	0.0002966	chloroplast (chloroplast)	✘
EP	EP	0.0007045	chloroplast (chloroplast)	✘
EPG	EPG	0.0003206	chloroplast (chloroplast)	✘
EPP	EPP	0.0002711	chloroplast (chloroplast)	✘
ER	ER	0.001201	chloroplast (chloroplast)	✘
F26BPc	F26BPc	0.0000000078	cytosol (cytosol)	✘
F6P	F6P	0.000733422825505269	chloroplast (chloroplast)	✔
F6Pc	F6Pc	0.00193356926724116	cytosol (cytosol)	✔
FBP	FBP	0.00067	chloroplast (chloroplast)	✘
FBPc	FBPc	0.002	cytosol (cytosol)	✘
G1P	G1P	0.000733034781579417	chloroplast (chloroplast)	✔
G1Pc	G1Pc	0.00193254624234574	cytosol (cytosol)	✔
G6P	G6P	0.000733542392915314	chloroplast (chloroplast)	✔
G6Pc	G6Pc	0.0019338844904131	cytosol (cytosol)	✔
GAP	GAP	0.000249908421059103	chloroplast (chloroplast)	✔
GAPc	GAPc	0.00114957873687187	cytosol (cytosol)	✔
H	H	0.0891250931577478	chloroplast (chloroplast)	✔
Hc	Hc	0.158489318357816	cytosol (cytosol)	✔
HeP	HeP	0.0022	chloroplast (chloroplast)	✘
HePc	HePc	0.0058	cytosol (cytosol)	✘
NADP	NADP	0.00029	chloroplast (chloroplast)	✔
NADPH	NADPH	0.00021	chloroplast (chloroplast)	✔
O2	O2	0.00026	chloroplast (chloroplast)	✔
PGA	PGA	0.0024	chloroplast (chloroplast)	✘
PGAc	PGAc	0.0	cytosol (cytosol)	✘
PeP	PeP	0.00025	chloroplast (chloroplast)	✘
Pi	Pi	0.0109928959090909	chloroplast (chloroplast)	✔
PiPi	PiPi	0.000001	chloroplast (chloroplast)	✔
PiPic	PiPic	0.00004	cytosol (cytosol)	✔
Pic	Pic	0.00157545454545454	cytosol (cytosol)	✔
R5P	R5P	0.0000833492525797069	chloroplast (chloroplast)	✔
Ru5P	Ru5P	0.0000833275817740362	chloroplast (chloroplast)	✔
RuBP	RuBP	0.002	chloroplast (chloroplast)	✘
S7P	S7P	0.002	chloroplast (chloroplast)	✘
SBP	SBP	0.0003	chloroplast (chloroplast)	✘
SucPc	SucPc	0.0	cytosol (cytosol)	✘
Succ	Succ	0.0	cytosol (cytosol)	✔
TP	TP	0.0005	chloroplast (chloroplast)	✘
TPc	TPc	0.0023	cytosol (cytosol)	✘
UDPGc	UDPGc	0.00057	cytosol (cytosol)	✘
UDPc	UDPc	0.00064	cytosol (cytosol)	✔
UTPc	UTPc	0.00036	cytosol (cytosol)	✔
X5P	X5P	0.0000833231656462569	chloroplast (chloroplast)	✔

Initial assignments 0

Initial assignments are expressions that are evaluated at time=0. It is not recommended to create initial assignments for all model entities. Restrict the use of initial assignments to cases where a value is expressed in terms of values or sizes of other model entities. Note that it is not permitted to have both an initial assignment and an assignment rule for a single model entity.

Definition

Reactions 27

Id	Name	Reaction Equation and Kinetic Law
AGPase	AGPase	HeP + ATP = ADPG + PiPi chloroplast * function_28(Vm, PGA, Pi, G1P, ATP, ADPG, PiPi, q, Ks1, Ks2, Kp1, Kp2)
ATP_S	ATP synthase	ADP + Pi = ATP chloroplast * function_24(ADP, Pi, ATP, q12, Ks1, Ks2, Kp12, Vm)
F26BPc_P	F26BPc phosphatase	F26BPc > HePc + Pic cytosol * function_33(Vm, F26BPc, Ks1, TPc, PGAc, Kr1, Pic, Kr3, HePc, Kr4)
F6P_TK	Transketolase (F6P)	HeP + TP = E4P + PeP chloroplast * Transketolase(Vm, q, F6P, GAP, E4P, X5P, K1, K2, K1s2, K2s1, S7P, K2r1, K2s2, R5P)
F6Pc_K	F6Pc kinase	HePc > F26BPc + ADPc cytosol * function_32(Vm, F6Pc, Ks1, Pic, Kr1, TPc, PGAc, Kr2)
FBP_A	Aldolase (FBP)	{2.0}TP = FBP chloroplast * function_24(GAP, DHAP, FBP, q, Ks1, Ks2, Kp1, Vm)
FBPase	FBPase	FBP = HeP + Pi chloroplast * function_25(Vm, FBP, F6P, Pi, q, Ks1, Kp1, Kp2)
FBPc_A	cyt. Aldolase (FBP)	{2.0}TPc = FBPc cytosol * function_24(GAPc, DHAPc, FBPc, q, Ks1, Ks2, Kp1, Vm)
FBPcase	cyt. FBPase	FBPc = HePc + Pic cytosol * function_30(Vm, FBPc, F6Pc, Pic, q, Ks1, F26BPc, Kr1, Kp1, Kp2)
PGA_K	PGA kinase	PGA + ATP = TP + ADP + Pi chloroplast * function_23(Vm, PGA, ATP, GAP, ADP, Pi, q1, Ks1, Ks2, Kp1, Kp2, Kp3)
Ru5P_K	Ru5P kinase	PeP + ATP = RuBP + ADP chloroplast * function_27(Ru5P, ATP, RuBP, ADP, q, Ks1, Ks2, Kp1, Kp2, Vm)
RuBisCO_1	RuBisCO (1)	RuBP + E = ER chloroplast * (k1 * RuBP * E - k2 * ER)
RuBisCO_2_CO2	RuBisCO (2) - V(CO2)	ER + CO2 > EPP chloroplast * k1 * ER * CO2
RuBisCO_4	RuBisCO (4)	EPP = PGA + EP chloroplast * (k1 * EPP - k2 * PGA * EP)
RuBisCO_5_EOP	RuBisCO (5) - EOP	EOP = ∅ chloroplast * function_34(W4)
RuBisCO_5_EP	RuBisCO (5) - EP	EP = PGA + E chloroplast * (k1 * EP - k2 * PGA * E)
RuBisCO_6_O2	RuBisCO (6) - V(O2)	ER + O2 + {0.5}ATP > EPG + {0.5}PGA + {0.5}ADP chloroplast * k1 * ER * O2
RuBisCO_7	RuBisCO (7)	EPG > EP chloroplast * k1 * EPG
S7P_TK	Transketolase (S7P)	S7P + TP = {2.0}PeP chloroplast * Transketolase(Vm, q, S7P, GAP, R5P, X5P, K1, K2, K1s2, K2s1, F6P, K2r1, K2s2, E4P)
SBP_A	Aldolase (SBP)	E4P + TP = SBP chloroplast * function_24(E4P, DHAP, SBP, q, Ks1, Ks2, Kp1, Vm)
SBPase	SBPase	SBP = S7P + Pi chloroplast * function_26(Vm, SBP, S7P, Pi, q, Ks1, Kp1, Kp2)
StS	Starch synthase	ADPG = ADP chloroplast * function_29(Vm, ADPG, ADP, q, Ks1, Kp1)
SucPc_P	Sucrose phosphatase	SucPc = Succ + Pic cytosol * function_25(Vm, SucPc, Succ, Pic, q, Ks1, Kp1, Kp2)
SucPc_S	Sucrose synthase	HePc + UDPGc = UDPc + SucPc + Hc cytosol * function_31(Vm, F6Pc, UDPGc, UDPc, SucPc, Hc, q, Ks1, Pic, Kr11, Ks2, Kp1, Kp2, Kr12)
TPT_PGA	TPT (PGA)	PGA = PGAc TPTout(Vm, PGA, Ks, TP, Kr1, Pi, Kr2, PGAc, Kp, TPc, Kr3, Pic, Kr4)
TPT_TP	TPT (TP)	TP = TPc TPTout(Vm, TP, Ks, PGA, Kr1, Pi, Kr2, TPc, Kp, PGAc, Kr3, Pic, Kr4)
UGPase	UGPase	HePc + UTPc = UDPGc + PiPic cytosol * function_27(G1Pc, UTPc, UDPGc, PiPic, q, Ks1, Ks2, Kp1, Kp2, Vm)

Parameters 123 12

Global parameters

Id	Value
ADT	0.0015
ADTc	0.001
Et	0.0028030303030303
Kp12	224014.808032967
NADPT	0.0005
PiT	0.0284090909090909
PiTc	0.0170454545454545
UDTc	0.001
V28	0.00007386364
W4	-0.00532314322950372
q1	0.129053067280279
q12	2227862541257.35

Local parameters

Id	Value	Reaction
Vm	0.00113636	AGPase (AGPase)
q	0.11059	AGPase (AGPase)
Ks1	0.0010398	AGPase (AGPase)
Ks2	0.00011023	AGPase (AGPase)
Kp1	0.00053013	AGPase (AGPase)
Kp2	0.01951	AGPase (AGPase)
Ks1	0.00031808	ATP_S (ATP synthase)
Ks2	0.00031612	ATP_S (ATP synthase)
Vm	0.0284091	ATP_S (ATP synthase)
Vm	0.000000000205284	F26BPc_P (F26BPc phosphatase)
Ks1	0.000000001	F26BPc_P (F26BPc phosphatase)
Kr1	0.002	F26BPc_P (F26BPc phosphatase)
Kr3	0.001	F26BPc_P (F26BPc phosphatase)
Kr4	0.0001	F26BPc_P (F26BPc phosphatase)
Vm	0.170455	F6P_TK (Transketolase (F6P))
q	0.99943	F6P_TK (Transketolase (F6P))
K1	0.00061349	F6P_TK (Transketolase (F6P))
K2	0.00011438	F6P_TK (Transketolase (F6P))
K1s2	0.00027035	F6P_TK (Transketolase (F6P))
K2s1	0.0005407	F6P_TK (Transketolase (F6P))
K2r1	0.00017677	F6P_TK (Transketolase (F6P))
K2s2	0.000090464	F6P_TK (Transketolase (F6P))
Vm	0.000000102614	F6Pc_K (F6Pc kinase)
Ks1	0.001	F6Pc_K (F6Pc kinase)
Kr1	0.001	F6Pc_K (F6Pc kinase)
Kr2	0.0015	F6Pc_K (F6Pc kinase)
q	1.18815	FBP_A (Aldolase (FBP))
Ks1	0.00027035	FBP_A (Aldolase (FBP))
Ks2	0.00036393	FBP_A (Aldolase (FBP))
Kp1	0.000020129	FBP_A (Aldolase (FBP))
Vm	0.022727	FBP_A (Aldolase (FBP))
Vm	0.011364	FBPase (FBPase)
q	0.77294	FBPase (FBPase)
Ks1	0.000032842	FBPase (FBPase)
Kp1	0.000063429	FBPase (FBPase)
Kp2	0.0017914	FBPase (FBPase)
q	1.00224	FBPc_A (cyt. Aldolase (FBP))
Ks1	0.000278407	FBPc_A (cyt. Aldolase (FBP))
Ks2	0.000374778	FBPc_A (cyt. Aldolase (FBP))
Kp1	0.0000210226	FBPc_A (cyt. Aldolase (FBP))
Vm	0.00568182	FBPc_A (cyt. Aldolase (FBP))
Vm	0.00113636	FBPcase (cyt. FBPase)
q	0.792367	FBPcase (cyt. FBPase)
Ks1	0.000022129	FBPcase (cyt. FBPase)
Kr1	0.0000011065	FBPcase (cyt. FBPase)
Kp1	0.000065319	FBPcase (cyt. FBPase)
Kp2	0.0018624	FBPcase (cyt. FBPase)
Vm	0.0170455	PGA_K (PGA kinase)
Ks1	0.0011122	PGA_K (PGA kinase)
Ks2	0.0003307	PGA_K (PGA kinase)
Kp1	0.00027035	PGA_K (PGA kinase)
Kp2	0.00053013	PGA_K (PGA kinase)
Kp3	0.0027397	PGA_K (PGA kinase)
q	1.05289	Ru5P_K (Ru5P kinase)
Ks1	0.0000363934	Ru5P_K (Ru5P kinase)
Ks2	0.00055117	Ru5P_K (Ru5P kinase)
Kp1	0.0000995868	Ru5P_K (Ru5P kinase)
Kp2	0.0000911825	Ru5P_K (Ru5P kinase)
Vm	0.568182	Ru5P_K (Ru5P kinase)
k1	50000.0	RuBisCO_1 (RuBisCO (1))
k2	0.9	RuBisCO_1 (RuBisCO (1))
k1	300000.0	RuBisCO_2_CO2 (RuBisCO (2) - V(CO2))
k1	6.0	RuBisCO_4 (RuBisCO (4))
k2	0.0	RuBisCO_4 (RuBisCO (4))
k1	6.0	RuBisCO_5_EP (RuBisCO (5) - EP)
k2	70000.0	RuBisCO_5_EP (RuBisCO (5) - EP)
k1	3030.3	RuBisCO_6_O2 (RuBisCO (6) - V(O2))
k1	3.0	RuBisCO_7 (RuBisCO (7))
Vm	0.0821023	S7P_TK (Transketolase (S7P))
q	0.99996	S7P_TK (Transketolase (S7P))
K1	0.00061349	S7P_TK (Transketolase (S7P))
K2	0.00011438	S7P_TK (Transketolase (S7P))
K1s2	0.00027035	S7P_TK (Transketolase (S7P))
K2s1	0.00017677	S7P_TK (Transketolase (S7P))
K2r1	0.0005407	S7P_TK (Transketolase (S7P))
K2s2	0.000090464	S7P_TK (Transketolase (S7P))
q	1.18815	SBP_A (Aldolase (SBP))
Ks1	0.00017677	SBP_A (Aldolase (SBP))
Ks2	0.00036393	SBP_A (Aldolase (SBP))
Kp1	0.000020129	SBP_A (Aldolase (SBP))
Vm	0.011364	SBP_A (Aldolase (SBP))
Vm	0.00568182	SBPase (SBPase)
q	0.77294	SBPase (SBPase)
Ks1	0.000012713	SBPase (SBPase)
Kp1	0.000015597	SBPase (SBPase)
Kp2	0.006744	SBPase (SBPase)
Vm	0.00284091	StS (Starch synthase)
q	1.00326	StS (Starch synthase)
Ks1	0.000212052	StS (Starch synthase)
Kp1	0.000636157	StS (Starch synthase)
Vm	0.0010267	SucPc_P (Sucrose phosphatase)
q	1.35286	SucPc_P (Sucrose phosphatase)
Ks1	0.00005354	SucPc_P (Sucrose phosphatase)
Kp1	0.01	SucPc_P (Sucrose phosphatase)
Kp2	0.002191	SucPc_P (Sucrose phosphatase)
Vm	0.0000738636	SucPc_S (Sucrose synthase)
q	1.00012	SucPc_S (Sucrose synthase)
Ks1	0.000278407	SucPc_S (Sucrose synthase)
Kr11	0.00920241	SucPc_S (Sucrose synthase)
Ks2	0.000110717	SucPc_S (Sucrose synthase)
Kp1	0.000642157	SucPc_S (Sucrose synthase)
Kp2	0.000374778	SucPc_S (Sucrose synthase)
Kr12	0.00164329	SucPc_S (Sucrose synthase)
Vm	0.0568182	TPT_PGA (TPT (PGA))
Ks	0.00089213	TPT_PGA (TPT (PGA))
Kr1	0.000093583	TPT_PGA (TPT (PGA))
Kr2	0.000098597	TPT_PGA (TPT (PGA))
Kp	0.00054107	TPT_PGA (TPT (PGA))
Kr3	0.000096372	TPT_PGA (TPT (PGA))
Kr4	0.000094837	TPT_PGA (TPT (PGA))
Vm	0.0568182	TPT_TP (TPT (TP))
Ks	0.000093583	TPT_TP (TPT (TP))
Kr1	0.00089213	TPT_TP (TPT (TP))
Kr2	0.000098597	TPT_TP (TPT (TP))
Kp	0.000096372	TPT_TP (TPT (TP))
Kr3	0.00054107	TPT_TP (TPT (TP))
Kr4	0.000094837	TPT_TP (TPT (TP))
q	1.6219	UGPase (UGPase)
Ks1	0.000032124	UGPase (UGPase)
Ks2	0.0002364	UGPase (UGPase)
Kp1	0.00014393	UGPase (UGPase)
Kp2	0.0013192	UGPase (UGPase)
Vm	0.00410568	UGPase (UGPase)

Rules 0 0 20

Assignment rules

Definition
Pic = PiTc - 2.0 * (FBPc + UTPc + ATPc + PiPic) - (PGAc + TPc + HePc + SucPc + UDPGc + UDPc + ADPc)
F6P = HeP * 0.999837 / (1.0 + 0.999837 + 0.999308)
Ru5P = PeP * 0.99974 / (1.0 + 0.99974 + 0.99974 / 1.000053)
GAP = TP / (1.0 + 1.0007329)
ADPc = ADTc - ATPc
W4 = 6.0 * EOP - 70000.0 * Pi * E
E = Et - (ER + EPP + EPG + EP + EOP)
CO2 = (1.2e-5 / (0.38 + 0.015) + 8.0 * 4.0 * 0.00055 * 3030.3 * ER * O2 / 2.0) / (1.0 / (0.38 + 0.015) + 8.0 * 4.0 * 0.00055 * 300000.0 * ER)
DHAPc = 1.0007329 * TPc / (1.0 + 1.0007329)
GAPc = TPc / (1.0 + 1.0007329)
Pi = PiT - 2.0 * (EPP + EPG + RuBP + FBP + SBP + ATP + PiPi) - (EP + PGA + TP + HeP + E4P + S7P + PeP + ADP + ADPG)
ADP = ADT - ATP
R5P = PeP / (1.0 + 0.99974 + 0.99974 / 1.000053)
F6Pc = HePc * 0.999837 / (1.0 + 0.999837 + 0.999308)
G6Pc = HePc / (1.0 + 0.999837 + 0.999308)
DHAP = 1.0007329 * TP / (1.0 + 1.0007329)
G1Pc = HePc * 0.999308 / (1.0 + 0.999837 + 0.999308)
G1P = HeP * 0.999308 / (1.0 + 0.999837 + 0.999308)
G6P = HeP / (1.0 + 0.999837 + 0.999308)
X5P = PeP * (0.99974 / 1.000053) / (1.0 + 0.99974 + 0.99974 / 1.000053)

Rate rules

Definition

Algebraic rules

Definition

Function definitions 14

Definition
function_28(Vm, r1, r2, s1, s2, p1, p2, q, Ks1, Ks2, Kp1, Kp2) = Vm * pow(r1 / r2, 2) * (s1 * s2 - p1 * p2 / q) / (Ks1 * Ks2 * ((1 + s1 / Ks1) * (1 + s2 / Ks2) + (1 + p1 / Kp1) * (1 + p2 / Kp2) - 1))
function_25(Vm, s1, p1, p2, q, Ks1, Kp1, Kp2) = Vm * (s1 - p1 * p2 / q) / (Ks1 * (1 + s1 / Ks1 + p1 / Kp1 + p1 * p2 / (Kp1 * Kp2)))
function_31(Vm, s1, s2, p1, p2, p3, q, Ks1, r1, Kr11, Ks2, Kp1, Kp2, Kr12) = Vm * s1 * (s1 * s2 - p1 * p2 * p3 / q) / (pow(Ks1 * (1 + r1 / Kr11), 2) * Ks2 * ((1 + pow(s1 / (Ks1 * (1 + r1 / Kr11)), 2)) * (1 + s2 / Ks2) + (1 + p1 / Kp1) * (1 + p2 / Kp2) - 1 + r1 / Kr12))
function_33(Vm, s1, Ks1, r1, r2, Kr1, r3, Kr3, r4, Kr4) = Vm * s1 / Ks1 * (1 + (r1 + r2) / Kr1) / (1 + r3 / Kr3 + r4 / Kr4)
function_26(Vm, s1, p1, p2, q, Ks1, Kp1, Kp2) = Vm * (s1 - p1 * p2 / q) / (Ks1 * (s1 / Ks1 + (1 + p1 / Kp1) * (1 + p2 / Kp2)))
function_24(s1, s2, p1, q, Ks1, Ks2, Kp1, Vm) = Vm * (s1 * s2 - p1 / q) / (Ks1 * Ks2 * ((1 + s1 / Ks1) * (1 + s2 / Ks2) + p1 / Kp1))
Transketolase(Vm, q, s1, s2, p1, p2, K1, K2, K1s2, K2s1, r1, K2r1, K2s2, r2) = Vm * (q * s1 * s2 - p1 * p2) / (K1 * K2 * (1 + (1 + s2 / K1s2) * (s1 / K2s1 + r1 / K2r1) + s2 / K2s2 + 1 / K2 * (p2 * (1 + p1 * r2 / K1) + p1 + r2)))
function_23(Vm, s1, s2, p1, p2, p3, q, Ks1, Ks2, Kp1, Kp2, Kp3) = Vm * (s1 * s2 - p1 * p2 * p3 / q) / (Ks1 * Ks2 * ((1 + s1 / Ks1) * (1 + s2 / Ks2) + p1 / Kp1 + p2 / Kp2 + p3 / Kp3 + p1 * p2 * p3 / (Kp1 * Kp2 * Kp3)))
function_27(s1, s2, p1, p2, q, Ks1, Ks2, Kp1, Kp2, Vm) = Vm * (s1 * s2 - p1 * p2 / q) / (Ks1 * Ks2 * ((1 + s1 / Ks1) * (1 + s2 / Ks2) + (1 + p1 / Kp1) * (1 + p2 / Kp2) - 1))
function_32(Vm, s1, Ks1, r1, Kr1, r2, r3, Kr2) = Vm * s1 / Ks1 * (1 + r1 / Kr1) / (1 + (r2 + r3) / Kr2)
function_29(Vm, s1, p1, q, Ks1, Kp1) = Vm * (s1 - p1 / q) / (Ks1 * (1 + s1 / Ks1 + p1 / Kp1))
TPTout(Vm, s, Ks, r1, Kr1, r2, Kr2, p, Kp, r3, Kr3, r4, Kr4) = Vm / (s / Ks + r1 / Kr1 + r2 / Kr2 + p / Kp + r3 / Kr3 + r4 / Kr4 + (s / Ks + r1 / Kr1 + r2 / Kr2) * (p / Kp + r3 / Kr3 + r4 / Kr4)) * (s * (p / Kp + r3 / Kr3 + r4 / Kr4) / Ks - p * (s / Ks + r1 / Kr1 + r2 / Kr2) / Kp)
function_30(Vm, s1, p1, p2, q, Ks1, r1, Kr1, Kp1, Kp2) = Vm * s1 * (s1 - p1 * p2 / q) / (pow(Ks1 * (1 + r1 / Kr1), 2) * (pow(s1 / (Ks1 * (1 + r1 / Kr1)), 2) + (1 + p1 / Kp1) * (1 + p2 / Kp2)))
function_34(v) = v

Events 0

Trigger	Assignments