The Matuszynska 2019 model is a kinetic model of photosynthesis that couples the photosynthetic electron transport chain (PETC) to the carbon-fixing reactions, developed to explore — from a theoretical perspective — how a plant balances the supply of energy (ATP) and reducing power (NADPH) against their demand. It resolves the full electron-transport chain — photosystem II (PSII), the plastoquinone (PQ) pool, the cytochrome b6f complex, plastocyanin, photosystem I (PSI), FNR and ATP synthase — together with the regulatory layers of non-photochemical quenching (PsbS protonation and the xanthophyll cycle) and light-harvesting complex state transitions, and connects this machinery to the Calvin-Benson-Bassham (CBB) cycle that consumes ATP and NADPH. By tracking lumen and stroma proton concentrations and the resulting pH, it links the proton motive force directly to ATP synthesis and to the activation of photoprotective quenching.
The model builds on the earlier Matuszyńska light-acclimation models (see the Matuszynska 2016 NPQ model and the fuller PhD-thesis version) by adding explicit carbon metabolism, which lets it address the mismatch between the fixed ATP:NADPH stoichiometry produced by linear electron flow and the different ratio required by the CBB cycle — a central question in understanding why mechanisms such as cyclic electron flow and quenching exist. It went on to serve as the foundation for the Saadat 2021 model, which extends it with the ascorbate-glutathione cycle and additional electron flows around PSI. Because it captures the coupling between light reactions and carbon fixation in a single, well-documented framework, this model is included in GreenSloth as a versatile base for studying the energetic regulation of photosynthesis.
| Symbol | ID | Initial value |
|---|---|---|
| _3PGA | 0.9928653922138561 | |
| BPGA | 0.0005297732935310749 | |
| GAP | 0.0062663539939955834 | |
| DHAP | 0.13785977143668732 | |
| FBP | 0.006133532145409954 | |
| F6P | 0.31271973359685457 | |
| G6P | 0.719255387166192 | |
| G1P | 0.041716812452951633 | |
| SBP | 0.013123745088361893 | |
| S7P | 0.15890073845176905 | |
| E4P | 0.007322797350442026 | |
| X5P | 0.022478763225333428 | |
| R5P | 0.037651927659696716 | |
| RUBP | 0.13184790283048484 | |
| RU5P | 0.015060770937455408 | |
| ATP | 1.612922506604933 | |
| Ferredoxine_oxidised | 3.8624032084329674 | |
| protons_lumen | 0.002208423037307405 | |
| Light_minus_harvesting_complex | 0.80137477470646 | |
| NADPH | 0.491395685599137 | |
| Plastocyanine_oxidised | 1.885391998090184 | |
| Plastoquinone_oxidised | 10.991562708096392 | |
| PsbS_de_minus_protonated | 0.9610220887579118 | |
| Violaxanthin | 0.9514408605906095 |
| Symbol | ID | Value |
|---|---|---|
| protons | 0.000012589254117941661 | |
| pH | 7.9 | |
| CO2_dissolved | 0.2 | |
| O2_dissolved_lumen | 8 | |
| PPFD | 100 | |
| bH | 100 | |
| F | 96.485 | |
| E0_PC | 0.38 | |
| E0_P700 | 0.48 | |
| E0_FA | -0.55 | |
| E0_Fd | -0.43 | |
| E0_NADP | -0.113 | |
| convf | 0.032 | |
| R | 0.0083 | |
| T | 298 | |
| Carotenoids_tot | 1 | |
| Fd_star | 5 | |
| PC_tot | 4 | |
| PSBS_tot | 1 | |
| LHC_tot | 1 | |
| gamma0 | 0.1 | |
| gamma1 | 0.25 | |
| gamma2 | 0.6 | |
| gamma3 | 0.15 | |
| kZSat | 0.12 | |
| E0_QA | -0.14 | |
| E0_PQ | 0.354 | |
| PQ_tot | 17.5 | |
| staticAntII | 0.1 | |
| staticAntI | 0.37 | |
| NADP_star | 0.8 | |
| A_star_P | 2.55 | |
| Pi_tot | 17.05 | |
| kf_atp_synthase | 20 | |
| HPR | 4.666666666666667 | |
| Pi_mol | 0.01 | |
| DeltaG0_ATP | 30.6 | |
| kcat_b6f | 2.5 | |
| kh_lhc_protonation | 3 | |
| kf_lhc_protonation | 0.0096 | |
| ksat_lhc_protonation | 5.8 | |
| kf_lhc_deprotonation | 0.0096 | |
| kf_cyclic_electron_flow | 1 | |
| kf_violaxanthin_deepoxidase | 0.0024 | |
| kh_violaxanthin_deepoxidase | 5 | |
| ksat_violaxanthin_deepoxidase | 5.8 | |
| kf_zeaxanthin_epoxidase | 0.00024 | |
| km_fnr_Ferredoxine_reduced | 1.56 | |
| km_fnr_NADP | 0.22 | |
| E0_fnr | 3 | |
| kcat_fnr | 500 | |
| kf_ndh | 0.002 | |
| PSII_total | 2.5 | |
| PSI_total | 2.5 | |
| kH0 | 500000000 | |
| kPQred | 250 | |
| kPCox | 2500 | |
| kFdred | 250000 | |
| k2 | 5000000000 | |
| kH | 5000000000 | |
| kF | 625000000 | |
| kf_proton_leak | 10 | |
| kPTOX | 0.01 | |
| kStt7 | 0.0035 | |
| km_lhc_state_transition_12 | 0.2 | |
| n_ST | 2 | |
| kPph1 | 0.0013 | |
| E0_rubisco | 1 | |
| kcat_rubisco_carboxylase | 2.72 | |
| km_rubisco_carboxylase_RUBP | 0.02 | |
| km_rubisco_carboxylase_CO2_dissolved | 0.0107 | |
| ki_rubisco_carboxylase_3PGA | 0.04 | |
| ki_rubisco_carboxylase_FBP | 0.04 | |
| ki_rubisco_carboxylase_SBP | 0.075 | |
| ki_rubisco_carboxylase_Orthophosphate | 0.9 | |
| ki_rubisco_carboxylase_NADPH | 0.07 | |
| kre_phosphoglycerate_kinase | 800000000 | |
| keq_phosphoglycerate_kinase | 0.00031 | |
| kre_gadph | 800000000 | |
| keq_gadph | 16000000 | |
| kre_triose_phosphate_isomerase | 800000000 | |
| keq_triose_phosphate_isomerase | 22 | |
| kre_aldolase_dhap_gap | 800000000 | |
| keq_aldolase_dhap_gap | 7.1 | |
| kre_aldolase_dhap_e4p | 800000000 | |
| keq_aldolase_dhap_e4p | 13 | |
| E0_fbpase | 1 | |
| kcat_fbpase | 1.6 | |
| km_fbpase_s | 0.03 | |
| ki_fbpase_F6P | 0.7 | |
| ki_fbpase_Orthophosphate | 12 | |
| kre_transketolase_gap_f6p | 800000000 | |
| keq_transketolase_gap_f6p | 0.084 | |
| kre_transketolase_gap_s7p | 800000000 | |
| keq_transketolase_gap_s7p | 0.85 | |
| E0_SBPase | 1 | |
| kcat_SBPase | 0.32 | |
| km_SBPase_s | 0.013 | |
| ki_SBPase_Orthophosphate | 12 | |
| kre_ribose_phosphate_isomerase | 800000000 | |
| keq_ribose_phosphate_isomerase | 0.4 | |
| kre_ribulose_phosphate_epimerase | 800000000 | |
| keq_ribulose_phosphate_epimerase | 0.67 | |
| E0_phosphoribulokinase | 1 | |
| kcat_phosphoribulokinase | 7.9992 | |
| km_phosphoribulokinase_RU5P | 0.05 | |
| km_phosphoribulokinase_ATP | 0.05 | |
| ki_phosphoribulokinase_3PGA | 2 | |
| ki_phosphoribulokinase_RUBP | 0.7 | |
| ki_phosphoribulokinase_Orthophosphate | 4 | |
| ki_phosphoribulokinase_4 | 2.5 | |
| ki_phosphoribulokinase_5 | 0.4 | |
| kre_g6pi | 800000000 | |
| keq_g6pi | 2.3 | |
| kre_phosphoglucomutase | 800000000 | |
| keq_phosphoglucomutase | 0.058 | |
| Orthophosphate_external | 0.5 | |
| km_ex_pga | 0.25 | |
| km_ex_gap | 0.075 | |
| km_ex_dhap | 0.077 | |
| km_N_translocator_Orthophosphate_external | 0.74 | |
| km_N_translocator_Orthophosphate | 0.63 | |
| kcat_N_translocator | 2 | |
| E0_N_translocator | 1 | |
| E0_ex_g1p | 1 | |
| km_ex_g1p_G1P | 0.08 | |
| km_ex_g1p_ATP | 0.08 | |
| ki_ex_g1p | 10 | |
| ki_ex_g1p_3PGA | 0.1 | |
| ki_ex_g1p_F6P | 0.02 | |
| ki_ex_g1p_FBP | 0.02 | |
| kcat_ex_g1p | 0.32 |
| Symbol | ID | Equation |
|---|---|---|
| RT | ||
| dG_pH | ||
| pH_lumen | ||
| Zeaxanthin | ||
| Ferredoxine_reduced | ||
| Plastocyanine_reduced | ||
| PsbS_protonated | ||
| Light_minus_harvesting_complex_protonated | ||
| Q | ||
| keq_Plastoquinone_reduced | ||
| Plastoquinone_reduced | ||
| PSII_cross_section | ||
| NADP | ||
| ADP | ||
| Orthophosphate | ||
| keq_atp_synthase | ||
| keq_b6f | ||
| keq_fnr | ||
| vmax_fnr | ||
| keq_PCP700 | ||
| keq_ferredoxin_reductase | ||
| A1 | ||
| vmax_rubisco_carboxylase | ||
| vmax_fbpase | ||
| vmax_SBPase | ||
| vmax_phosphoribulokinase | ||
| vmax_ex_pga | ||
| N_translocator | ||
| vmax_ex_g1p | ||
| B0 | ||
| B1 | ||
| B2 | ||
| B3 | ||
| PQ_ox_div_tot | ||
| Fd_ox_div_tot | ||
| PC_ox_div_tot | ||
| NADPH_div_tot | ||
| ATP_div_tot | ||
| Fluo |
| Symbol | ID | Rate | Stoichiometry |
|---|---|---|---|
| atp_synthase | |||
| b6f | |||
| lhc_protonation | |||
| lhc_deprotonation | |||
| cyclic_electron_flow | |||
| violaxanthin_deepoxidase | |||
| zeaxanthin_epoxidase | |||
| fnr | |||
| ndh | |||
| PSII | |||
| PSI | |||
| proton_leak | |||
| PTOX | |||
| lhc_state_transition_12 | |||
| lhc_state_transition_21 | |||
| rubisco_carboxylase | |||
| phosphoglycerate_kinase | |||
| gadph | |||
| triose_phosphate_isomerase | |||
| aldolase_dhap_gap | |||
| aldolase_dhap_e4p | |||
| fbpase | |||
| transketolase_gap_f6p | |||
| transketolase_gap_s7p | |||
| SBPase | |||
| ribose_phosphate_isomerase | |||
| ribulose_phosphate_epimerase | |||
| phosphoribulokinase | |||
| g6pi | |||
| phosphoglucomutase | |||
| ex_pga | |||
| ex_gap | |||
| ex_dhap | |||
| ex_g1p |
The original model calculates the pH as log10(proton_concentration).
To avoid numerical instabilities, we clamped the pH to the range of 1-14 using log10(clamp(0.1, proton_concentration, 1e-14))..
This model was validated by reproducing the following figures of the original publication.