Photosynthetica X:X | DOI: 10.32615/ps.2025.030
Characterizing the CO2-response curve in photosynthesis and photorespiration: an innovative model for C3 plant species
- 1 Wenzhou Academy of Agricultural Sciences, Wenzhou, Zhejiang, China
- 2 Wenzhou Key Laboratory of Agricultural & Forestry Carbon Sequestration and Tea Resource Development, Wenzhou, Zhejiang, China
- 3 New Quality Productivity Research Center, Guangdong ATV College of Performing Arts, Deqing, Guangdong, China
- 4 Medical College, Guangdong ATV College of Performing Arts, Deqing, Guangdong, China
- 5 School of Life Sciences, Nantong University, Nantong, Jiangsu, China
- 6 State Key Laboratory of Environmental Chemistry and Ecotoxicology, Research Center for Eco-Environmental Sciences, Chinese Academy of Sciences, Beijing, China
- 7 Institute of Biophysics, Math & Physics College, Jinggangshan University, Ji'an, Jiangxi, China
We introduce a novel model for characterizing the CO₂-response curve in photosynthesis, addressing the limitations of the Farquhar-von Caemmerer-Berry (FvCB) model by providing a more comprehensive framework for understanding photosynthetic responses to varying CO2 concentrations in C3 plants. The FvCB model, while instrumental in interpreting the photosynthetic response to CO2, does not directly estimate critical parameters such as maximum net photosynthetic rate, transit point from RuBP- to TPU-limited photosynthesis, and the CO2 compensation point in the presence of day respiration (Rday). Our new model, referred to as Model I, incorporates these parameters and accounts for the Rday, offering a nuanced understanding of plant physiological responses to CO₂ concentrations. The research also developed Model II, which does not require an explicit Rday, addressing the challenges in measuring Rday and providing an alternative for analyzing the CO₂-response curve. Both models were validated against empirical data, demonstrating their effectiveness in studying plant photosynthesis and photorespiration. The study concludes that the new models advance the FvCB model by predicting photosynthetic and photorespiratory responses under varying conditions, which is crucial for agricultural practices and ecosystem management in the context of climate change.
Additional key words: C3 plants; C4 plants; climate change; CO2-response curve; Farquhar-von Caemmerer-Berry model; photosynthesis.
Received: July 5, 2025; Revised: July 5, 2025; Accepted: September 25, 2025; Prepublished online: October 30, 2025
References
- Atkin O.K., Evans J.R., Ball M.C. et al.: Leaf respiration of snow gum in the light and dark. Interactions between temperature and irradiance. - Plant Physiol. 122: 915-924, 2000.
Go to original source... - Baker N.R.: Chlorophyll fluorescence: a probe of photosynthesis in vivo. - Annu. Rev. Plant Biol. 59: 89-113, 2008.
Go to original source... - Barbour M.M., Evans J.R., Simonin K.A., von Caemmerer S.: Online CO2 and H2O oxygen isotope fractionation allows estimation of mesophyll conductance in C4 plants, and reveals that mesophyll conductance decreases as leaves age in both C4 and C3 plants. - New Phytol. 210: 875-889, 2016.
Go to original source... - Bernacchi C.J., Bagley J.E., Serbin S.P. et al.: Modelling C3 photosynthesis from the chloroplast to the ecosystem. - Plant Cell Environ. 36: 1641-1657, 2013.
Go to original source... - Busch F.A., Sage R.F.: The sensitivity of photosynthesis to O2 and CO2 concentration identifies strong Rubisco control above the thermal optimum. - New Phytol. 213: 1036-1051, 2017.
Go to original source... - Chang Y., Latham J., Licht M., Wang L.: A data-driven crop model for maize yield prediction. Commun. Biol. 6: 439, 2023.
Go to original source... - Drake B.G., Gonzàlez-Meler M.A., Long S.P.: More efficient plants: a consequence of rising atmospheric CO₂? - Annu. Rev. Plant Biol. 48: 609-639, 1997.
Go to original source... - Dubois J.-J.B., Fiscus E.L., Booker F.L. et al.: Optimizing the statistical estimation of the parameters of the Farquhar-von Caemmerer-Berry model of photosynthesis. - New Phytol. 176: 402-414, 2007.
Go to original source... - Farazdaghi H., Edwards G.E.: A model for photosynthesis and photorespiration in C3 plants based on the biochemistry and stoichiometry of the pathways. - Plant Cell Environ. 11: 799-809, 1988.
Go to original source... - Farquhar G.D., Busch F.A.: Changes in the chloroplastic CO2 concentration explain much of the observed Kok effect: a model. - New Phytol. 214: 570-584, 2017.
Go to original source... - Farquhar G.D., von Caemmerer S., Berry J.A.: A biochemical model of photosynthetic CO2 assimilation in leaves of C3 species. - Planta 149: 78-90, 1980.
Go to original source... - Fila G., Badeck F.-W., Meyer S. et al.: Relationships between leaf conductance to CO2 diffusion and photosynthesis in micropropagated grapevine plants, before and after ex vitro acclimatization. - J. Exp. Bot. 57: 2687-2695, 2006.
Go to original source... - Flexas J., Díaz-Espejo A., Conesa M.A. et al.: Mesophyll conductance to CO2 and Rubisco as targets for improving intrinsic water use efficiency in C3 plants. - Plant Cell Environ. 39: 965-982, 2016.
Go to original source... - Gregory L.M., McClain A.M., Kramer D.M. et al.: The triose phosphate utilization limitation of photosynthetic rate: out of global models but important for leaf models. - Plant Cell Environ. 44: 3223-3226, 2021.
Go to original source... - Han T., Zhu G., Ma J. et al.: Sensitivity analysis and estimation using a hierarchical Bayesian method for the parameters of the FvCB biochemical photosynthetic model. - Photosynth. Res. 143: 45-66, 2020.
Go to original source... - Harley P.C., Sharkey T.D.: An improved model of C3 photosynthesis at high CO2: reversed O2 sensitivity explained by lack of glycerate reentry into the chloroplast. - Photosynth. Res. 27: 169-178, 1991.
Go to original source... - Hu H., Jiang W., Fan X.: Estimating CO2 response in a mixed broadleaf forest using the dynamic assimilation technique. - BMC Plant Biol. 25: 79, 2025.
Go to original source... - Joshi J., Stocker B.D., Hofhansl F. et al.: Towards a unified theory of plant photosynthesis and hydraulics. - Nat. Plants 8: 1304-1316, 2022.
Go to original source... - Joubert D., Zhang N., Berman S.R. et al.: A small dynamic leaf-level model predicting photosynthesis in greenhouse tomatoes. - PLoS ONE 18: e0275047, 2023.
Go to original source... - Kieffer C., Kaur N., Li J. et al.: Photosynthetic responses of switchgrass to light and CO2 under different precipitation treatments. - Glob. Change Biol.-Bioenergy 16: e13138, 2024.
Go to original source... - Kromdijk J., G³owacka K., Long S.P.: Predicting light-induced stomatal movements based on the redox state of plastoquinone: theory and validation. - Photosynth. Res. 141: 83-97, 2019.
Go to original source... - Lawson T., Oxborough K., Morison J.I.L., Baker N.R.: Responses of photosynthetic electron transport in stomatal guard cells and mesophyll cells in intact leaves to light, CO2, and humidity. - Plant Physiol. 128: 52-62, 2002.
Go to original source... - Leakey A.D.B., Uribelarrea M., Ainsworth E.A. et al.: Photosynthesis, productivity, and yield of maize are not affected by open-air elevation of CO2 concentration in the absence of drought. - Plant Physiol. 140: 779-790, 2006.
Go to original source... - Li Y.L., Liu X.G., Hao K. et al.: Light-response curve of photosynthesis and model fitting in leaves of Mangifera indica under different soil water conditions. - Photosynthetica 57: 796-803, 2019.
Go to original source... - Liu J., Ryu Y., Luo X. et al.: Evidence for widespread thermal acclimation of canopy photosynthesis. - Nat. Plants 10: 1919-1927, 2024.
Go to original source... - Long S.P., Bernacchi C.J.: Gas exchange measurements, what can they tell us about the underlying limitations to photosynthesis? Procedures and sources of error. - J. Exp. Bot. 54: 2393-2401, 2003.
Go to original source... - Norby R.J., Gu L., Haworth I.C. et al.: Informing models through empirical relationships between foliar phosphorus, nitrogen and photosynthesis across diverse woody species in tropical forests of Panama. - New Phytol. 215: 1425-1437, 2017.
Go to original source... - Peisker M.: Models of carbon metabolism in C3-C4 intermediate plants as applied to the evolution of C4 photosynthesis. - Plant Cell Environ. 9: 627-635, 1986.
Go to original source... - Poirier-Pocovi M., Lothier J., Buck-Sorlin G.: Modelling temporal variation of parameters used in two photosynthesis models: influence of fruit load and girdling on leaf photosynthesis in fruit-bearing branches of apple. - Ann. Bot.-London 121: 821-832, 2018.
Go to original source... - Rizza F., Pagani D., Stanca A.M., Cattivelli L.: Use of chlorophyll fluorescence to evaluate the cold acclimation and freezing tolerance of winter and spring oats. - Plant Breeding 120: 389-396, 2001.
Go to original source... - Rogers A., Serbin S.P., Ely K.S. et al.: Terrestrial biosphere models underestimate photosynthetic capacity and CO2 assimilation in the Arctic. - New Phytol. 216: 1090-1103, 2017.
Go to original source... - Sharkey T.D., Bernacchi C.J., Farquhar G.D., Singsaas E.L.: Fitting photosynthetic carbon dioxide response curves for C3 leaves. - Plant Cell Environ. 30: 1035-1040, 2007.
Go to original source... - Silva-Pérez V., Furbank R.T., Condon A.G., Evans J.R.: Biochemical model of C3 photosynthesis applied to wheat at different temperatures. - Plant Cell Environ. 40: 1552-1564, 2017.
Go to original source... - Siqueira J.A., Martins A.O., Wakin T. et al.: The modulation of growth and metabolism in Solanum lycopersicum contrast with the leaf-specific regulation of wild tomato species. - Plant Cell Environ. 48: 1201-1214, 2025.
Go to original source... - Smith N.G., Keenan T.F., Prentice I.C. et al.: Global photosynthetic capacity is optimized to the environment. - Ecol. Lett. 22: 506-517, 2019.
Go to original source... - Sun J., Sun J., Feng Z.: Modelling photosynthesis in flag leaves of winter wheat (Triticum aestivum) considering the variation in photosynthesis parameters during development. - Funct. Plant Biol. 42: 1036-1044, 2015.
Go to original source... - Tcherkez G., Gauthier P., Buckley T.N. et al.: Leaf day respiration: low CO2 flux but high significance for metabolism and carbon balance. - New Phytol. 216: 986-1001, 2017.
Go to original source... - Vialet-Chabrand S.R.M., Matthews J.S.A., McAusland L. et al.: Temporal dynamics of stomatal behavior: modelling and implications for photosynthesis and water use. - Plant Physiol. 174: 603-613, 2017.
Go to original source... - Walker A.P., Quaife T., van Bodegom P.M. et al.: The impact of alternative trait-scaling hypotheses for the maximum photosynthetic carboxylation rate (Vcmax) on global gross primary production. - New Phytol. 215: 1370-1386, 2017.
Go to original source... - Wu Q., Zhang T., Li C.R. et al.: Photosynthetic CO2 response to soil water and its simulation using different models in leaves of two species. - Photosynthetica 58: 790-798, 2020.
Go to original source... - Xue W., Luo H., Carriquí M. et al.: Quantitative expression of mesophyll conductance temperature response in the FvCB model and impacts on plant gas exchange estimations. - Agr. Forest Meteorol. 325: 109153, 2022.
Go to original source... - Ye Z.P., He J.Q., An T. et al.: Influences of residual stomatal conductance on the intrinsic water use efficiency of two C3 and two C4 species. - Agr. Water Manage. 306: 109136, 2024a.
Go to original source... - Ye Z.P., Hu W.H., Zhou S.X. et al.: Limitations of the Farquhar-von Caemmerer-Berry model in estimating the maximum electron transport rate: evidence from four C3 species. - Biology 14: 630, 2025a.
Go to original source... - Ye Z.P., Yang X.L., Ye Z.W.Y. et al.: Evaluating photosynthetic models and their potency in assessing plant responses to changing oxygen concentrations: a comparative analysis of An-Ca and An-Ci curves in Lolium perenne and Triticum aestivum. - Front. Plant Sci. 16: 1575217, 2025b.
Go to original source... - Ye Z.P., Zhou S.X., Yang X.L. et al.: Light curve parametrization of three rice (Oryza sativa L.) cultivars based on mechanistic models. - Photosynthetica 62: 305-313, 2024b.
Go to original source... - Yin X.Y., Amthor J.S.: Estimating leaf day respiration from conventional gas exchange measurements. - New Phytol. 241: 52-58, 2024.
Go to original source... - Yin X.Y., Busch F.A., Struik P.C., Sharkey T.D.: Evolution of a biochemical model of steady-state photosynthesis. - Plant Cell Environ. 44: 2811-2837, 2021.
Go to original source... - Yin X.Y., Struik P.C.: Mathematical review of the energy transduction stoichiometries of C4 leaf photosynthesis under limiting light. - Plant Cell Environ. 35: 1299-1312, 2012.
Go to original source... - Yin X.Y., Struik P.C., Romero P. et al.: Using combined measurements of gas exchange and chlorophyll fluorescence to estimate parameters of a biochemical C3 photosynthesis model: a critical appraisal and a new integrated approach applied to leaves in a wheat (Triticum aestivum) canopy. - Plant Cell Environ. 32: 448-464, 2009.
Go to original source... - Yin X.Y., Sun Z.P., Struik P.C. et al.: Using a biochemical C4 photosynthesis model and combined gas exchange and chlorophyll fluorescence measurements to estimate bundle-sheath conductance of maize leaves differing in age and nitrogen content. - Plant Cell Environ. 34: 2183-2199, 2011.
Go to original source... - Yin X.Y., van Oijen M., Schapendonk A.H.C.M.: Extension of a biochemical model for the generalized stoichiometry of electron transport limited C3 photosynthesis. - Plant Cell Environ. 27: 1211-1222, 2004.
Go to original source... - Zheng Y., Zhao Z., Zhou J.J., Zhou H.: Evaluations of different leaf and canopy photosynthesis models: a case study with black locust (Robinia pseudoacacia) plantations on a Loess Plateau. - Pak. J. Bot. 44: 531-539, 2012.




