1.De Kauwe, M. G. et al. Do land surface models need to include differential plant species responses to drought? Examining model predictions across a mesic-xeric gradient in Europe. Biogeosciences 12, 7503–7518 (2015).Article
Google Scholar
2.Smith, N. G. & Keenan, T. F. Mechanisms underlying leaf photosynthetic acclimation to warming and elevated CO2 as inferred from least‐cost optimality theory. Global Change Biol. 26, 5202–5216 (2020).Article
Google Scholar
3.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).CAS
PubMed
Article
PubMed Central
Google Scholar
4.Wullschleger, S. D. Biochemical limitations to carbon assimilation in C3 plants—a retrospective analysis of the A/Ci curves from 109 Species. J. Exp. Bot. 44, 907–920 (1993).CAS
Article
Google Scholar
5.Lloyd, J., Bloomfield, K., Domingues, T. F. & Farquhar, G. D. Photosynthetically relevant foliar traits correlating better on a mass vs an area basis: of ecophysiological relevance or just a case of mathematical imperatives and statistical quicksand? New Phytol. 199, 311–321 (2013).CAS
PubMed
Article
PubMed Central
Google Scholar
6.De Kauwe, M. G. et al. A test of the ‘one-point method’ for estimating maximum carboxylation capacity from field-measured, light-saturated photosynthesis. New Phytol. 210, 1130–1144 (2015).PubMed
Article
CAS
PubMed Central
Google Scholar
7.Ferreira Domingues, T. et al. Biome-specific effects of nitrogen and phosphorus on the photosynthetic characteristics of trees at a forest-savanna boundary in Cameroon. Oecologia 178, 659–672 (2015).PubMed Central
Article
Google Scholar
8.Domingues, T. F. et al. Co-limitation of photosynthetic capacity by nitrogen and phosphorus in West Africa woodlands. Plant, Cell Environ. 33, 959–980 (2010).CAS
Article
Google Scholar
9.Walker, A. P. et al. The relationship of leaf photosynthetic traits -VcmaxandJmax- to leaf nitrogen, leaf phosphorus, and specific leaf area: a meta-analysis and modeling study. Ecol. Evol. 4, 3218–3235 (2014).PubMed
PubMed Central
Article
Google Scholar
10.Wang, H. et al. Towards a universal model for carbon dioxide uptake by plants. Nat. Plants 3, 734–741 (2017).CAS
PubMed
Article
PubMed Central
Google Scholar
11.Smith, N. G. et al. Global photosynthetic capacity is optimized to the environment. Ecol. Lett. 22, 506–517 (2019).PubMed
PubMed Central
Article
Google Scholar
12.Prentice, I. C., Dong, N., Gleason, S. M., Maire, V. & Wright, I. J. Balancing the costs of carbon gain and water transport: testing a new theoretical framework for plant functional ecology. Ecol. Lett. 17, 82–91 (2014).PubMed
Article
PubMed Central
Google Scholar
13.Givnish, T. J. On the Economy of Plant Form and Function: Proceedings of the Sixth Maria Moors Cabot Symposium, Vol. 6 (Cambridge University Press, 1986).14.Maire, V. et al. Global effects of soil and climate on leaf photosynthetic traits and rates. Glob. Ecol. Biogeogr. 24, 706–717 (2015).Article
Google Scholar
15.Franklin, O. et al. Organizing principles for vegetation dynamics. Nat. Plants 6, 444–453 (2020).PubMed
Article
PubMed Central
Google Scholar
16.Ali, A. A. et al. A global scale mechanistic model of photosynthetic capacity (LUNA V1.0). Geosci. Model Dev. 9, 587–606 (2016).Article
Google Scholar
17.Dewar, R. et al. New insights into the covariation of stomatal, mesophyll and hydraulic conductances from optimization models incorporating nonstomatal limitations to photosynthesis. New Phytol. 217, 571–585 (2017).PubMed
Article
CAS
PubMed Central
Google Scholar
18.Caldararu, S., Thum, T., Yu, L. & Zaehle, S. Whole-plant optimality predicts changes in leaf nitrogen under variable CO 2 and nutrient availability. New Phytol. 225, 2331–2346 (2019).PubMed
Article
CAS
PubMed Central
Google Scholar
19.Wright, I. J. et al. The worldwide leaf economics spectrum. Nature 428, 821–827 (2004).CAS
PubMed
PubMed Central
Article
Google Scholar
20.Wang, H. et al. The China Plant Trait Database: toward a comprehensive regional compilation of functional traits for land plants. Ecology 99, 500–500 (2017).PubMed
Article
PubMed Central
Google Scholar
21.Wang, H. et al. Photosynthetic responses to altitude: an explanation based on optimality principles. New Phytol. 213, 976–982 (2016).PubMed
Article
PubMed Central
Google Scholar
22.Lavergne, A. et al. Historical changes in the stomatal limitation of photosynthesis: empirical support for an optimality principle. New Phytol. 225, 2484–2497 (2019).PubMed
Article
CAS
PubMed Central
Google Scholar
23.Maire, V. et al. The coordination of leaf photosynthesis links C and N fluxes in C3 plant species. PLoS ONE 7, e38345 (2012).CAS
PubMed
PubMed Central
Article
Google Scholar
24.Fürstenau Togashi, H. et al. Thermal acclimation of leaf photosynthetic traits in an evergreen woodland, consistent with the coordination hypothesis. Biogeosciences 15, 3461–3474 (2018).Article
CAS
Google Scholar
25.Kumarathunge, D. P. et al. Acclimation and adaptation components of the temperature dependence of plant photosynthesis at the global scale. New Phytol. 222, 768–784 (2019).CAS
PubMed
Article
PubMed Central
Google Scholar
26.Wright, I. J., Reich, P. B. & Westoby, M. Strategy shifts in leaf physiology, structure and nutrient content between species of high- and low-rainfall and high- and low-nutrient habitats. Funct. Ecol. 15, 423–434 (2001).Article
Google Scholar
27.Rogers, A. The use and misuse of V c,max in earth system models. Photosynth. Res. 119, 15–29 (2013).PubMed
Article
CAS
PubMed Central
Google Scholar
28.Dong, N. et al. Leaf nitrogen from first principles: field evidence for adaptive variation with climate. Biogeosciences 14, 481–495 (2017).CAS
Article
Google Scholar
29.Reich, P. B. & Schoettle, A. W. Role of phosphorus and nitrogen in photosynthetic and whole plant carbon gain and nutrient use efficiency in eastern white pine. Oecologia 77, 25–33 (1988).CAS
PubMed
Article
PubMed Central
Google Scholar
30.Raaimakers, D., Boot, R. G. A., Dijkstra, P. & Pot, S. Photosynthetic rates in relation to leaf phosphorus content in pioneer versus climax tropical rainforest trees. Oecologia 102, 120–125 (1995).CAS
PubMed
Article
PubMed Central
Google Scholar
31.Goll, D. S. et al. Nutrient limitation reduces land carbon uptake in simulations with a model of combined carbon, nitrogen and phosphorus cycling. Biogeosciences 9, 3547–3569 (2012).CAS
Article
Google Scholar
32.Reich, P. B., Oleksyn, J. & Wright, I. J. Leaf phosphorus influences the photosynthesis–nitrogen relation: a cross-biome analysis of 314 species. Oecologia 160, 207–212 (2009).PubMed
Article
PubMed Central
Google Scholar
33.Evans, J. R. Photosynthesis and nitrogen relationships in leaves of C3 plants. Oecologia 78, 9–19 (1989).PubMed
Article
PubMed Central
Google Scholar
34.Reich, P. B., Walters, M. B., Ellsworth, D. S. & Uhl, C. Photosynthesis-nitrogen relations in Amazonian tree species. Oecologia 97, 62–72 (1994).CAS
PubMed
Article
PubMed Central
Google Scholar
35.Kattge, J., Knorr, W., Raddatz, T. & Wirth, C. Quantifying photosynthetic capacity and its relationship to leaf nitrogen content for global-scale terrestrial biosphere models. Global Change Biol. 15, 976–991 (2009).Article
Google Scholar
36.Evans, J. R. & Clarke, V. C. The nitrogen cost of photosynthesis. J. Exp. Bot. 70, 7–15 (2018).Article
CAS
Google Scholar
37.Marschner, H. in Mineral Nutrition of Higher Plants, 405–435 (Elsevier, 1995).38.Niinemets, Ü., Wright, I. J. & Evans, J. R. Leaf mesophyll diffusion conductance in 35 Australian sclerophylls covering a broad range of foliage structural and physiological variation. J. Exp. Bot. 60, 2433–2449 (2009).CAS
PubMed
Article
PubMed Central
Google Scholar
39.Malhi, Y. et al. The variation of productivity and its allocation along a tropical elevation gradient: a whole carbon budget perspective. New Phytol. 214, 1019–1032 (2016).PubMed
Article
CAS
PubMed Central
Google Scholar
40.Wang, H. et al. Acclimation of leaf respiration consistent with optimal photosynthetic capacity. Global Change Biol. 26, 2573–2583 (2020).Article
Google Scholar
41.Peng, Y., Bloomfield, K. J. & Prentice, I. C. A theory of plant function helps to explain leaf-trait and productivity responses to elevation. New Phytol. 226, 1274–1284, (2020).CAS
PubMed
Article
PubMed Central
Google Scholar
42.Gvozdevaite, A. et al. Leaf-level photosynthetic capacity dynamics in relation to soil and foliar nutrients along forest–savanna boundaries in Ghana and Brazil. Tree Physiol. 38, 1912–1925 (2018).CAS
PubMed
Article
PubMed Central
Google Scholar
43.Terrer, C. et al. Nitrogen and phosphorus constrain the CO 2 fertilization of global plant biomass. Nat. Clim. Change 9, 684–689 (2019).CAS
Article
Google Scholar
44.Meir, P. et al. in Advances in Photosynthesis and Respiration, 89–105 (Springer International Publishing, 2017).45.Luo, X. & Keenan, T. F. Global evidence for the acclimation of ecosystem photosynthesis to light. Nat. Ecol. Evol. 4, 1351–1357 (2020).PubMed
Article
PubMed Central
Google Scholar
46.Stocker, B. D. et al. P-model v1.0: an optimality-based light use efficiency model for simulating ecosystem gross primary production. Geosci. Model Dev. 13, 1545–1581 (2020).Article
Google Scholar
47.Lavergne, A., Sandoval, D., Hare, V. J., Graven, H. & Prentice, I. C. Impacts of soil water stress on the acclimated stomatal limitation of photosynthesis: insights from stable carbon isotope data. Global Change Biol. 26, 7158–7172 (2020).48.Zhou, S., Duursma, R. A., Medlyn, B. E., Kelly, J. W. G. & Prentice, I. C. How should we model plant responses to drought? An analysis of stomatal and non-stomatal responses to water stress. Agric. For. Meteorol. 182-183, 204–214 (2013).Article
Google Scholar
49.Zhou, S. et al. Short-term water stress impacts on stomatal, mesophyll and biochemical limitations to photosynthesis differ consistently among tree species from contrasting climates. Tree Physiol. 34, 1035–1046 (2014).CAS
PubMed
Article
PubMed Central
Google Scholar
50.Katul, G., Manzoni, S., Palmroth, S. & Oren, R. A stomatal optimization theory to describe the effects of atmospheric CO2 on leaf photosynthesis and transpiration. Ann. Bot. 105, 431–442 (2009).PubMed
PubMed Central
Article
Google Scholar
51.Manzoni, S. et al. Optimizing stomatal conductance for maximum carbon gain under water stress: a meta-analysis across plant functional types and climates. Funct. Ecol. 25, 456–467 (2011).Article
Google Scholar
52.Medlyn, B. E. et al. Reconciling the optimal and empirical approaches to modelling stomatal conductance. Global Change Biol. 17, 2134–2144 (2011).Article
Google Scholar
53.Crous, K. Y. et al. Photosynthesis of temperate Eucalyptus globulus trees outside their native range has limited adjustment to elevated CO2 and climate warming. Global Change Biol. 19, 3790–3807 (2013).Article
Google Scholar
54.Zhou, S.-X., Medlyn, B. E. & Prentice, I. C. Long-term water stress leads to acclimation of drought sensitivity of photosynthetic capacity in xeric but not riparian Eucalyptus species. Ann. Bot. 117, 133–144 (2015).PubMed
PubMed Central
Article
CAS
Google Scholar
55.Smith, N. G. & Dukes, J. S. Short-term acclimation to warmer temperatures accelerates leaf carbon exchange processes across plant types. Global Change Biol. 23, 4840–4853 (2017).Article
Google Scholar
56.Katul, G., Leuning, R. & Oren, R. Relationship between plant hydraulic and biochemical properties derived from a steady‐state coupled water and carbon transport model. Plant, Cell Environ. 26, 339–350 (2003).CAS
Article
Google Scholar
57.Stocker, B. D. et al. Quantifying soil moisture impacts on light use efficiency across biomes. New Phytol. 218, 1430–1449 (2018).PubMed
PubMed Central
Article
Google Scholar
58.Kattge, J. & Knorr, W. Temperature acclimation in a biochemical model of photosynthesis: a reanalysis of data from 36 species. Plant, Cell Environ. 30, 1176–1190 (2007).CAS
Article
Google Scholar
59.van der Plas, F. et al. Plant traits alone are poor predictors of ecosystem properties and long-term ecosystem functioning. Nat. Ecol. Evol. 4, 1602–1611 (2020).PubMed
Article
PubMed Central
Google Scholar
60.Quesada, M. et al. Succession and management of tropical dry forests in the Americas: review and new perspectives. For. Ecol. Manag. 258, 1014–1024 (2009).Article
Google Scholar
61.Phillips, O. L. et al. Drought–mortality relationships for tropical forests. New Phytol. 187, 631–646 (2010).PubMed
Article
PubMed Central
Google Scholar
62.Laliberté, E., Lambers, H., Burgess, T. I. & Wright, S. J. Phosphorus limitation, soil-borne pathogens and the coexistence of plant species in hyperdiverse forests and shrublands. New Phytol. 206, 507–521 (2014).PubMed
Article
CAS
PubMed Central
Google Scholar
63.Conroy, J. P., Smillie, R. M., Küppers, M., Bevege, D. I. & Barlow, E. W. Chlorophyll a fluorescence and photosynthetic and growth responses of pinus radiata to phosphorus deficiency, drought stress, and high CO2. Plant Physiol. 81, 423–429 (1986).CAS
PubMed
PubMed Central
Article
Google Scholar
64.Loustau, D., Brahim, M. B., Gaudillere, J. P. & Dreyer, E. Photosynthetic responses to phosphorus nutrition in two-year-old maritime pine seedlings. Tree Physiol. 19, 707–715 (1999).PubMed
Article
PubMed Central
Google Scholar
65.Warren, C. R. & Adams, M. A. Phosphorus affects growth and partitioning of nitrogen to Rubisco in Pinus pinaster. Tree Physiol. 22, 11–19 (2002).CAS
PubMed
Article
PubMed Central
Google Scholar
66.Bloomfield, K. J., Farquhar, G. D. & Lloyd, J. Photosynthesis–nitrogen relationships in tropical forest tree species as affected by soil phosphorus availability: a controlled environment study. Funct. Plant Biol. 41, 820–832 (2014).CAS
PubMed
Article
PubMed Central
Google Scholar
67.Crous, K. Y., Ósvaldsson, A. & Ellsworth, D. S. Is phosphorus limiting in a mature Eucalyptus woodland? Phosphorus fertilisation stimulates stem growth. Plant Soil 391, 293–305 (2015).CAS
Article
Google Scholar
68.Sivak, M. N. & Walker, D. A. Photosynthesis in vivo can be limited by phosphate supplY. New Phytol. 102, 499–512 (1986).CAS
Article
Google Scholar
69.Kiirats, O., Cruz, J. A., Edwards, G. E. & Kramer, D. M. Feedback limitation of photosynthesis at high CO2 acts by modulating the activity of the chloroplast ATP synthase. Funct. Plant Biol. 36, 893–901 (2009).CAS
PubMed
Article
PubMed Central
Google Scholar
70.Ellsworth, D. S., Crous, K. Y., Lambers, H. & Cooke, J. Phosphorus recycling in photorespiration maintains high photosynthetic capacity in woody species. Plant, Cell Environ. 38, 1142–1156 (2015).CAS
Article
Google Scholar
71.Zhang, S. & Dang, Q. L. Effects of carbon dioxide concentration and nutrition on photosynthetic functions of white birch seedlings. Tree Physiol. 26, 1457–1467 (2006).CAS
PubMed
Article
PubMed Central
Google Scholar
72.Lambers, H. et al. Proteaceae from severely phosphorus-impoverished soils extensively replace phospholipids with galactolipids and sulfolipids during leaf development to achieve a high photosynthetic phosphorus-use-efficiency. New Phytol. 196, 1098–1108 (2012).CAS
PubMed
Article
PubMed Central
Google Scholar
73.Meir, P., Levy, P. E., Grace, J. & Jarvis, P. G. Photosynthetic parameters from two contrasting woody vegetation types in West Africa. Plant Ecol. 192, 277–287 (2007).Article
Google Scholar
74.Kull, O. Acclimation of photosynthesis in canopies: models and limitations. Oecologia 133, 267–279 (2002).PubMed
Article
PubMed Central
Google Scholar
75.Field, C. & Mooney, H. in On the Economy of Plant Form and Function: Proceedings of the Sixth Maria Moors Cabot Symposium, Evolutionary Constraints on Primary Productivity, Adaptive Patterns of Energy Capture in Plants, Harvard Forest, August 1983 (Cambridge University Press, 1986).76.Niinemets, Ü. Research review. Components of leaf dry mass per area – thickness and density – alter leaf photosynthetic capacity in reverse directions in woody plants. New Phytol. 144, 35–47 (1999).Article
Google Scholar
77.Lloyd, J. et al. Optimisation of photosynthetic carbon gain and within-canopy gradients of associated foliar traits for Amazon forest trees. Biogeosciences 7, 1833–1859 (2010).CAS
Article
Google Scholar
78.Anten, N. P. R. Optimal photosynthetic characteristics of individual plants in vegetation stands and implications for species coexistence. Ann. Bot. 95, 495–506 (2004).PubMed
PubMed Central
Article
CAS
Google Scholar
79.Alton, P. B. & North, P. Interpreting shallow, vertical nitrogen profiles in tree crowns: a three-dimensional, radiative-transfer simulation accounting for diffuse sunlight. Agric. For. Meteorol. 145, 110–124 (2007).Article
Google Scholar
80.Rogers, A. et al. A roadmap for improving the representation of photosynthesis in Earth system models. New Phytol. 213, 22–42 (2017).PubMed
Article
PubMed Central
Google Scholar
81.Tosens, T. & Laanisto, L. Mesophyll conductance and accurate photosynthetic carbon gain calculations. J. Exp. Bot. 69, 5315–5318 (2018).CAS
PubMed
Article
PubMed Central
Google Scholar
82.Niinemets, Ü., Díaz-Espejo, A., Flexas, J., Galmés, J. & Warren, C. R. Importance of mesophyll diffusion conductance in estimation of plant photosynthesis in the field. J. Exp. Bot. 60, 2271–2282 (2009).CAS
PubMed
Article
PubMed Central
Google Scholar
83.Farquhar, G. D., O’Leary, M. H. & Berry, J. A. On the relationship between carbon isotope discrimination and the intercellular carbon dioxide concentration in leaves. Funct. Plant Biol. 9, 121–137 (1982).CAS
Article
Google Scholar
84.Bernacchi, C. J., Singsaas, E. L., Pimentel, C., Portis, A. R. Jr & Long, S. P. Improved temperature response functions for models of Rubisco-limited photosynthesis. Plant, Cell Environ. 24, 253–259 (2001).CAS
Article
Google Scholar
85.Bernacchi, C. J., Pimentel, C. & Long, S. P. In vivo temperature response functions of parameters required to model RuBP-limited photosynthesis. Plant, Cell Environ. 26, 1419–1430 (2003).CAS
Article
Google Scholar
86.Scafaro, A. P. et al. Strong thermal acclimation of photosynthesis in tropical and temperate wet-forest tree species: the importance of altered Rubisco content. Global Change Biol. 23, 2783–2800 (2017).Article
Google Scholar
87.Warton, D. I., Wright, I. J., Falster, D. S. & Westoby, M. Bivariate line-fitting methods for allometry. Biol. Rev. 81, 259–291 (2006).PubMed
Article
PubMed Central
Google Scholar
88.Team, R. C. R.: a Language and Environment for Statistical Computing (R Foundation for Statistical Computing, 2018).89.Atkin, O. K. et al. Global variability in leaf respiration in relation to climate, plant functional types and leaf traits. New Phytol. 206, 614–636 (2015).CAS
PubMed
Article
PubMed Central
Google Scholar
90.Bahar, N. H. A. et al. Leaf-level photosynthetic capacity in lowland Amazonian and high-elevation Andean tropical moist forests of Peru. New Phytol. 214, 1002–1018 (2016).PubMed
Article
CAS
PubMed Central
Google Scholar
91.Bloomfield, K. J. et al. The validity of optimal leaf traits modelled on environmental conditions. New Phytol. 221, 1409–1423 (2018).PubMed
Article
CAS
PubMed Central
Google Scholar
92.Cernusak, L. A., Hutley, L. B., Beringer, J., Holtum, J. A. M. & Turner, B. L. Photosynthetic physiology of eucalypts along a sub-continental rainfall gradient in northern Australia. Agric. For. Meteorol. 151, 1462–1470 (2011).Article
Google Scholar
93.Xu, H. Y., et al. Predictability of leaf traits with climate and elevation: a case study in Gongga Mountain, China. Tree Physiol. https://doi.org/10.1093/treephys/tpab003 (2021).94.Walker, A. P., et al. A Global Data Set of Leaf Photosynthetic Rates, Leaf N and P, and Specific Leaf Area (Oak Ridge National Laboratory Distributed Active Archive Center, 2014). https://doi.org/10.3334/ORNLDAAC/1224.95.Kattge, J. et al. TRY–a global database of plant traits. Global Change Biol. 17, 2905–2935 (2011).Article
Google Scholar
96.Collatz, G. J., Ribas-Carbo, M. & Berry, J. A. Coupled photosynthesis-stomatal conductance model for leaves of C4 plants. Funct. Plant Biol. 19, 519 (1992).Article
Google Scholar
97.Rogers, A., Serbin, S. P., Ely, K. S., Sloan, V. L. & Wullschleger, S. D. Terrestrial biosphere models underestimate photosynthetic capacity and CO2 assimilation in the Arctic. New Phytol. 216, 1090–1103 (2017).CAS
PubMed
Article
PubMed Central
Google Scholar
98.Burnett, A. C., Davidson, K. J., Serbin, S. P. & Rogers, A. The “one‐point method” for estimating maximum carboxylation capacity of photosynthesis: a cautionary tale. Plant, Cell Environ. 42, 2472–2481 (2019).CAS
Article
Google Scholar
99.Harris, I., Jones, P. D., Osborn, T. J. & Lister, D. H. Updated high-resolution grids of monthly climatic observations–the CRU TS3.10 Dataset. Int. J. Climatol. 34, 623–642 (2013).Article
Google Scholar
100.Jones, H. G. Plants and Microclimate (Cambridge University Press, 2009).101.Weedon, G. P. et al. The WFDEI meteorological forcing data set: WATCH forcing data methodology applied to ERA-interim reanalysis data. Water Resour. Res. 50, 7505–7514 (2014).Article
Google Scholar
102.Davis, T. W. et al. Simple process-led algorithms for simulating habitats (SPLASH v.1.0): robust indices of radiation, evapotranspiration and plant-available moisture. Geosci. Model Dev. 10, 689–708 (2017).Article
Google Scholar
103.Berberan-Santos, M. N., Bodunov, E. N. & Pogliani, L. On the barometric formula. Am. J. Phys. 65, 404–412 (1997).Article
Google Scholar
104.Peng, Y., et al. Dataset of Global Climate and Nutrient Controls of Photosynthetic Capacity (Zenodo, 2021). https://doi.org/10.5281/zenodo.4568148. More