[1]

Lynch J, Cain M, Frame D, Pierrehumbert R. 2021. Agriculture's contribution to climate change and role in mitigation is distinct from predominantly fossil CO2-emitting sectors. Frontiers in Sustainable Food Systems 4:518039

doi: 10.3389/fsufs.2020.518039
[2]

Intergovernmental Panel on Climate Change (IPCC). 2023. Summary for Policymakers. Climate Change 2023: Synthesis Report. IPCC, Geneva, Switzerland. doi: 10.59327/ipcc/ar6-9789291691647.001

[3]

Niu S, Wu M, Han Y, Xia J, Li L, et al. 2008. Water‐mediated responses of ecosystem carbon fluxes to climatic change in a temperate steppe. New Phytologist 177:209−219

doi: 10.1111/j.1469-8137.2007.02237.x
[4]

Yu G, Song X, Wang Q, Liu Y, Guan D, et al. 2008. Water-use efficiency of forest ecosystems in Eastern China and its relations to climatic variables. New Phytologist 177:927−937

doi: 10.1111/j.1469-8137.2007.02316.x
[5]

Lesk C, Rowhani P, Ramankutty N. 2016. Influence of extreme weather disasters on global crop production. Nature 529:84−87

doi: 10.1038/nature16467
[6]

Qaseem MF, Qureshi R, Shaheen H. 2019. Effects of pre-anthesis drought, heat and their combination on the growth, yield and physiology of diverse wheat (Triticum aestivum L.) genotypes varying in sensitivity to heat and drought stress. Scientific Reports 9:6955

doi: 10.1038/s41598-019-43477-z
[7]

Anderegg WRL, Berry JA, Smith DD, Sperry JS, Anderegg LDL, et al. 2012. The roles of hydraulic and carbon stress in a widespread climate-induced forest die-off. Proceedings of the National Academy of Sciences of the United States of America 109:233−237

doi: 10.1073/pnas.1107891109
[8]

Cowan I. 2002. Fit, fitter, fittest; where does optimisation fit in? Silva Fennica 36:745−754

doi: 10.14214/sf.536
[9]

Gilbert ME, Hernandez MI. 2019. How should crop water-use efficiency be analyzed? A warning about spurious correlations. Field Crops Research 235:59−67

doi: 10.1016/j.fcr.2019.02.017
[10]

Niu S, Xing X, Zhang Z, Xia J, Zhou X, et al. 2011. Water-use efficiency in response to climate change: from leaf to ecosystem in a temperate steppe. Global Change Biology 17:1073−1082

doi: 10.1111/j.1365-2486.2010.02280.x
[11]

Shao X, Chen S, Chen Y, Yang Z, Liu Q, et al. 2024. A review of the relationship between plant water use efficiency and plant functional community structure. World Forestry Research 37:37−44 (in Chinese)

doi: 10.13348/j.cnki.sjlyyj.2024.0003.y
[12]

Tang J, Bolstad PV, Ewers BE, Desai AR, Davis KJ, et al. 2006. Sap flux-upscaled canopy transpiration, stomatal conductance, and water use efficiency in an old growth forest in the Great Lakes region of the United States. Journal of Geophysical Research: Biogeosciences 111:G02009

doi: 10.1029/2005jg000083
[13]

Wang W, Xiao J, Ollinger SV, Desai AR, Chen J, et al. 2014. Quantifying the effects of harvesting on carbon fluxes and stocks in northern temperate forests. Biogeosciences 11:6667−6682

doi: 10.5194/bg-11-6667-2014
[14]

Schleser GH. 1990. Investigations of the δ13C pattern in leaves of Fagus sylvatica L. Journal of Experimental Botany 41:565−572

doi: 10.1093/jxb/41.5.565
[15]

Quan Q, Zhang F, Meng C, Ma F, Zhou Q, et al. 2020. Shifting biomass allocation determines community water use efficiency under climate warming. Environmental Research Letters 15:094041

doi: 10.1088/1748-9326/aba472
[16]

Loader NJ, Switsur VR, Field EM. 1995. High-resolution stable isotope analysis of tree rings: implications of microdendroclimatology for palaeoenvironmental research. The Holocene 5:457−460

doi: 10.1177/095968369500500408
[17]

Xue BL, Guo Q, Otto A, Xiao J, Tao S, et al. 2015. Global patterns, trends, and drivers of water use efficiency from 2000 to 2013. Ecosphere 6:1−18

doi: 10.1890/es14-00416.1
[18]

Nicotra AB, Cosgrove MJ, Cowling A, Schlichting CD, Jones CS. 2008. Leaf shape linked to photosynthetic rates and temperature optima in South African Pelargonium species. Oecologia 154:625−635

doi: 10.1007/s00442-007-0865-1
[19]

Purdy AJ, Fisher JB, Goulden ML, Colliander A, Halverson G, et al. 2018. SMAP soil moisture improves global evapotranspiration. Remote Sensing of Environment 219:1−14

doi: 10.1016/j.rse.2018.09.023
[20]

Liu X, Feng X, Fu B. 2020. Changes in global terrestrial ecosystem water use efficiency are closely related to soil moisture. Science of the Total Environment 698:134165

doi: 10.1016/j.scitotenv.2019.134165
[21]

Hao X, Zhang J, Fan X, Hao H, Li Y. 2021. Quantifying soil moisture impacts on water use efficiency in terrestrial ecosystems of China. Remote Sensing 13:4257

doi: 10.3390/rs13214257
[22]

Baker NR. 2008. Chlorophyll fluorescence: a probe of photosynthesis in vivo. Annual Review of Plant Biology 59:89−113

doi: 10.1146/annurev.arplant.59.032607.092759
[23]

Baret F, de Solan B, Lopez-Lozano R, Ma K, Weiss M. 2010. GAI estimates of row crops from downward looking digital photos taken perpendicular to rows at 57.5° zenith angle: theoretical considerations based on 3D architecture models and application to wheat crops. Agricultural and Forest Meteorology 150:1393−1401

doi: 10.1016/j.agrformet.2010.04.011
[24]

Zhang F, Ju W, Shen S, Wang S, Yu G, et al. 2014. How recent climate change influences water use efficiency in East Asia. Theoretical and Applied Climatology 116:359−370

doi: 10.1007/s00704-013-0949-2
[25]

Li W, Liu M, Li M, Sun R, Zhou T, et al. 2024. Influence of nitrogen water interaction on leaf functional traits of dominant species in warm temperate forest. Forestry Research 4:e009

doi: 10.48130/forres-0024-0006
[26]

Xiao B, Bai X, Zhao C, Tan Q, Li Y, et al. 2023. Responses of carbon and water use efficiencies to climate and land use changes in China's karst areas. Journal of Hydrology 617:128968

doi: 10.1016/j.jhydrol.2022.128968
[27]

Xiao J, Sun G, Chen J, Chen H, Chen S, et al. 2013. Carbon fluxes, evapotranspiration, and water use efficiency of terrestrial ecosystems in China. Agricultural and Forest Meteorology 182−183:76−90

doi: 10.1016/j.agrformet.2013.08.007
[28]

Canadell J, Jackson RB, Ehleringer JB, Mooney HA, Sala OE, et al. 1996. Maximum rooting depth of vegetation types at the global scale. Oecologia 108:583−595

doi: 10.1007/BF00329030
[29]

Wright IJ, Reich PB, Westoby M, Ackerly DD, Baruch Z, et al. 2004. The worldwide leaf economics spectrum. Nature 428:821−827

doi: 10.1038/nature02403
[30]

Craufurd PQ, Wheeler TR, Ellis RH, Summerfield RJ, Williams JH. 1999. Effect of temperature and water deficit on water-use efficiency, carbon isotope discrimination, and specific leaf area in peanut. Crop Science 39:136−142

doi: 10.2135/cropsci1999.0011183x003900010022x
[31]

Querejeta JI, Ren W, Prieto I. 2021. Vertical decoupling of soil nutrients and water under climate warming reduces plant cumulative nutrient uptake, water-use efficiency and productivity. New Phytologist 230:1378−1393

doi: 10.1111/nph.17258
[32]

Rose MT, Patti AF, Little KR, Brown AL, Jackson WR, et al. 2014. A meta-analysis and review of plant-growth response to humic substances: practical implications for agriculture. Advances in Agronomy 124:37−89

doi: 10.1016/b978-0-12-800138-7.00002-4
[33]

Hedges LV, Gurevitch J, Curtis PS. 1999. The meta-analysis of response ratios in experimental ecology. Ecology 80:1150−1156

doi: 10.1890/0012-9658(1999)080[1150:TMAORR]2.0.CO;2
[34]

Veroniki AA, Jackson D, Viechtbauer W, Bender R, Bowden J, et al. 2016. Methods to estimate the between-study variance and its uncertainty in meta-analysis. Research Synthesis Methods 7:55−79

doi: 10.1002/jrsm.1164
[35]

R Core Team. 2026. R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. www.R-project.org/

[36]

Song F, Eastwood AJ, Gilbody S, Duley L, Sutton AJ. 2000. Publication and related biases: a review. Health Technology Assessment 4:1−115

doi: 10.3310/hta4100
[37]

Viechtbauer W. 2010. Conducting meta-analyses in r with the metafor package. Journal of Statistical Software 36:1−48

doi: 10.18637/jss.v036.i03
[38]

Zhang J, Deng L, Jiang H, Peng C, Huang C, et al. 2021. The effects of elevated CO2, elevated O3, elevated temperature, and drought on plant leaf gas exchanges: a global meta-analysis of experimental studies. Environmental Science and Pollution Research 28:15274−15289

doi: 10.1007/s11356-020-11728-6
[39]

Streit K, Siegwolf RTW, Hagedorn F, Schaub M, Buchmann N. 2014. Lack of photosynthetic or stomatal regulation after 9 years of elevated [CO2] and 4 years of soil warming in two conifer species at the alpine treeline. Plant, Cell & Environment 37:315−326

doi: 10.1111/pce.12197
[40]

Li S, Li X, Wei Z, Liu F. 2020. ABA-mediated modulation of elevated CO2 on stomatal response to drought. Current Opinion in Plant Biology 56:174−180

doi: 10.1016/j.pbi.2019.12.002
[41]

Wang D, Wang H, Wang P, Ling T, Tao W, et al. 2019. Warming treatment methodology affected the response of plant ecophysiological traits to temperature increases: a quantitive meta-analysis. Frontiers in Plant Science 10:957

doi: 10.3389/fpls.2019.00957
[42]

Li Y, Liu J, Zhou G, Huang W, Duan H. 2016. Warming effects on photosynthesis of subtropical tree species: a translocation experiment along an altitudinal gradient. Scientific Reports 6:24895

doi: 10.1038/srep24895
[43]

Dong T, Liu J, He P, Shi M, Chi Y, et al. 2024. Time lag and cumulative effects of extreme climate on coastal vegetation in China. Remote Sensing 16:528

doi: 10.3390/rs16030528
[44]

Sun ZP, Li TL, Liu YL. 2011. Effects of elevated CO2 applied to potato roots on the anatomy and ultrastructure of leaves. Biologia Plantarum 55:675−680

doi: 10.1007/s10535-011-0167-7
[45]

Maire V, Wright IJ, Prentice IC, Batjes NH, Bhaskar R, et al. 2015. Global effects of soil and climate on leaf photosynthetic traits and rates. Global Ecology and Biogeography 24:706−717

doi: 10.1111/geb.12296
[46]

Rosbakh S, Auerswald K, Poschlod P. 2021. Rising CO2 concentrations reduce nitrogen availability in alpine grasslands. Ecological Indicators 129:107990

doi: 10.1016/j.ecolind.2021.107990
[47]

Krämer K, Kepp G, Brock J, Stutz S, Heyer AG. 2022. Acclimation to elevated CO2 affects the C/N balance by reducing de novo N-assimilation. Physiologia Plantarum 174:e13615

doi: 10.1111/ppl.13615
[48]

Perkowski EA, Ezekannagha E, Smith NG. 2025. Nitrogen demand, availability, and acquisition strategy control plant responses to elevated CO2. Journal of Experimental Botany 76:2908−2923

doi: 10.1093/jxb/eraf118
[49]

Yang K, Huang Y, Yang J, Yu L, Hu Z, et al. 2023. The determiner of photosynthetic acclimation induced by biochemical limitation under elevated CO2 in Japonica rice. Journal of Plant Physiology 280:153889

doi: 10.1016/j.jplph.2022.153889
[50]

Babenko LM, Futorna OA, Akimov YA, Romanenko KO, Kosakivska IV, et al. 2024. How short-term temperature stresses affect leaf micromorphology and ultrastructure of mesophyll cells in winter rye Secale cereale L. Acta Physiologiae Plantarum 46:111

doi: 10.1007/s11738-024-03743-8
[51]

Cao L, Chen Y, Xiao K, Chen L. 2025. FaNAC047-FaNAC058 module coordinately promotes chlorophyll degradation and reactive oxygen species production during heat-induced leaf senescence in tall fescue. Journal of Integrative Plant Biology 67:1009−1027

doi: 10.1111/jipb.13897
[52]

Xu C, Wang M, Yang Z, Han W, Zheng S. 2021. Effects of high temperature on photosynthetic physiological characteristics of strawberry seedlings in greenhouse and construction of stress level. Chinese Journal Applied Ecology 32:231−240 (in Chinese)

doi: 10.13287/j.1001-9332.202101.028
[53]

Wu J, Xu G, Li H, Zeng X, Jiang J, et al. 2021. Effects of heat stress on chlorophyll fluorescence and photosynthetic characteristic parameters in grape (Vitisvinifera L. 'Manicure finger'). Xinjiang Agricultural Sciences 58:2274−2281

doi: 10.6048/j.issn.1001-4330.2021.12.015
[54]

White PJ, Broadley MR. 2003. Calcium in plants. Annals of Botany 92:487−511

doi: 10.1093/aob/mcg164
[55]

Zhou N, Li H, Wang B, Rengel Z, Li H. 2024. Differential root nutrient-acquisition strategies underlie biogeochemical niche separation between grasses and forbs across grassland biomes. Functional Ecology 38:2286−2299

doi: 10.1111/1365-2435.14629
[56]

Yan W, Zhong Y, Shangguan, Z. 2017. Contrasting responses of leaf stomatal characteristics to climate change: a considerable challenge to predict carbon and water cycles. Global Change Biology 23:3781−3793

doi: 10.1111/gcb.13654
[57]

Habermann E, Dias de Oliveira EA, Contin DR, San Martin JAB, Curtarelli L, et al. 2019. Stomatal development and conductance of a tropical forage legume are regulated by elevated [CO2] under moderate warming. Frontiers in Plant Science 10:609

doi: 10.3389/fpls.2019.00609
[58]

Dai M, Wang T, Wang Y, Xu J. 2022. Effects of warming and phosphorus enrichment on the C: N: P stoichiometry of Potamogeton crispus organs. Frontiers in Plant Science 13:814255

doi: 10.3389/fpls.2022.814255
[59]

Ji Y, Zeng S, Tang Q, Yan L, Wu S, et al. 2023. Spatiotemporal variations and driving factors of China's ecosystem water use efficiency. Ecological Indicators 148:110077

doi: 10.1016/j.ecolind.2023.110077
[60]

Li F, Xiao J, Chen J, Ballantyne A, Jin K, et al. 2023. Global water use efficiency saturation due to increased vapor pressure deficit. Science 381:672−677

doi: 10.1126/science.adf5041
[61]

Hsu PK, Takahashi Y, Merilo E, Costa A, Zhang L, et al. 2021. Raf-like kinases and receptor-like (pseudo) kinase GHR1 are required for stomatal vapor pressure difference response. Proceedings of the National Academy of Sciences of the United States of America 118:e2107280118

doi: 10.1073/pnas.2107280118
[62]

Tu Y, Wang X, Zhou J, Wang X, Jia Z, et al. 2024. Atmospheric water demand dominates terrestrial ecosystem productivity in China. Agricultural and Forest Meteorology 355:110151

doi: 10.1016/j.agrformet.2024.110151
[63]

Li B, Zhao X, Geng J, Zhang S, Li J. 2026. SlTIP2;3 mediates H2O2 transport to activate GA signaling and maintain stomatal conductance under high vapor pressure deficit. The Plant Journal 125:e70705

doi: 10.1111/tpj.70705
[64]

Denissen JMC, Teuling AJ, Pitman AJ, Koirala S, Migliavacca M, et al. 2022. Widespread shift from ecosystem energy to water limitation with climate change. Nature Climate Change 12:677−684

doi: 10.1038/s41558-022-01403-8
[65]

Xu X, Jiao F, Gong H, Xue P, Lin N, et al. 2023. Observed divergence in the trends of temperature controls on Chinese ecosystem water use efficiency. Ecological Indicators 157:111241

doi: 10.1016/j.ecolind.2023.111241
[66]

Bai J, Zhai D, Xu Y, Chen D, Wang W, et al. 2026. Altitude-adaptive water use strategies of grassland are constrained by air dryness and stoichiometry in southwest of China. Frontiers in Plant Science 17:1773262

doi: 10.3389/fpls.2026.1773262
[67]

Tissink M, Radolinski J, Reinthaler D, Venier S, Pötsch EM, et al. 2025. Individual versus combined effects of warming, elevated CO2 and drought on grassland water uptake and fine root traits. Plant, Cell & Environment 48:2083−2098

doi: 10.1111/pce.15274
[68]

Mathias JM, Thomas RB. 2021. Global tree intrinsic water use efficiency is enhanced by increased atmospheric CO2 and modulated by climate and plant functional types. Proceedings of the National Academy of Sciences of the United States of America 118:e2014286118

doi: 10.1073/pnas.2014286118
[69]

Wang X, Fu Z, Ciais P, Wang L, Buchmann N, et al. 2026. Global distribution and changes of leaf-level intrinsic water use efficiency and their responses to water stress. Nature Communications 17:1530

doi: 10.1038/s41467-025-68252-9
[70]

Liu L, Zhang X, Song B, Xiao Y, Zhuang W. 2026. Oasis or trap: divergent survival strategies of two desert herbs under shrub fertile islands. BMC Plant Biology 26:346

doi: 10.1186/s12870-026-08160-2
[71]

Sugiura D, Wang Y, Kono M, Mizokami Y. 2024. Exploring the responses of crop photosynthesis to CO2 elevation at the molecular, physiological, and morphological levels toward increasing crop production. Crop and Environment 3:75−83

doi: 10.1016/j.crope.2023.11.006
[72]

Smith MD, Wilkins KD, Holdrege MC, Wilfahrt P, Collins SL, et al. 2024. Extreme drought impacts have been underestimated in grasslands and shrublands globally. Proceedings of the National Academy of Sciences of the United States of America 121:e2309881120

doi: 10.1073/pnas.2309881120
[73]

Herberich MM, Schädle JE, Tielbörger K. 2023. Plant community productivity and soil water are not resistant to extreme experimental drought in temperate grasslands but in the understory of temperate forests. Science of the Total Environment 891:164625

doi: 10.1016/j.scitotenv.2023.164625
[74]

Liu L, Xia H, Quan X, Wang Y. 2023. Plant trait-based life strategies of overlapping species vary in different succession stages of subtropical forests, Eastern China. Frontiers in Ecology and Evolution 10:1103937

doi: 10.3389/fevo.2022.1103937
[75]

Li F, Guo D, Gao X, Zhao X. 2021. Water deficit modulates the CO2 fertilization effect on plant gas exchange and leaf-level water use efficiency: a global meta-analysis. Frontiers in Plant Science 12:775477

doi: 10.3389/fpls.2021.775477
[76]

Wang H, Prentice IC, Davis TW, Keenan TF, Wright IJ, et al. 2017. Photosynthetic responses to altitude: an explanation based on optimality principles. New Phytologist 213:976−982

doi: 10.1111/nph.14332
[77]

Forstner V, Vremec M, Herndl M, Birk S. 2023. Effects of dry spells on soil moisture and yield anomalies at a montane managed grassland site: a lysimeter climate experiment. Ecohydrology 16:e2518

doi: 10.1002/eco.2518
[78]

Cusack DF, Christoffersen B, Smith-Martin CM, Andersen KM, Cordeiro AL, et al. 2024. Toward a coordinated understanding of hydro-biogeochemical root functions in tropical forests for application in vegetation models. New Phytologist 242:351−371

doi: 10.1111/nph.19561