Abstract
Droughts have intensified under climate change, threatening ecosystem stability. While rising atmospheric CO2 concentrations may enhance vegetation drought resistance, the net effect remains uncertain amid concurrent warming. Here we combine ecological modeling with multi-source observations to investigate how CO2 and warming jointly regulate vegetation drought responses on the Qinghai-Tibetan Plateau, a sensitive alpine region exposed to escalating drought threats under changing precipitation regimes. Using factorial scenarios to isolate individual forcings, we show that 40-year CO2 rise mitigated drought-induced productivity losses by 5.7 ± 0.9% under constant temperature. However, in the presence of warming, rising CO2 intensifies drought stress by 5.2 ± 0.5%, reflecting increased plant water demand and disrupted regional water supply-demand balance. Permafrost areas experienced the strongest CO2-driven drought alleviation under constant temperature, but also the greatest warming-induced reversal. These findings reveal interacting CO2-warming impacts on alpine vegetation drought responses, highlighting ecological risks for the plateau and other permafrost-dominant regions under future warming.
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Data availability
Source data used to generate figures have been deposited in the Science Data Bank and are publicly available at https://doi.org/10.57760/sciencedb.36264. Other publicly available datasets include: the China Meteorological Forcing Data (CMFD) (https://doi.org/10.11888/AtmosphericPhysics.tpe.249369.file); the Climatic Research Unit gridded Time Series (CRU TS) v4.07 (https://crudata.uea.ac.uk/cru/data/hrg/cru_ts_4.07/); soil data obtained from SoilGrids (https://www.isric.org/explore/soilgrids); annual time series of [CO2] between 1979 and 2018 obtained from the National Oceanic and Atmospheric Administration Earth System Research Laboratory (NOAA) (https://www.esrl.noaa.gov/gmd/ccgg/trends/); the dataset of annual GPP over China’s terrestrial ecosystems during 2000–2020 (ChinaFLUX20) (https://doi.org/10.11922/11-6035.csd.2023.0037.zh); a global OCO-2-based solar-induced chlorophyll fluorescence dataset (GOSIF v2) (https://globalecology.unh.edu/); a long-term global GPP dataset based on the near-infrared (NIR) reflectance of vegetation (NIRv) (https://doi.org/10.6084/m9.figshare.12981977.v2); the Global Land Evaporation Amsterdam Model (GLEAM) v3 (https://www.gleam.eu/); the Simple Terrestrial Hydrosphere Model (SiTHv2) (https://doi.org/10.11888/Terre.tpdc.300751); records from the China River Sediment Bulletin published by the Ministry of Water Resources of China (http://www.mwr.gov.cn/); the Global Runoff Data Centre (GRDC) (https://www.bafg.de/GRDC/); TRENDY-v12 dataset (https://mdosullivan.github.io/GCB/); the vegetation map of China (1:1,000,000) (https://www.plantplus.cn/en/doi/10.12282/plantdata.0155).
Code availability
Scripts used to generate Figs. 2–5 are available at https://github.com/helv716/code_for_figures_lpjguess_QTP. All scripts were written in Python (v3.10). Figure 1 is a hand-drawn conceptual diagram and is therefore not included in the repository.
References
Walker, A. P. et al. Integrating the evidence for a terrestrial carbon sink caused by increasing atmospheric CO2. New Phytol. 229, 2413–2445 (2021).
Ainsworth, E. A. & Long, S. P. What have we learned from 15 years of free-air CO2 enrichment (FACE)? A meta-analytic review of the responses of photosynthesis, canopy properties and plant production to rising CO2. New Phytol. 165, 351–372 (2005).
Wang, Z., Wang, C. & Liu, S. Elevated CO2 alleviates adverse effects of drought on plant water relations and photosynthesis: a global meta-analysis. J. Ecol. 110, 2836–2849 (2022).
Zeng, Z. et al. Impact of Earth Greening on the Terrestrial Water Cycle. https://doi.org/10.1175/JCLI-D-17-0236.1 (2018).
Vicente-Serrano, S. M. et al. The uncertain role of rising atmospheric CO2 on global plant transpiration. Earth-Sci. Rev. 230, 104055 (2022).
Dai, A. Increasing drought under global warming in observations and models. Nat. Clim. Change 3, 52–58 (2013).
Chai, Y. et al. Global reduction in sensitivity of vegetation water use efficiency to increasing CO2. J. Hydrol. 641, 131844 (2024).
Yao, Y. et al. Declining tradeoff between resistance and resilience of ecosystems to drought. Earth’s Future 12, e2024EF004665 (2024).
Jiang, M., Kelly, J. W. G., Atwell, B. J., Tissue, D. T. & Medlyn, B. E. Drought by CO2 interactions in trees: a test of the water savings mechanism. New Phytol. 230, 1421–1434 (2021).
De Kauwe, M. G., Medlyn, B. E. & Tissue, D. T. To what extent can rising [CO2] ameliorate plant drought stress? New Phytol. 231, 2118–2124 (2021).
van der Kooi, C. J., Reich, M., Löw, M., de Kok, L. J. & Tausz, M. Growth and yield stimulation under elevated CO2 and drought: a meta-analysis on crops. Environ. Exp. Bot. 122, 150–157 (2016).
McDowell, N. G. et al. Mechanisms of woody-plant mortality under rising drought, CO2 and vapour pressure deficit. Nat. Rev. Earth Environ. 3, 294–308 (2022).
Yao, T. et al. From Tibetan Plateau to Third Pole and Pan-Third Pole. Bull. Chin. Acad. Sci. 32, 924–931 (2017).
Yao, T. et al. The imbalance of the Asian water tower. Nat. Rev. Earth Environ. 3, 618–632 (2022).
Wang, T. et al. The current and future of terrestrial carbon balance over the Tibetan Plateau. Sci. China Earth Sci. 53, 1506–1516 (2023).
Liu, Y. et al. Recent centennial drought on the Tibetan Plateau is outstanding within the past 3500 years. Nat. Commun. 16, 1311 (2025).
Gao, Q. et al. On the multi-scale spatiotemporal characteristics of drought in China based on SPEI. Water Sav. Irrig. 111–120 https://doi.org/10.12396/jsgg.2023488 (2024).
Wang, S., Che, L., Zhu, H. & Wang, M. Temporal and spatial variation characteristics of meteorological drought in the crop growing season in the Qinghai-Xizang Plateau from 1961 to 2020. Chin. J. Appl. Environ. Biol. 30, 1093–1100 (2024).
Liu, D. et al. Deciphering impacts of climate extremes on Tibetan grasslands in the last fifteen years. Sci. Bull. 64, 446–454 (2019).
Wang, C., Huang, M. & Zhai, P. Change in drought conditions and its impacts on vegetation growth over the Tibetan Plateau. Adv. Clim. Change Res. 12, 333–341 (2021).
Piao, S. et al. Characteristics, drivers and feedbacks of global greening. Nat. Rev. Earth Environ. 1, 14–27 (2020).
Cuo, L., Zhang, Y., Xu-Ri & Zhou, B. Decadal change and inter-annual variability of net primary productivity on the Tibetan Plateau. Clim. Dyn. 56, 1837–1857 (2021).
Jia, B., Luo, X., Wang, L. & Lai, X. Changes in water use efficiency caused by climate change, CO2 fertilization, and land use changes on the Tibetan Plateau. Adv. Atmos. Sci. 40, 144–154 (2023).
Xia, J. et al. CO2 enrichment accelerates alpine plant growth via increasing water-use efficiency. Agric. For. Meteorol. 352, 110036 (2024).
Guo, Y. et al. Enhanced leaf turnover and nitrogen recycling sustain CO2 fertilization effect on tree-ring growth. Nat. Ecol. Evol. 6, 1271–1278 (2022).
Wang, Y. et al. Response of vegetation to drought in the Tibetan Plateau: Elevation differentiation and the dominant factors. Agric. For. Meteorol. 306, 108468 (2021).
Liu, Y. et al. Soil moisture drought and diverse impacts on vegetation across the Tibetan Plateau in recent three decades. Sci. Total Environ. 963, 178367 (2025).
Fu, G. et al. A meta-analysis of the effects of experimental warming on plant physiology and growth on the Tibetan Plateau. J. Plant Growth Regul. 34, 57–65 (2015).
Wang, R. et al. Recent increase in the observation-derived land evapotranspiration due to global warming. Environ. Res. Lett. 17, 024020 (2022).
Baig, S., Medlyn, B. E., Mercado, L. M. & Zaehle, S. Does the growth response of woody plants to elevated CO2 increase with temperature? A model-oriented meta-analysis. Glob. Change Biol. 21, 4303–4319 (2015).
Ehlers, T. A. et al. Past, present, and future geo-biosphere interactions on the Tibetan Plateau and implications for permafrost. Earth-Sci. Rev. 234, 104197 (2022).
Wang, T. et al. Unsustainable water supply from thawing permafrost on the Tibetan Plateau in a changing climate. Sci. Bull. 68, 1105–1108 (2023).
Wang, X. et al. Contrasting characteristics changes and linkages of permafrost between the Arctic and the Third Pole. Earth-Sci. Rev. 230, 104042 (2020).
Sitch, S. et al. Trends and drivers of terrestrial sources and sinks of carbon dioxide: an overview of the TRENDY Project. Glob. Biogeochem. Cycles 38, e2024GB008102 (2024).
Smith, B. et al. Implications of incorporating N cycling and N limitations on primary production in an individual-based dynamic vegetation model. Biogeosciences 11, 2027–2054 (2014).
Hickler, T. et al. Using a generalized vegetation model to simulate vegetation dynamics in Northeastern USA. Ecology 85, 519–530 (2004).
Chaudhary, N., Miller, P. A. & Smith, B. Modelling Holocene peatland dynamics with an individual-based dynamic vegetation model. Biogeosciences 14, 2571–2596 (2017).
Smith, S. D. et al. Elevated CO2 increases productivity and invasive species success in an arid ecosystem. Nature 408, 79–82 (2000).
Zak, D. R., Pregitzer, K. S., Kubiske, M. E. & Burton, A. J. Forest productivity under elevated CO2 and O3: positive feedbacks to soil N cycling sustain decade-long net primary productivity enhancement by CO2. Ecol. Lett. 14, 1220–1226 (2011).
Bai, P., Liu, X., Zhang, Y. & Liu, C. Assessing the impacts of vegetation greenness change on evapotranspiration and water yield in China. Water Resour. Res. 56, e2019WR027019 (2020).
Gimeno, T. E., McVicar, T. R., O’Grady, A. P., Tissue, D. T. & Ellsworth, D. S. Elevated CO2 did not affect the hydrological balance of a mature native Eucalyptus woodland. Glob. Change Biol. 24, 3010–3024 (2018).
Rustad, L. et al. A meta-analysis of the response of soil respiration, net nitrogen mineralization, and aboveground plant growth to experimental ecosystem warming. Oecologia 126, 543–562 (2001).
Shi, Z. et al. Evidence for long-term shift in plant community composition under decadal experimental warming. J. Ecol. 103, 1131–1140 (2015).
Ganjurjav, H. et al. Differential response of alpine steppe and alpine meadow to climate warming in the central Qinghai–Tibetan Plateau. Agric. For. Meteorol. 223, 233–240 (2016).
Zou, F., Li, H. & Hu, Q. Responses of vegetation greening and land surface temperature variations to global warming on the Qinghai-Tibetan Plateau, 2001–2016. Ecol. Indic. 119, 106867 (2020).
Bafitlhile, T. M. & Liu, Y. Temperature contributes more than precipitation to the greening of the Tibetan Plateau during 1982–2019. Theor. Appl. Climatol. 147, 1471–1488 (2022).
Wang, Y. et al. Vegetation structural shift tells environmental changes on the Tibetan Plateau over 40 years. Sci. Bull. 68, 1928–1937 (2023).
Chen, X. et al. A doubled increasing trend of evapotranspiration on the Tibetan Plateau. Sci. Bull. 69, 1980–1990 (2024).
Cheng, T. F., Chen, D., Wang, B., Ou, T. & Lu, M. Human-induced warming accelerates local evapotranspiration and precipitation recycling over the Tibetan Plateau. Commun. Earth Environ. 5, 1–15 (2024).
Ganjurjav, H. et al. Warming tends to decrease ecosystem carbon and water use efficiency in dissimilar ways in an alpine meadow and a cultivated grassland in the Tibetan Plateau. Agric. For. Meteorol. 323, 109079 (2022).
Wolkovich, E. M. et al. Warming experiments underpredict plant phenological responses to climate change. Nature 485, 494–497 (2012).
Yao, T. et al. Different glacier status with atmospheric circulations in Tibetan Plateau and surroundings. Nat. Clim. Change 2, 663–667 (2012).
Li, J. et al. Weakening warming on spring freeze–thaw cycle caused greening Earth’s third pole. Proc. Natl. Acad. Sci. USA 121, e2319581121 (2024).
Abdelhakim, L. O. A., Zhou, R. & Ottosen, C.-O. Physiological Responses of Plants to Combined Drought and Heat under Elevated CO2. Agronomy 12, 2526 (2022).
Duan, H. et al. Elevated [CO2] does not ameliorate the negative effects of elevated temperature on drought-induced mortality in Eucalyptus radiata seedlings. Plant Cell Environ. 37, 1598–1613 (2014).
Meyer, B. F. et al. Simulating the drought response of European tree species with the dynamic vegetation model LPJ-GUESS (v4.1, 97c552c5). EGUsphere, 1–37, https://doi.org/10.5194/egusphere-2024-3352 (2024).
Monteith, J. L. Accommodation between transpiring vegetation and the convective boundary layer. J. Hydrol. 166, 251–263 (1995).
Haxeltine, A. & Prentice, I. C. BIOME3: an equilibrium terrestrial biosphere model based on ecophysiological constraints, resource availability, and competition among plant functional types. Glob. Biogeochem. Cycles 10, 693–709 (1996).
De Kauwe, M. G. et al. Forest water use and water use efficiency at elevated CO2: a model-data intercomparison at two contrasting temperate forest FACE sites. Glob. Change Biol. 19, 1759–1779 (2013).
Medlyn, B. E. et al. Using models to guide field experiments: a priori predictions for the CO2 response of a nutrient- and water-limited native Eucalypt woodland. Glob. Change Biol. 22, 2834–2851 (2016).
Sitch, S. et al. Evaluation of ecosystem dynamics, plant geography and terrestrial carbon cycling in the LPJ dynamic global vegetation model. Glob. Change Biol. 9, 161–185 (2003).
Zhou, H., Tang, J., Olin, S. & Miller, P. A. A comprehensive evaluation of hydrological processes in a second-generation dynamic vegetation model. Hydrol. Process. 38, e15152 (2024).
Liu, C., Li, C. & Li, L. Climate warming benefits plant growth but not net carbon uptake: simulation of Alaska Tundra and Needle Leaf Forest Using LPJ-GUESS. Land 13, 632 (2024).
Quan, Q. et al. Drought-induced peatland carbon loss exacerbated by elevated CO2 and warming. Science 390, 367–370 (2025).
He, J. et al. The first high-resolution meteorological forcing dataset for land process studies over China. Sci. Data 7, 25 (2020).
Yang, K. et al. China Meteorological Forcing Dataset v1.6 (1979-2018). National Tibetan Plateau Data Center. https://doi.org/10.11888/AtmosphericPhysics.tpe.249369.file (2025).
Harris, I., Osborn, T. J., Jones, P. & Lister, D. Version 4 of the CRU TS monthly high-resolution gridded multivariate climate dataset. Sci. Data 7, 109 (2020).
Fan, R. et al. A dataset of annual gross primary productivity in China’s terrestrial ecosystems during 2000-2020. China Sci. Data 8, 1–13 (2023).
Fan, R. et al. A dataset of annual gross primary productivity in China’s terrestrial ecosystems during 2000-2020. Science Data Bank. https://doi.org/10.11922/11-6035.csd.2023.0037.zh (2023).
Li, X. & Xiao, J. Mapping photosynthesis solely from solar-induced chlorophyll fluorescence: a global, fine-resolution dataset of gross primary production derived from OCO-2. Remote Sens. 11, 2563 (2019).
Wang, S., Zhang, Y., Ju, W., Qiu, B. & Zhang, Z. Tracking the seasonal and inter-annual variations of global gross primary production during last four decades using satellite near-infrared reflectance data. Sci. Total Environ. 755, 142569 (2021).
Martens, B. et al. GLEAM v3: satellite-based land evaporation and root-zone soil moisture. Geosci. Model Dev. 10, 1903–1925 (2017).
Zhang, K. et al. A global dataset of terrestrial evapotranspiration and soil moisture dynamics from 1982 to 2020. Sci. Data 11, 445 (2024).
Zhang, K. & Zhu, G. A global dataset of terrestrial evapotranspiration (multi-component) and soil moisture (multi-layer) from 1982 to 2022. National Tibetan Plateau Data Center. https://doi.org/10.11888/Terre.tpdc.300751 (2025).
Wei, D. et al. Plant uptake of CO2 outpaces losses from permafrost and plant respiration on the Tibetan Plateau. Proc. Natl. Acad. Sci. USA 118, e2015283118 (2021).
Wania, R., Ross, I. & Prentice, I. C. Integrating peatlands and permafrost into a dynamic global vegetation model: 2. Evaluation and sensitivity of vegetation and carbon cycle processes. Glob. Biogeochem. Cycles 23, GB3015 (2009).
Zhang, X. Vegetation map of the People’s Republic of China (1:1,000,000) and its illustration put to press. Acta Ecol. Sin. http://en.cnki.com.cn/Article_en/CJFDTotal-STXB200803053.htm (2007).
Zhang, X. Vegetation Map of the People’s Republic of China (1:1000000). Plant Data Center of Chinese Academy of Sciences. https://doi.org/10.12282/plantdata.0155 (2021).
Nelson, F. E. & Outcalt, S. I. A computational method for prediction and regionalization of permafrost. Arct. Alp. Res. 19, 279–288 (1987).
Zou, D. et al. A new map of permafrost distribution on the Tibetan Plateau. Cryosphere 11, 2527–2542 (2017).
Monserud, R. A. & Leemans, R. Comparing global vegetation maps with the Kappa statistic. Ecol. Model. 62, 275–293 (1992).
Moreira, E. E., Martins, D. S. & Pereira, L. S. Assessing drought cycles in SPI time series using a Fourier analysis. Nat. Hazards Earth Syst. Sci. 15, 571–585 (2015).
Zhai, J. et al. Spatial variation and trends in PDSI and SPI indices and their relation to streamflow in 10 large regions of China. J. Clim. 23, 649–663 (2010).
Svoboda, M. et al. The drought monitor. Bull. Am. Meteorol. Soc. 83, 1181–1190 (2002).
McKee, T. The relationship of drought frequency and duration to time scales. In Proc. 8th Conference on Applied Climatology, Anaheim, 17-22 January 1993 179–184 (Scientific Research, 1993).
Khan, S., Gabriel, H. F. & Rana, T. Standard precipitation index to track drought and assess impact of rainfall on watertables in irrigation areas. Irrig. Drain. Syst. 22, 159–177 (2008).
Yang, J. & Huang, X. The 30 m annual land cover datasets and its dynamics in China from 1985 to 2023. Zenodo, https://doi.org/10.5281/zenodo.12779975 (2024).
Dash, C. H. S. K., Behera, A. K., Dehuri, S. & Ghosh, A. An outliers detection and elimination framework in classification task of data mining. Decis. Anal. J. 6, 100164 (2023).
Acknowledgements
This study was supported by the Ministry of Science and Technology of China National R&D Program (2022YFF0801904), National Natural Science Foundation of China (32301383, 32471669), Zhejiang Provincial Natural Science Foundation (LR24C030001, LZ23C030001), Key Research and Development Program of Zhejiang (2024C03244), and Fundamental Research Funds for the Central Universities (226-2024-00187). The authors acknowledge technical support by Paul Miller, Jing Tang, Johan Nord, Stefan Olin, Thomas Pugh, and Drew Holzworth. The authors also acknowledge all the modelers involved in the TRENDY project for providing access to their simulation data.
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M.J. and H.L. conceived the study. H.L., D.W., and J.K. modified the model. H.L., X.Z., and C.C. collected the data. H.L. and X.Z. preprocessed the data for model simulation and evaluation. H.L. conducted model simulations and performed the analysis. H.L. wrote the first draft with contributions from J.S., X.Z., and M.J. All co-authors (H.L., X.Z., J.S., D.W., J.K., L.T., J.C., X.X., C.C., J.Z., J.N., S.S., Y.F., B.M., B.S., Y.Y., and M.J.) contributed to the discussion and revision of the manuscript.
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J.C. is an Editorial Board Member for Communications Earth & Environment, but was not involved in the editorial review of, nor the decision to publish this article. The authors declare no other competing interests.
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Lyu, H., Zhang, X., Su, J. et al. Warming overwhelms CO2-driven drought mitigation in alpine vegetation on the Qinghai-Tibetan Plateau.
Commun Earth Environ (2026). https://doi.org/10.1038/s43247-026-03308-2
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DOI: https://doi.org/10.1038/s43247-026-03308-2
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