García-Palacios, P. et al. Evidence for large microbial-mediated losses of soil carbon under anthropogenic warming. Nat. Rev. Earth Environ. 2, 507–517 (2021).

Article 

Google Scholar
 

Bossio, D. A. et al. The role of soil carbon in natural climate solutions. Nat. Sustain. 3, 391–398 (2020).

Article 

Google Scholar
 

IPCC. Climate Change 2021—The Physical Science Basis (eds Masson-Delmotte, V. et al.) (Cambridge Univ. Press, 2023).

Zhou, J. et al. Microbial mediation of carbon-cycle feedbacks to climate warming. Nat. Clim. Change 2, 106–110 (2012).

Article 
CAS 

Google Scholar
 

Luo, Y. Terrestrial carbon-cycle feedback to climate warming. Annu. Rev. Ecol. Evol. System. 38, 683–712 (2007).

Article 

Google Scholar
 

Friedlingstein, P. et al. Uncertainties in CMIP5 climate projections due to carbon cycle feedbacks. J. Clim. 27, 511–526 (2014).

Cox, P. M., Betts, R. A., Jones, C. D., Spall, S. A. & Totterdell, I. J. Acceleration of global warming due to carbon-cycle feedbacks in a coupled climate model. Nature 408, 184–187 (2000).

Article 
CAS 

Google Scholar
 

Crowther, T. W. et al. Quantifying global soil carbon losses in response to warming. Nature 540, 104–108 (2016).

Article 
CAS 

Google Scholar
 

Liu, W., Zhang, Z. H. E. & Wan, S. Predominant role of water in regulating soil and microbial respiration and their responses to climate change in a semiarid grassland. Glob. Change Biol. 15, 184–195 (2009).

Article 

Google Scholar
 

Zhang, Z. et al. Effect of climate warming on the annual terrestrial net ecosystem CO2 exchange globally in the boreal and temperate regions. Sci. Rep. 7, 3108 (2017).

Article 
CAS 

Google Scholar
 

D’Orangeville, L. et al. Northeastern North America as a potential refugium for boreal forests in a warming climate. Science 352, 1452–1455 (2016).

Article 

Google Scholar
 

Chen, Y. et al. Warming has a minor effect on surface soil organic carbon in alpine meadow ecosystems on the Qinghai–Tibetan Plateau. Glob. Change Biol. 28, 1618–1629 (2022).

Article 
CAS 

Google Scholar
 

Ziegler, S. E. et al. Climate warming can accelerate carbon fluxes without changing soil carbon stocks. Front. Earth Sci. 5, 2 (2017).

van Gestel, N. et al. Predicting soil carbon loss with warming. Nature 554, E4–E5 (2018).

Article 

Google Scholar
 

Bai, T., Wang, P., Qiu, Y., Zhang, Y. & Hu, S. Nitrogen availability mediates soil carbon cycling response to climate warming: a meta-analysis. Glob. Change Biol. 29, 2608–2626 (2023).

Article 
CAS 

Google Scholar
 

Carney, K. M., Hungate, B. A., Drake, B. G. & Megonigal, J. P. Altered soil microbial community at elevated CO2 leads to loss of soil carbon. Proc. Natl Acad. Sci. USA 104, 4990–4995 (2007).

Article 
CAS 

Google Scholar
 

Wu, L. et al. Reduction of microbial diversity in grassland soil is driven by long-term climate warming. Nat. Microbiol. 7, 1054–1062 (2022).

Article 
CAS 

Google Scholar
 

Reich, P. B. et al. Synergistic effects of four climate change drivers on terrestrial carbon cycling. Nat. Geosci. 13, 787–793 (2020).

Article 
CAS 

Google Scholar
 

Yuan, X. et al. Plant and microbial regulations of soil carbon dynamics under warming in two alpine swamp meadow ecosystems on the Tibetan Plateau. Sci. Total Environ. 790, 148072 (2021).

Article 
CAS 

Google Scholar
 

Huntington, T. G. Evidence for intensification of the global water cycle: review and synthesis. J. Hydrol. 319, 83–95 (2006).

Article 

Google Scholar
 

Reichstein, M. et al. Climate extremes and the carbon cycle. Nature 500, 287–295 (2013).

Article 
CAS 

Google Scholar
 

Easterling, D. R. et al. Climate extremes: observations, modeling, and impacts. Science 289, 2068–2074 (2000).

Article 
CAS 

Google Scholar
 

Wang, J. et al. Precipitation manipulation and terrestrial carbon cycling: the roles of treatment magnitude, experimental duration and local climate. Glob. Ecol. Biogeogr. 30, 1909–1921 (2021).

Article 

Google Scholar
 

Schuur, E. A. G. et al. Climate change and the permafrost carbon feedback. Nature 520, 171–179 (2015).

Article 
CAS 

Google Scholar
 

Rineau, F. et al. Towards more predictive and interdisciplinary climate change ecosystem experiments. Nat. Clim. Change 9, 809–816 (2019).

Article 

Google Scholar
 

Wei, X. et al. Responses of soil C pools to combined warming and altered precipitation regimes: a meta-analysis. Glob. Ecol. Biogeogr. 32, 1660–1675 (2023).

Article 

Google Scholar
 

Song, B. et al. Light and heavy fractions of soil organic matter in response to climate warming and increased precipitation in a temperate steppe. PLoS ONE 7, e33217 (2012).

Article 
CAS 

Google Scholar
 

Poeplau, C. Grassland soil organic carbon stocks along management intensity and warming gradients. Grass Forage Sci. 76, 186–195 (2021).

Article 
CAS 

Google Scholar
 

Pulido, M., Barrena-González, J., Badgery, W., Rodrigo-Comino, J. & Cerdà, A. Sustainable grazing. Curr. Opin. Environ. Sci. Health 5, 42–46 (2018).

Article 

Google Scholar
 

Guo, X. et al. Climate warming leads to divergent succession of grassland microbial communities. Nat. Clim. Change 8, 813–818 (2018).

Article 

Google Scholar
 

Guo, X. et al. Climate warming accelerates temporal scaling of grassland soil microbial biodiversity. Nat. Ecol. Evol. 3, 612–619 (2019).

Article 

Google Scholar
 

Zhang, Y. et al. Experimental warming leads to convergent succession of grassland archaeal community. Nat. Clim. Change 13, 561–569 (2023).

Article 

Google Scholar
 

Chen, H. et al. Carbon and nitrogen cycling on the Qinghai–Tibetan Plateau. Nat. Rev. Earth Environ. 3, 701–716 (2022).

Article 
CAS 

Google Scholar
 

Mishra, U. & Riley, W. J. Scaling impacts on environmental controls and spatial heterogeneity of soil organic carbon stocks. Biogeosciences 12, 3993–4004 (2015).

Article 

Google Scholar
 

Melillo, J. M. et al. Long-term pattern and magnitude of soil carbon feedback to the climate system in a warming world. Science 358, 101–105 (2017).

Article 
CAS 

Google Scholar
 

Rocci, K. S., Lavallee, J. M., Stewart, C. E. & Cotrufo, M. F. Soil organic carbon response to global environmental change depends on its distribution between mineral-associated and particulate organic matter: a meta-analysis. Sci. Total Environ. 793, 148569 (2021).

Article 
CAS 

Google Scholar
 

Schmidt, M. W. I. et al. Persistence of soil organic matter as an ecosystem property. Nature 478, 49–56 (2011).

Article 
CAS 

Google Scholar
 

Soong, J. L. et al. Five years of whole-soil warming led to loss of subsoil carbon stocks and increased CO2 efflux. Sci. Adv. 7, eabd1343 (2021).

Article 
CAS 

Google Scholar
 

Guo, X. et al. Particulate and mineral-associated organic carbon turnover revealed by modelling their long-term dynamics. Soil Biol. Biochem. 173, 108780 (2022).

Article 
CAS 

Google Scholar
 

Chen, Y. et al. Long-term warming reduces surface soil organic carbon by reducing mineral-associated carbon rather than “free” particulate carbon. Soil Biol. Biochem. 177, 108905 (2023).

Article 
CAS 

Google Scholar
 

Xu, X. et al. Global pattern and controls of soil microbial metabolic quotient. Ecol. Monogr. 87, 429–441 (2017).

Article 

Google Scholar
 

Guo, X. et al. Gene-informed decomposition model predicts lower soil carbon loss due to persistent microbial adaptation to warming. Nat. Commun. 11, 4897 (2020).

Article 
CAS 

Google Scholar
 

Chari, N. R. & Taylor, B. N. Soil organic matter formation and loss are mediated by root exudates in a temperate forest. Nat. Geosci. 15, 1011–1016 (2022).

Article 
CAS 

Google Scholar
 

Wieder, W. R., Bonan, G. B. & Allison, S. D. Global soil carbon projections are improved by modelling microbial processes. Nat. Clim. Change 3, 909–912 (2013).

Article 
CAS 

Google Scholar
 

Tao, X. et al. Experimental warming accelerates positive soil priming in a temperate grassland ecosystem. Nat. Commun. 15, 1178 (2024).

Article 
CAS 

Google Scholar
 

Wang, G. et al. Soil enzymes as indicators of soil function: a step toward greater realism in microbial ecological modeling. Glob. Change Biol. 28, 1935–1950 (2022).

Article 
CAS 

Google Scholar
 

Luo, Y. et al. Modeled interactive effects of precipitation, temperature, and CO2 on ecosystem carbon and water dynamics in different climatic zones. Glob. Change Biol. 14, 1986–1999 (2008).

Article 

Google Scholar
 

Tao, F. et al. Microbial carbon use efficiency promotes global soil carbon storage. Nature 618, 981–985 (2023).

Article 
CAS 

Google Scholar
 

Kikstra, J. S. et al. The IPCC Sixth Assessment Report WGIII climate assessment of mitigation pathways: from emissions to global temperatures. Geosci. Model Dev. 15, 9075–9109 (2022).

Article 

Google Scholar
 

Matthews, H. D. & Wynes, S. Current global efforts are insufficient to limit warming to 1.5 °C. Science 376, 1404–1409 (2022).

Article 
CAS 

Google Scholar
 

Zhang, S. et al. Reconciling carbon quality with availability predicts temperature sensitivity of global soil carbon mineralization. Proc. Natl Acad. Sci. USA 121, e2313842121 (2024).

Article 
CAS 

Google Scholar
 

Zhang, Q. et al. Water limitation regulates positive feedback of increased ecosystem respiration. Nat. Ecol. Evol. 8, 1870–1876 (2024).

Article 

Google Scholar
 

Metze, D. et al. Microbial growth under drought is confined to distinct taxa and modified by potential future climate conditions. Nat. Commun. 14, 5895 (2023).

Article 
CAS 

Google Scholar
 

AghaKouchak, A. et al. Climate extremes and compound hazards in a warming world. Annu. Rev. Earth Planet. Sci. 48, 519–548 (2020).

Article 
CAS 

Google Scholar
 

Maestre, F. T. et al. Increasing aridity reduces soil microbial diversity and abundance in global drylands. Proc. Natl Acad. Sci. USA 112, 15684–15689 (2015).

Article 
CAS 

Google Scholar
 

Xu, X. et al. Unchanged carbon balance driven by equivalent responses of production and respiration to climate change in a mixed-grass prairie. Glob. Change Biol. 22, 1857–1866 (2016).

Article 

Google Scholar
 

Cotrufo, M. F., Ranalli, M. G., Haddix, M. L., Six, J. & Lugato, E. Soil carbon storage informed by particulate and mineral-associated organic matter. Nat. Geosci. 12, 989–994 (2019).

Article 
CAS 

Google Scholar
 

Leuthold, S., Haddix, M., Lavallee, J. & Cotrufo, M. F. Physical fractionation techniques. In Reference Module in Earth Systems and Environmental Sciences 1–13 (Elsevier, 2022).

McLean, E. Soil pH and lime requirement. In Methods of soil Analysis. Part 2. Chemical and Microbiological Properties, 2nd ed. (ed. Page, A. L.) Ch. 12 (American Society of Agronomy & Soil Science Society of America, 1982).

Frank, D. A. & McNaughton, S. J. Aboveground biomass estimation with the canopy intercept method: a plant growth form caveat. Oikos 57, 57–60 (1990).

Article 

Google Scholar
 

Xu, X. et al. Plant community structure regulates responses of prairie soil respiration to decadal experimental warming. Glob. Change Biol. 21, 3846–3853 (2015).

Article 

Google Scholar
 

Gong, H. et al. Soil microbial DNA concentration is a powerful indicator for estimating soil microbial biomass C and N across arid and semi-arid regions in northern China. Appl. Soil Ecol. 160, 103869 (2021).

Article 

Google Scholar
 

Rosinger, C., Rousk, J., Bonkowski, M., Rethemeyer, J. & Jaeschke, A. Rewetting the hyper-arid Atacama Desert soil reactivates a carbon-starved microbial decomposer community and also triggers archaeal metabolism. Sci. Total Environ. 892, 164785 (2023).

Article 
CAS 

Google Scholar
 

Nilsson, R. H. et al. The UNITE database for molecular identification of fungi: handling dark taxa and parallel taxonomic classifications. Nucleic Acids Res. 47, D259–D264 (2019).

Article 
CAS 

Google Scholar
 

Zhou, J. et al. High-throughput metagenomic technologies for complex microbial community analysis: open and closed formats. MBio 6, e02288-02214 (2015).

Article 

Google Scholar
 

Bates, D., Mächler, M., Bolker, B. & Walker, S. Fitting linear mixed-effects models using lme4. J. Stat. Softw. 67, 1–48 (2015).

Article 

Google Scholar
 

Oksanen, J. et al. Vegan: community ecology package. R package version 2.0-10 (2013).

Lefcheck, J., Byrnes, J. & Grace, J. Package ‘piecewiseSEM’. R package version 1 (2016).

Wang, G. et al. Soil moisture drives microbial controls on carbon decomposition in two subtropical forests. Soil Biol. Biochem. 130, 185–194 (2019).

Article 
CAS 

Google Scholar
 

Wang, G., Li, W., Wang, K. & Huang, W. Uncertainty quantification of the soil moisture response functions for microbial dormancy and resuscitation. Soil Biol. Biochem. 160, 108337 (2021).

Article 
CAS 

Google Scholar
 

Liang, J. et al. Evaluating the E3SM land model version 0 (ELMv0) at a temperate forest site using flux and soil water measurements. Geosci. Model Dev. 12, 1601–1612 (2019).

Article 
CAS 

Google Scholar
 

Li, J. et al. Reduced carbon use efficiency and increased microbial turnover with soil warming. Glob. Change Biol. 25, 900–910 (2019).

Article 

Google Scholar
 

Huang, W. et al. High carbon losses from oxygen-limited soils challenge biogeochemical theory and model assumptions. Glob. Change Biol. 27, 6166–6180 (2021).

Article 

Google Scholar
 

Zhou, S. et al. Enhanced understanding of soil methane processes through modeling microbial kinetics and taxonomy. Soil Biol. Biochem. 207, 109838 (2025).

Article 
CAS 

Google Scholar
 

Guisan, A. & Zimmermann, N. E. Predictive habitat distribution models in ecology. Ecol. Model. 135, 147–186 (2000).

Article 

Google Scholar
 

Ellsworth, D. S. et al. Photosynthesis, carboxylation and leaf nitrogen responses of 16 species to elevated pCO2 across four free-air CO2 enrichment experiments in forest, grassland and desert. Glob. Change Biol. 10, 2121–2138 (2004).

Article 

Google Scholar
 

Batstone, D. J., Pind, P. F. & Angelidaki, I. Kinetics of thermophilic, anaerobic oxidation of straight and branched chain butyrate and valerate. Biotechnol. Bioeng. 84, 195–204 (2003).

Article 
CAS 

Google Scholar
 

Yang, Z. zhifengyang-ou/MEND-warming-soil-C: MEND for simulating soil C feedback under climate change (1.0.0). Zenodo https://doi.org/10.5281/zenodo.18396578 (2026).