Abstract
Plant biomass tends to increase under nutrient addition and decrease under drought. Biotic and abiotic factors influence responses to both, making the combined impact of nutrient addition and drought difficult to predict. Using a globally distributed network of manipulative field experiments, we assessed grassland aboveground biomass response to both drought and increased nutrient availability at 26 sites across nine countries. Overall, drought reduced biomass by 19% and nutrient addition increased it by 24%, resulting in no net impact under combined drought and nutrient addition. Among the plant functional groups, only graminoids responded positively to nutrients during drought. However, these general responses depended on local conditions, especially aridity. Nutrient effects were stronger in arid grasslands and weaker in humid regions and nitrogen-rich soils, although nutrient addition alleviated drought effects the most in subhumid sites. Biomass responses were weaker with higher precipitation variability. Biomass increased more with increased nutrient availability and declined more with drought at high-diversity sites than at low-diversity sites. Our findings highlight the importance of local abiotic and biotic conditions in predicting grassland responses to anthropogenic nutrient and climate changes.
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Data availability
All data, including the measured biomass in the studied grasslands, as well as the applied biotic and abiotic data used for analysis, are available via figshare at https://doi.org/10.6084/m9.figshare.27249012 (ref. 59).
Code availability
The R code used for analysis is available via figshare at https://doi.org/10.6084/m9.figshare.27249012 (ref. 59).
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Acknowledgements
This work used data from NPK-D Net (https://www.bayceer.uni-bayreuth.de/npkd/index.php?lang=en) experiment, funded at the site level by individual researchers following protocols from the Nutrient Network (http://www.nutnet.org) experiment and Drought-Net (https://droughtnet.weebly.com/). Q.Y. and his working collaboration group received funding from the National Key R&D Program of China (2022YFE0128000, 2022YFF1300603), the National Natural Science Foundation of China (32171592, 32061123005) and the Scientific and Technological Innovation Project of China Academy of Chinese Medical Sciences (CI2024C003YN). V.F.B. thanks the Alexander von Humboldt Foundation for the Georg Forster Research Fellowship that provided the support necessary to complete most of this work during her research stay at the University of Bayreuth, Germany. L.Y. acknowledges PICT 2019-02324 from the National Agency of Scientific Promotion. A.J. received funding from the Federal Ministry of Education and Research Germany (BMBF, FKZ 031B1067C). E.W.S., E.T.B., F.I. and J.G. acknowledge support by grants from the National Science Foundation Research Coordination Network (NSF-DEB-1042132) and Long-Term Ecological Research (NSF-DEB-1234162 and NSF-DEB-1831944 to Cedar Creek LTER) programmes and the Institute on the Environment (DG-0001-13). They thank P.W. and A. Asmus for data coordination and management. M.D.S. was supported by a National Science Foundation (NSF) Research Coordination Network grant (DEB-1354732), US Department of Agriculture’s National Institute of Food and Agriculture (USDA-NIFA) Postdoctoral Fellowship grant (2020-67034-31898), USDA-NIFA Conference Grant (2020-67019-31757), US Geological Survey John Wesley Powell Center for Analysis and Synthesis grant, US Geological Survey grant (G21AC10266-00), and a Global Drought Synthesis Group grant funded by the NSF Long-term Ecological Research Network Office (LNO) and the National Center for Ecological Analysis and Synthesis, University of California-Santa Barbara. J.A. and P.D. acknowledge grants from CONICET, FONCyT and UNMdP. H.A. received funding from the Helmholtz-Centre for Environmental Research - UFZ and thanks the staff of the Bad Lauchstädt Research Station for maintaining the plots and infrastructures. N.E. received support from iDiv funded by the German Research Foundation (DFG– FZT 118, 202548816). A.C.G., E.V. and G.M.W. received funding from the Hermon Slade Foundation (HSF 19103) and the Australian National Landcare Program: DigiFarm, the Holsworth Wildlife Research Endowment, and they thank the staff at the Narrabri Plant Breeding Institute for site maintenance. Y.M. and M.T. acknowledge support from a grant from the National Research Foundation (grant no: 116262). Y.H. thanks the Dutch state forestry (Staatsbosbeheer) for providing access to the study site.
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Y.H. and Q.Y. developed and framed the research question. Y.H., A.J., Q.Y., L.Y., E.T.B., E.W.S. and M.D.S. coordinated the NPK-D Net collaboration. V.F.B. and C.X. led the writing of the paper. V.F.B., G.R.O. and H.D. analysed data. C.X., P.W., J.A., M.C., A.K., S.A.P., T.O. and L.B. contributed to data analysis. P.W., L.Y., Q.Y., E.T.B., A.J., E.W.S., J.A., G.R.O., H.D., M.C., A.K., S.A.P., N.E., F.I., H. Auge, M.H.C., A.C.C., P.D., T.F., A.C.G., S.E.K., T.O., P.P., A.P., D.S., M.T., A.V., E.V., G.M.W., C.W. and G.R.W. contributed to writing the paper. V.F.B., C.X., A.J., J.A., M.C., A.K., S.A.P., N.E., F.I., H. Auge, M.H.C., A.C.C., P.D., T.F., A.C.G., S.E.K., P.P., A.P., D.S., M.T., A.V., E.V., G.M.W., C.W., G.W., H. An, H.J.D., J.G., L.B.H., Y.G.K., J.L.L., Y.M., D.S.T., D.T., S.W., C.Z.W., K.W., H.H.W., A.Y., F.W.Z., B.Z., J.Z., N.Z. and X.Z. contributed to data collection and were site-level coordinators. Further details of each author’s contribution can be found in Supplementary Table 10.
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Bondaruk, V.F., Xu, C., Wilfahrt, P. et al. Aridity modulates grassland biomass responses to combined drought and nutrient addition. Nat Ecol Evol 9, 937–946 (2025). https://doi.org/10.1038/s41559-025-02705-8
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DOI: https://doi.org/10.1038/s41559-025-02705-8
