Jackson, R. B. et al. The ecology of soil carbon: pools, vulnerabilities, and biotic and abiotic controls. Annu. Rev. Ecol. Evol. Syst. 48, 419–445 (2017).
Jenkinson, D. S., Adams, D. E. & Wild, A. Model estimates of CO2 emissions from soil in response to global warming. Nature 351, 304–306 (1991).
Lal, R. Soil carbon sequestration impacts on global climate change and food security. Science 304, 1623–1627 (2004).
Amelung, W. et al. Towards a global-scale soil climate mitigation strategy. Nat. Commun. 11, 5427 (2020).
Keller, A. B. et al. Root-derived inputs are major contributors to soil carbon in temperate forests, but vary by mycorrhizal type. Ecol. Lett. 24, 626–635 (2021).
Panchal, P., Preece, C., Peñuelas, J. & Giri, J. Soil carbon sequestration by root exudates. Trends Plant Sci. 27, 749–757 (2022).
Villarino, S. H., Pinto, P., Jackson, R. B. & Piñeiro, G. Plant rhizodeposition: a key factor for soil organic matter formation in stable fractions. Sci. Adv. 7, eabd3176 (2021).
Vives-Peris, V., de Ollas, C., Gomez-Cadenas, A. & Perez-Clemente, R. M. Root exudates: from plant to rhizosphere and beyond. Plant Cell Rep. 39, 3–17 (2020).
Wen, Z., White, P. J., Shen, J. & Lambers, H. Linking root exudation to belowground economic traits for resource acquisition. New Phytol. 233, 1620–1635 (2021).
McLaughlin, S., Zhalnina, K., Kosina, S., Northen, T. R. & Sasse, J. The core metabolome and root exudation dynamics of three phylogenetically distinct plant species. Nat. Commun. 14, 1649 (2023).
Williams, A. & de Vries, F. T. Plant root exudation under drought: implications for ecosystem functioning. New Phytol. 225, 1899–1905 (2020).
Bais, H. P., Weir, T. L., Perry, L. G., Gilroy, S. & Vivanco, J. M. The role of root exudates in rhizosphere interactions with plants and other organisms. Annu. Rev. Plant Biol. 57, 233–266 (2006).
Zhalnina, K. et al. Dynamic root exudate chemistry and microbial substrate preferences drive patterns in rhizosphere microbial community assembly. Nat. Microbiol. 3, 470–480 (2018).
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).
Meier, I. C., Finzi, A. C. & Phillips, R. P. Root exudates increase N availability by stimulating microbial turnover of fast-cycling N pools. Soil Biol. Biochem. 106, 119–128 (2017).
Lambers, H., Hayes, P. E., Laliberté, E., Oliveira, R. S. & Turner, B. L. Leaf manganese accumulation and phosphorus-acquisition efficiency. Trends Plant Sci. 20, 83–90 (2015).
Lambers, H., Raven, J. A., Shaver, G. R. & Smith, S. E. Plant nutrient-acquisition strategies change with soil age. Trends Ecol. Evol. 23, 95–103 (2008).
Chari, N. R. et al. Estimating the global root exudate carbon flux. Biogeochemistry 167, 895–908 (2024).
Pausch, J. & Kuzyakov, Y. Carbon input by roots into the soil: quantification of rhizodeposition from root to ecosystem scale. Global Change Biol. 24, 1–12 (2018).
Walker, T. S., Bais, H. P., Grotewold, E. & Vivanco, J. M. Root exudation and rhizosphere biology. Plant Physiol. 132, 44–51 (2003).
Haichar, F. E. Z. et al. Plant host habitat and root exudates shape soil bacterial community structure. ISME J. 2, 1221–1230 (2008).
Brunn, M. et al. Carbon allocation to root exudates is maintained in mature temperate tree species under drought. New Phytol. 235, 965–977 (2022).
Liese, R., Lubbe, T., Albers, N. W. & Meier, I. C. The mycorrhizal type governs root exudation and nitrogen uptake of temperate tree species. Tree Physiol. 38, 83–95 (2018).
Rog, I. et al. Increased belowground tree carbon allocation in a mature mixed forest in a dry versus a wet year. Global Change Biol. 30, e17172 (2024).
Sun, L. et al. Root exudation as a major competitive fine-root functional trait of 18 coexisting species in a subtropical forest. New Phytol. 229, 259–271 (2021).
Meier, I. C. et al. Root exudation of mature beech forests across a nutrient availability gradient: the role of root morphology and fungal activity. New Phytol. 226, 583–594 (2020).
Yang, L. et al. Root exudation rates decrease with increasing latitude in some tree species. Forests 11, 1045 (2020).
Nakayama, M. & Tateno, R. Solar radiation strongly influences the quantity of forest tree root exudates. Trees 32, 871–879 (2018).
Spohn, M., Ermak, A. & Kuzyakov, Y. Microbial gross organic phosphorus mineralization can be stimulated by root exudates-A 33P isotopic dilution study. Soil Biol. Biochem. 65, 254–263 (2013).
Dijkstra, F. A., Zhu, B. & Cheng, W. Root effects on soil organic carbon: a double-edged sword. New Phytol. 230, 60–65 (2021).
Phillips, R. P., Brzostek, E. & Midgley, M. G. The mycorrhizal-associated nutrient economy: a new framework for predicting carbon-nutrient couplings in temperate forests. New Phytol. 199, 41–51 (2013).
Dı́az, S. & Cabido, M. Vive la différence: plant functional diversity matters to ecosystem processes. Trends Ecol. Evol. 16, 646–655 (2001).
Augusto, L. & Boča, A. Tree functional traits, forest biomass, and tree species diversity interact with site properties to drive forest soil carbon. Nat. Commun. 13, 1097 (2022).
Pichon, N. A. et al. Nitrogen availability and plant functional composition modify biodiversity-multifunctionality relationships. Ecol. Lett. 27, e14361 (2024).
Hagan, J. G., Henn, J. J. & Osterman, W. H. A. Plant traits alone are good predictors of ecosystem properties when used carefully. Nat. Ecol. Evol. 7, 332–334 (2023).
Wright, I. J. et al. The worldwide leaf economics spectrum. Nature 428, 821–827 (2004).
Díaz, S. et al. The global spectrum of plant form and function. Nature 529, 167–171 (2016).
Bergmann, J. et al. The fungal collaboration gradient dominates the root economics space in plants. Sci. Adv. 6, eaba3756 (2020).
Yaffar, D., Cabugao, K. G. & Meier, I. C. Representing root physiological traits in the root economic space framework. New Phytol. 234, 773–775 (2022).
Weigelt, A. et al. An integrated framework of plant form and function: the belowground perspective. New Phytol. 232, 42–59 (2021).
Bardgett, R. D., Mommer, L. & De Vries, F. T. Going underground: root traits as drivers of ecosystem processes. Trends Ecol. Evol. 29, 692–699 (2014).
Freschet, G. T., Cornelissen, J. H. C., van Logtestijn, R. S. P. & Aerts, R. Evidence of the ‘plant economics spectrum’ in a subarctic flora. J. Ecol. 98, 362–373 (2010).
Weemstra, M. et al. Towards a multidimensional root trait framework: a tree root review. New Phytol. 211, 1159–1169 (2016).
Williams, A. et al. Root functional traits explain root exudation rate and composition across a range of grassland species. J. Ecol. 110, 21–33 (2022).
Hou, F. et al. Root exudates from drought-affected plants increase soil respiration across a range of grassland species. Soil Biol. Biochem. 203, 109731 (2025).
Jiang, Z. et al. Plant growth strategy determines the magnitude and direction of drought-induced changes in root exudates in subtropical forests. Global Change Biol. 29, 3476–3488 (2023).
Kramer-Walter, K. R. et al. Root traits are multidimensional: specific root length is independent from root tissue density and the plant economic spectrum. J. Ecol. 104, 1299–1310 (2016).
McCormack, M. L. & Iversen, C. M. Physical and functional constraints on viable belowground acquisition strategies. Front. Plant Sci. 10, 1215 (2019).
Valverde-Barrantes, O. J., Freschet, G. T., Roumet, C. & Blackwood, C. B. A worldview of root traits: the influence of ancestry, growth form, climate and mycorrhizal association on the functional trait variation of fine-root tissues in seed plants. New Phytol. 215, 1562–1573 (2017).
Comas, L. H. et al. Evolutionary patterns and biogeochemical significance of angiosperm root traits. Int. J. Plant Sci. 173, 584–595 (2012).
Vitousek, P. M., Porder, S., Houlton, B. Z. & Chadwick, O. A. Terrestrial phosphorus limitation: mechanisms, implications, and nitrogen–phosphorus interactions. Ecol. Appl. 20, 5–15 (2010).
Filippelli, G. M. The global phosphorus cycle: past, present, and future. Elements 4, 89–95 (2008).
Han, M. et al. The latitudinal pattern of fine root intraspecific trait variation among species in plant communities. Nat. Commun. 16, 9340 (2025).
Ma, Z. et al. Evolutionary history resolves global organization of root functional traits. Nature 555, 94–97 (2018).
Zemunik, G., Turner, B. L., Lambers, H. & Laliberte, E. Diversity of plant nutrient-acquisition strategies increases during long-term ecosystem development. Nat. Plants 1, 15050 (2015).
See, C. R. et al. Global patterns in fine root decomposition: climate, chemistry, mycorrhizal association and woodiness. Ecol. Lett. 22, 946–953 (2019).
Schenk, H. J. & Jackson, R. B. Rooting depths, lateral root spreads and below-ground/above-ground allometries of plants in water-limited ecosystems. J. Ecol. 90, 480–494 (2002).
Tumber-Dávila, S. J., Schenk, H. J., Du, E. & Jackson, R. B. Plant sizes and shapes above and belowground and their interactions with climate. New Phytol. 235, 1032–1056 (2022).
Yin, H., Wheeler, E. & Phillips, R. P. Root-induced changes in nutrient cycling in forests depend on exudation rates. Soil Biol. Biochem. 78, 213–221 (2014).
Phillips, R. P. & Fahey, T. J. Patterns of rhizosphere carbon flux in sugar maple (Acer saccharum) and yellow birch (Betula allegheniensis) saplings. Global Change Biol. 11, 983–995 (2005).
Steidinger, B. S. et al. Climatic controls of decomposition drive the global biogeography of forest-tree symbioses. Nature 569, 404–408 (2019).
Reichert, T. et al. Plant phosphorus-use and -acquisition strategies in Amazonia. New Phytol. 234, 1126–1143 (2022).
Walker, T. W. & Syers, J. K. The fate of phosphorus during pedogenesis. Geoderma 15, 1–19 (1976).
Crews, T. E. et al. Changes in soil phosphorus fractions and ecosystem dynamics across a long chronosequence in Hawaii. Ecology 76, 1407–1424 (1995).
Ryan, M. H. et al. Carbon trading for phosphorus gain: the balance between rhizosphere carboxylates and arbuscular mycorrhizal symbiosis in plant phosphorus acquisition. Plant Cell Environ. 35, 2170–2180 (2012).
Courty, P.-E. et al. The role of ectomycorrhizal communities in forest ecosystem processes: New perspectives and emerging concepts. Soil Biol. Biochem. 42, 679–698 (2010).
Smith, S. E. & Smith, F. A. Roles of arbuscular mycorrhizas in plant nutrition and growth: new paradigms from cellular to ecosystem scales. Annu. Rev. Plant Biol. 62, 227–250 (2011).
Iversen, C. M. et al. A global Fine-Root Ecology Database to address below-ground challenges in plant ecology. New Phytol. 215, 15–26 (2017).
Ma, H. et al. The global distribution and environmental drivers of aboveground versus belowground plant biomass. Nat. Ecol. Evol. 5, 1110–1122 (2021).
Spawn, S. A., Sullivan, C. C., Lark, T. J. & Gibbs, H. K. Harmonized global maps of above and belowground biomass carbon density in the year 2010. Sci. Data 7, 112 (2020).
Brunn, M. et al. Tree carbon allocation to root exudates: implications for carbon budgets, soil sequestration and drought response. Tree Physiol. 45, tpaf026 (2025).
Du, E. et al. Global patterns of terrestrial nitrogen and phosphorus limitation. Nat. Geosci. 13, 221–226 (2020).
Zhu, J., Wu, A. & Zhou, G. Spatial distribution patterns of soil total phosphorus influenced by climatic factors in China’s forest ecosystems. Sci. Rep. 11, 5357 (2021).
George, T. S. et al. Organic phosphorus in the terrestrial environment: a perspective on the state of the art and future priorities. Plant Soil 427, 191–208 (2018).
Galván-Tejada, N. C., Peña-Ramírez, V., Mora-Palomino, L. & Siebe, C. Soil P fractions in a volcanic soil chronosequence of Central Mexico and their relationship to foliar P in pine trees. J. Plant Nutr. Soil Sci. 177, 792–802 (2014).
Prescott, C. E. et al. Surplus carbon drives allocation and plant–soil interactions. Trends Ecol. Evol. 35, 1110–1118 (2020).
Martínez-Vilalta, J. et al. Dynamics of non-structural carbohydrates in terrestrial plants: a global synthesis. Ecol. Monogr. 86, 495–516 (2016).
Körner, C. Carbon limitation in trees. J. Ecol. 91, 4–17 (2003).
Wang, H. J. et al. Factors determining soil nutrient distribution in a small-scaled watershed in the purple soil region of Sichuan Province, China. Soil Tillage Res. 105, 300–306 (2009).
Dar, G. H., Bhat, R. A., Mehmood, M. A. & Hakeem, K. R. Microbiota and Biofertilizers, Vol 2: Ecofriendly Tools for Reclamation of Degraded Soil Environs (Springer, 2021).
Smith, N. G. et al. Global photosynthetic capacity is optimized to the environment. Ecol. Lett. 22, 506–517 (2019).
Wright, I. J. et al. Global climatic drivers of leaf size. Science 357, 917–921 (2017).
Locosselli, G. M. et al. Global tree-ring analysis reveals rapid decrease in tropical tree longevity with temperature. PNAS 117, 33358–33364 (2020).
Freschet, G. T. et al. Climate, soil and plant functional types as drivers of global fine-root trait variation. J. Ecol. 105, 1182–1196 (2017).
Matthus, E. et al. Revisiting the root economics space—its applications, extensions and nuances advance our understanding of fine-root functioning. Plant Soil 514, 1–27 (2025).
Zhang, Y. et al. The origin of bi-dimensionality in plant root traits. Trends Ecol. Evol. 39, 78–88 (2024).
Kong, D. et al. Nonlinearity of root trait relationships and the root economics spectrum. Nat. Commun. 10, 2203 (2019).
Laughlin, D. C. et al. Root traits explain plant species distributions along climatic gradients yet challenge the nature of ecological trade-offs. Nat. Ecol. Evol. 5, 1123–1134 (2021).
Han, M. et al. Root phosphatase activity aligns with the collaboration gradient of the root economics space. New Phytol. 234, 837–849 (2022).
Chen, M. et al. Changes in Chinese fir plantations root exudation strategies seasonally and as tree age. For. Ecol. Manag. 545, 121239 (2023).
Rees, F. et al. Deciphering spatiotemporal patterns of rhizodeposition with a functional-structural root model: RhizoDep. Plant Soil 516, 777–795 (2025).
Li, M., Duncan, K., Topp, C. N. & Chitwood, D. H. Persistent homology and the branching topologies of plants. Am. J. Bot. 104, 349–353 (2017).
Dinerstein, E. et al. An ecoregion-based approach to protecting half the terrestrial realm. BioScience 67, 534–545 (2017).
Olson, D. M. et al. Terrestrial ecoregions of the world: a new map of life on Earth: a new global map of terrestrial ecoregions provides an innovative tool for conserving biodiversity. BioScience 51, 933–938 (2001).
Phillips, R. P., Erlitz, Y., Bier, R. & Bernhardt, E. S. New approach for capturing soluble root exudates in forest soils. Funct. Ecol. 22, 990–999 (2008).
Han, M., Sun, L., Gan, D., Fu, L. & Zhu, B. Root functional traits are key determinants of the rhizosphere effect on soil organic matter decomposition across 14 temperate hardwood species. Soil Biol. Biochem. 151, 108019 (2020).
Soudzilovskaia, N. A. et al. FungalRoot: global online database of plant mycorrhizal associations. New Phytol. 227, 955–966 (2020).
Wang, B. & Qiu, Y. L. Phylogenetic distribution and evolution of mycorrhizas in land plants. Mycorrhiza 16, 299–363 (2006).
Guerrero-Ramírez, N. R. et al. Global root traits (GRooT) database. Global Ecol. Biogeogr. 30, 25–37 (2021).
Bates, D., Mächler, M., Bolker, B. & Walker, S. Fitting linear mixed-effects models using lme4. J. Stat. Softw. 67, 1–48 (2015).
Liaw, A. & Wiener, M. Classification and regression by randomForest. R News 2, 18–22 (2002).
Ma, H. et al. The global biogeography of tree leaf form and habit. Nat. Plants 9, 1795–1809 (2023).
Karger, D. N. et al. Climatologies at high resolution for the Earth’s land surface areas. Sci. Data 4, 170122 (2017).
Poggio, L. et al. SoilGrids 2.0: producing soil information for the globe with quantified spatial uncertainty. Soil 7, 217–240 (2021).
McDowell, R. W., Noble, A., Pletnyakov, P. & Haygarth, P. M. A global database of soil plant available phosphorus. Sci. Data 10, 125 (2023).
Tuanmu, M. N. & Jetz, W. A global 1-km consensus land-cover product for biodiversity and ecosystem modelling. Global Ecol. Biogeogr. 23, 1031–1045 (2014).
Valavi, R., Elith, J., Lahoz-Monfort, J. J. & Guillera-Arroita, G. blockCV: An r package for generating spatially or environmentally separated folds for k-fold cross-validation of species distribution models. Methods Ecol. Evol. 10, 225–232 (2019).
Gurmesa, G. A. et al. Retention of deposited ammonium and nitrate and its impact on the global forest carbon sink. Nat. Commun. 13, 880 (2022).
Hijmans, R. terra: spatial data analysis. R package version (1.8-8) (R Foundation for Statistical Computing, 2025).
Lê, S., Josse, J. & Husson, F. FactoMineR: an R package for multivariate analysis. J. Stat. Softw. 25, 1–18 (2008).
Revell, L. J. phytools 2.0: an updated R ecosystem for phylogenetic comparative methods (and other things). Peerj 12, e16505 (2024).
Cornelis, J. T. & de Tombeur, F. Soil controls on carboxylate-driven processes and opportunities. Plant Soil 476, 239–250 (2022).
Semchenko, M., Xue, P. & Leigh, T. Functional diversity and identity of plant genotypes regulate rhizodeposition and soil microbial activity. New Phytol. 232, 776–787 (2021).
Heiberger, R. H. H. Statistical analysis and data display: Heiberger and Holland. R package version 3, 1–52 (2024).
Jin, Y. & Qian, H. V. PhyloMaker: an R package that can generate very large phylogenies for vascular plants. Ecography 42, 1353–1359 (2019).
Han, M. et al. Data for latitudinal patterns and environmental correlates of plant root exudation. figshare https://doi.org/10.6084/m9.figshare.29254439 (2026).