in

Latitudinal patterns and environmental correlates of plant root exudation


Abstract

Root exudation is a key trait involved in the plant carbon economy that exerts an important control over ecosystem functioning. Yet, our understanding of global patterns of root exudation and associated influencing factors remains limited. Here we assembled a comprehensive dataset of specific root exudation rate (that is, organic carbon exuded per unit root biomass), root traits and environmental covariates from in situ measurements and a literature synthesis. Our results show that non-woody plants have a higher specific root exudation rate than woody plants. Specific root exudation rate exhibits an increasing trend from high to low latitudes, and soil available phosphorus emerges as the primary influencing factor. We also show that specific root exudation rate primarily falls at the ‘fast’ end of the conservation axis of the root economics space, reflecting a trade-off between carbon allocation to root metabolism and tissue construction. These findings provide a basis for predictive understanding of plant impacts on belowground biogeochemical cycles.

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Fig. 1: Data distribution of plant root exudation.
The alternative text for this image may have been generated using AI.
Fig. 2: The estimated spatial distribution of plant root exudation in temperate and subtropical biomes.
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Fig. 3: Relationships between root exudation and root traits across species.
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Fig. 4: Factors that influence plant root exudation in the field.
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Mycorrhizal type modifies the position of exudation carbon within the root economics space

Data availability

The data that support the findings of this study are available via figshare at https://doi.org/10.6084/m9.figshare.29254439 (ref. 116), and the data sources are listed in Supplementary Tables 1 and 7.

Code availability

The R codes needed to reproduce the results of this study are available via figshare at https://doi.org/10.6084/m9.figshare.29254439 (ref. 116).

References

  1. 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).

    Article 

    Google Scholar 

  2. 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).

    Article 
    CAS 

    Google Scholar 

  3. Lal, R. Soil carbon sequestration impacts on global climate change and food security. Science 304, 1623–1627 (2004).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  4. Amelung, W. et al. Towards a global-scale soil climate mitigation strategy. Nat. Commun. 11, 5427 (2020).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 

  5. 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).

    Article 
    PubMed 

    Google Scholar 

  6. Panchal, P., Preece, C., Peñuelas, J. & Giri, J. Soil carbon sequestration by root exudates. Trends Plant Sci. 27, 749–757 (2022).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  7. 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).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 

  8. 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).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  9. 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).

    Article 
    PubMed 

    Google Scholar 

  10. 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).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 

  11. Williams, A. & de Vries, F. T. Plant root exudation under drought: implications for ecosystem functioning. New Phytol. 225, 1899–1905 (2020).

    Article 
    PubMed 

    Google Scholar 

  12. 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).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  13. 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).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  14. 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 

  15. 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).

    Article 
    CAS 

    Google Scholar 

  16. 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).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  17. 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).

    Article 
    PubMed 

    Google Scholar 

  18. Chari, N. R. et al. Estimating the global root exudate carbon flux. Biogeochemistry 167, 895–908 (2024).

    Article 
    CAS 

    Google Scholar 

  19. 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).

    Article 

    Google Scholar 

  20. Walker, T. S., Bais, H. P., Grotewold, E. & Vivanco, J. M. Root exudation and rhizosphere biology. Plant Physiol. 132, 44–51 (2003).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 

  21. Haichar, F. E. Z. et al. Plant host habitat and root exudates shape soil bacterial community structure. ISME J. 2, 1221–1230 (2008).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  22. Brunn, M. et al. Carbon allocation to root exudates is maintained in mature temperate tree species under drought. New Phytol. 235, 965–977 (2022).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  23. 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).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  24. 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).

    Article 
    CAS 

    Google Scholar 

  25. 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).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  26. 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).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  27. Yang, L. et al. Root exudation rates decrease with increasing latitude in some tree species. Forests 11, 1045 (2020).

    Article 

    Google Scholar 

  28. Nakayama, M. & Tateno, R. Solar radiation strongly influences the quantity of forest tree root exudates. Trees 32, 871–879 (2018).

    Article 
    CAS 

    Google Scholar 

  29. 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).

    Article 
    CAS 

    Google Scholar 

  30. Dijkstra, F. A., Zhu, B. & Cheng, W. Root effects on soil organic carbon: a double-edged sword. New Phytol. 230, 60–65 (2021).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  31. 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).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  32. Dı́az, S. & Cabido, M. Vive la différence: plant functional diversity matters to ecosystem processes. Trends Ecol. Evol. 16, 646–655 (2001).

    Article 

    Google Scholar 

  33. 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).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 

  34. Pichon, N. A. et al. Nitrogen availability and plant functional composition modify biodiversity-multifunctionality relationships. Ecol. Lett. 27, e14361 (2024).

    Article 
    PubMed 

    Google Scholar 

  35. 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).

    Article 
    PubMed 

    Google Scholar 

  36. Wright, I. J. et al. The worldwide leaf economics spectrum. Nature 428, 821–827 (2004).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  37. Díaz, S. et al. The global spectrum of plant form and function. Nature 529, 167–171 (2016).

    Article 
    PubMed 

    Google Scholar 

  38. Bergmann, J. et al. The fungal collaboration gradient dominates the root economics space in plants. Sci. Adv. 6, eaba3756 (2020).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 

  39. Yaffar, D., Cabugao, K. G. & Meier, I. C. Representing root physiological traits in the root economic space framework. New Phytol. 234, 773–775 (2022).

    Article 
    PubMed 

    Google Scholar 

  40. Weigelt, A. et al. An integrated framework of plant form and function: the belowground perspective. New Phytol. 232, 42–59 (2021).

    Article 
    PubMed 

    Google Scholar 

  41. 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).

    Article 
    PubMed 

    Google Scholar 

  42. 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).

    Article 

    Google Scholar 

  43. Weemstra, M. et al. Towards a multidimensional root trait framework: a tree root review. New Phytol. 211, 1159–1169 (2016).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  44. 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).

    Article 

    Google Scholar 

  45. 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).

    Article 
    CAS 

    Google Scholar 

  46. 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).

    Article 
    CAS 

    Google Scholar 

  47. 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).

    Article 

    Google Scholar 

  48. McCormack, M. L. & Iversen, C. M. Physical and functional constraints on viable belowground acquisition strategies. Front. Plant Sci. 10, 1215 (2019).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar 

  49. 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).

    Article 
    PubMed 

    Google Scholar 

  50. Comas, L. H. et al. Evolutionary patterns and biogeochemical significance of angiosperm root traits. Int. J. Plant Sci. 173, 584–595 (2012).

    Article 

    Google Scholar 

  51. 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).

    Article 
    PubMed 

    Google Scholar 

  52. Filippelli, G. M. The global phosphorus cycle: past, present, and future. Elements 4, 89–95 (2008).

    Article 
    CAS 

    Google Scholar 

  53. Han, M. et al. The latitudinal pattern of fine root intraspecific trait variation among species in plant communities. Nat. Commun. 16, 9340 (2025).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 

  54. Ma, Z. et al. Evolutionary history resolves global organization of root functional traits. Nature 555, 94–97 (2018).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  55. 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).

    Article 
    CAS 

    Google Scholar 

  56. See, C. R. et al. Global patterns in fine root decomposition: climate, chemistry, mycorrhizal association and woodiness. Ecol. Lett. 22, 946–953 (2019).

    Article 
    PubMed 

    Google Scholar 

  57. 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).

    Article 

    Google Scholar 

  58. 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).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar 

  59. 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).

    Article 
    CAS 

    Google Scholar 

  60. 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).

    Article 

    Google Scholar 

  61. Steidinger, B. S. et al. Climatic controls of decomposition drive the global biogeography of forest-tree symbioses. Nature 569, 404–408 (2019).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  62. Reichert, T. et al. Plant phosphorus-use and -acquisition strategies in Amazonia. New Phytol. 234, 1126–1143 (2022).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  63. Walker, T. W. & Syers, J. K. The fate of phosphorus during pedogenesis. Geoderma 15, 1–19 (1976).

    Article 
    CAS 

    Google Scholar 

  64. Crews, T. E. et al. Changes in soil phosphorus fractions and ecosystem dynamics across a long chronosequence in Hawaii. Ecology 76, 1407–1424 (1995).

    Article 

    Google Scholar 

  65. 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).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  66. 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).

    Article 
    CAS 

    Google Scholar 

  67. 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).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  68. 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).

    Article 
    PubMed 

    Google Scholar 

  69. Ma, H. et al. The global distribution and environmental drivers of aboveground versus belowground plant biomass. Nat. Ecol. Evol. 5, 1110–1122 (2021).

    Article 
    PubMed 

    Google Scholar 

  70. 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).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar 

  71. Brunn, M. et al. Tree carbon allocation to root exudates: implications for carbon budgets, soil sequestration and drought response. Tree Physiol. 45, tpaf026 (2025).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  72. Du, E. et al. Global patterns of terrestrial nitrogen and phosphorus limitation. Nat. Geosci. 13, 221–226 (2020).

    Article 
    CAS 

    Google Scholar 

  73. 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).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 

  74. 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).

    Article 
    CAS 

    Google Scholar 

  75. 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).

    Article 

    Google Scholar 

  76. Prescott, C. E. et al. Surplus carbon drives allocation and plant–soil interactions. Trends Ecol. Evol. 35, 1110–1118 (2020).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  77. Martínez-Vilalta, J. et al. Dynamics of non-structural carbohydrates in terrestrial plants: a global synthesis. Ecol. Monogr. 86, 495–516 (2016).

    Article 

    Google Scholar 

  78. Körner, C. Carbon limitation in trees. J. Ecol. 91, 4–17 (2003).

    Article 

    Google Scholar 

  79. 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).

    Article 

    Google Scholar 

  80. 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).

  81. Smith, N. G. et al. Global photosynthetic capacity is optimized to the environment. Ecol. Lett. 22, 506–517 (2019).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar 

  82. Wright, I. J. et al. Global climatic drivers of leaf size. Science 357, 917–921 (2017).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  83. Locosselli, G. M. et al. Global tree-ring analysis reveals rapid decrease in tropical tree longevity with temperature. PNAS 117, 33358–33364 (2020).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 

  84. 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).

    Article 

    Google Scholar 

  85. 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).

    Article 
    CAS 

    Google Scholar 

  86. Zhang, Y. et al. The origin of bi-dimensionality in plant root traits. Trends Ecol. Evol. 39, 78–88 (2024).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  87. Kong, D. et al. Nonlinearity of root trait relationships and the root economics spectrum. Nat. Commun. 10, 2203 (2019).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar 

  88. 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).

    Article 
    PubMed 

    Google Scholar 

  89. Han, M. et al. Root phosphatase activity aligns with the collaboration gradient of the root economics space. New Phytol. 234, 837–849 (2022).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  90. Chen, M. et al. Changes in Chinese fir plantations root exudation strategies seasonally and as tree age. For. Ecol. Manag. 545, 121239 (2023).

    Article 

    Google Scholar 

  91. Rees, F. et al. Deciphering spatiotemporal patterns of rhizodeposition with a functional-structural root model: RhizoDep. Plant Soil 516, 777–795 (2025).

    Article 
    CAS 

    Google Scholar 

  92. 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).

    Article 
    PubMed 

    Google Scholar 

  93. Dinerstein, E. et al. An ecoregion-based approach to protecting half the terrestrial realm. BioScience 67, 534–545 (2017).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar 

  94. 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).

    Article 

    Google Scholar 

  95. 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).

    Article 

    Google Scholar 

  96. 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).

    Article 
    CAS 

    Google Scholar 

  97. Soudzilovskaia, N. A. et al. FungalRoot: global online database of plant mycorrhizal associations. New Phytol. 227, 955–966 (2020).

    Article 
    PubMed 

    Google Scholar 

  98. Wang, B. & Qiu, Y. L. Phylogenetic distribution and evolution of mycorrhizas in land plants. Mycorrhiza 16, 299–363 (2006).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  99. Guerrero-Ramírez, N. R. et al. Global root traits (GRooT) database. Global Ecol. Biogeogr. 30, 25–37 (2021).

    Article 

    Google Scholar 

  100. 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 

  101. Liaw, A. & Wiener, M. Classification and regression by randomForest. R News 2, 18–22 (2002).

    Google Scholar 

  102. Ma, H. et al. The global biogeography of tree leaf form and habit. Nat. Plants 9, 1795–1809 (2023).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar 

  103. Karger, D. N. et al. Climatologies at high resolution for the Earth’s land surface areas. Sci. Data 4, 170122 (2017).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar 

  104. Poggio, L. et al. SoilGrids 2.0: producing soil information for the globe with quantified spatial uncertainty. Soil 7, 217–240 (2021).

    Article 
    CAS 

    Google Scholar 

  105. McDowell, R. W., Noble, A., Pletnyakov, P. & Haygarth, P. M. A global database of soil plant available phosphorus. Sci. Data 10, 125 (2023).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 

  106. 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).

    Article 

    Google Scholar 

  107. 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).

    Article 

    Google Scholar 

  108. 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).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 

  109. Hijmans, R. terra: spatial data analysis. R package version (1.8-8) (R Foundation for Statistical Computing, 2025).

  110. Lê, S., Josse, J. & Husson, F. FactoMineR: an R package for multivariate analysis. J. Stat. Softw. 25, 1–18 (2008).

    Article 

    Google Scholar 

  111. Revell, L. J. phytools 2.0: an updated R ecosystem for phylogenetic comparative methods (and other things). Peerj 12, e16505 (2024).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar 

  112. Cornelis, J. T. & de Tombeur, F. Soil controls on carboxylate-driven processes and opportunities. Plant Soil 476, 239–250 (2022).

    Article 
    CAS 

    Google Scholar 

  113. 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).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  114. Heiberger, R. H. H. Statistical analysis and data display: Heiberger and Holland. R package version 3, 1–52 (2024).

    Google Scholar 

  115. Jin, Y. & Qian, H. V. PhyloMaker: an R package that can generate very large phylogenies for vascular plants. Ecography 42, 1353–1359 (2019).

    Article 

    Google Scholar 

  116. 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).

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Acknowledgements

We express our gratitude to H. Cornelissen for providing valuable comments on an early version of this paper. We acknowledge the help from the Maoershan Forest Research Station of Northeast Forestry University (J. Gu), the Shennongjia Station of the Chinese Ecosystem Research Network (Z. Xie), and the Taiyanghe Nature Reserve (J. Su and S. Li) for access permission and field support. We are grateful for our laboratory members for carrying out field work (L. Sun, D. Gan, Y. Han, M. Yu and R. Li) and all the scientists who contributed to collecting and sharing the root trait data used in this study.

Funding

This study was financially supported by the National Natural Science Foundation of China (grant nos. 32425038 and 32588202, B.Z.; 32301348, M.H.), the Fundamental Research Funds for the Central Universities (grant nos. 2572025BR64 and 2572025JT09, M.H.), the Heilongjiang Provincial Natural Science Foundation of China (grant no. YQ2025C016, M.H.) and the China Postdoctoral Science Foundation (grant nos. BX20220003 and 2022M720006, M.H.).

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B.Z. and M.H. conceived of the idea; M.H. performed field experiments and collected literature data; M.H. and F.J. conducted statistical analyses; F.T.d.V., R.D.B., F.J. and B.Z. provided substantial intellectual inputs and contributed to paper developments; M.H. wrote the first draft of the paper, and all authors contributed to subsequent revisions.

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Biao Zhu.

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Nature Ecology & Evolution thanks Doug Aubrey, Awaz Mohamed, Andrea Schnepf and the other, anonymous, reviewer(s) for their contribution to the peer review of this work.

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Han, M., de Vries, F.T., Bardgett, R.D. et al. Latitudinal patterns and environmental correlates of plant root exudation.
Nat Ecol Evol (2026). https://doi.org/10.1038/s41559-026-03103-4

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