in

The overlooked role of root water content in the root economics space


Abstract

Root traits are fundamental to plant survival, growth and adaptation to environmental changes. Despite increasing attention to the root economics space, a quantitative understanding of global patterns and key drivers of root trait variation remains elusive. By combining metabolic theory with global trait datasets, we reveal universal nonlinear relationships of five key root traits with root water content regardless of plant growth form or climate zone. Root water content emerges as a stronger predictor of growth-related root traits and shows a closer association with the conservation gradient than the widely considered root nitrogen, thereby better defining ‘fast’ resource acquisition strategies. Moreover, replacing nitrogen with tissue water content in analyses reveals a closer alignment of leaf and fine-root traits than expected. Our findings highlight general quantitative biotic and abiotic controls on plant trait variation, offering broader insights into plant economics strategies, community dynamics and ecosystem functioning under changing climate and resource availability.

Access through your institution

Buy or subscribe

This is a preview of subscription content, access via your institution

Access options

Access through your institution

Buy this article

USD 39.95

Prices may be subject to local taxes which are calculated during checkout

Fig. 1: General nonlinear relationships between key root traits and dry-mass-based RWC.
Fig. 2: RWC is a more robust predictor of other root traits than RN.
Fig. 3: RWC better defines the fast end of the conservation gradient of the RES than the widely considered trait RN.
Fig. 4: Replacing nitrogen with plant water content improves the alignment between leaf and fine-root traits.

Similar content being viewed by others

Aboveground and belowground sizes are aligned in the unified spectrum of plant form and function

Growth form and lifespan of herbaceous species mediate the role of traits in short-term drought response

Three-dimensional in vivo analysis of water uptake and translocation in maize roots by fast neutron tomography

Data availability

The TRY plant trait database is publicly available at https://www.try-db.org. The GRooT database is publicly available at https://github.com/GRooT-Database/GRooT-Data. The WorldClim version 2.1 database is publicly available at https://www.worldclim.org/. Global soil temperature maps are available via Zenodo at https://doi.org/10.5281/zenodo.4558732 (ref. 59). The Global Aridity Index and Potential Evapotranspiration Database version 3 is publicly available via figshare at https://doi.org/10.6084/m9.figshare.7504448.v6 (ref. 60). The datasets generated and analysed in this study are available via the Open Science Framework at https://osf.io/65s3c/.

Code availability

The R code used for data analysis in this study is available via the Open Science Framework at https://osf.io/65s3c/.

References

  1. 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 

  2. Cavender-Bares, J., Kozak, K. H., Fine, P. V. A. & Kembel, S. W. The merging of community ecology and phylogenetic biology. Ecol. Lett. 12, 693–715 (2009).

    Article 
    PubMed 

    Google Scholar 

  3. 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 

  4. Maurel, C. & Nacry, P. Root architecture and hydraulics converge for acclimation to changing water availability. Nat. Plants 6, 744–749 (2020).

    Article 
    PubMed 

    Google Scholar 

  5. 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 

  6. Carmona, C. P. et al. Fine-root traits in the global spectrum of plant form and function. Nature 597, 683–687 (2021).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  7. Matthus, E. et al. Revisiting the root economics space—its applications, extensions and nuances advance our understanding of fine-root functioning. Plant Soil https://doi.org/10.1007/s11104-025-07379-6 (2025).

    Article 

    Google Scholar 

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

    Article 
    PubMed 
    PubMed Central 

    Google Scholar 

  9. Donoghue, M. J. A phylogenetic perspective on the distribution of plant diversity. Proc. Natl Acad. Sci. USA 105, 11549–11555 (2008).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 

  10. 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 

  11. Weigelt, A. et al. The importance of trait selection in ecology. Nature 618, E29–E30 (2023).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  12. Bueno, C. G. et al. Reply to: The importance of trait selection in ecology. Nature 618, E31–E34 (2023).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  13. Slade, S. Elements in Living Organisms (Rosen Publishing Group’s PowerKids Press, 2006).

  14. Huang, H. et al. A general model for seed and seedling respiratory metabolism. Am. Nat. 195, 534–546 (2020).

    Article 
    PubMed 

    Google Scholar 

  15. Huang, H. et al. Water content quantitatively affects metabolic rates over the course of plant ontogeny. New Phytol. 228, 1524–1534 (2020).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  16. Wang, Z. Q. et al. Leaf water content contributes to global leaf trait relationships. Nat. Commun. 13, 5525 (2022).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 

  17. Stears, A. E. et al. Water availability dictates how plant traits predict demographic rates. Ecology 103, e3799 (2022).

    Article 
    PubMed 

    Google Scholar 

  18. Smart, S. M. et al. Leaf dry matter content is better at predicting above-ground net primary production than specific leaf area. Funct. Ecol. 31, 1336–1344 (2017).

    Article 

    Google Scholar 

  19. Osnas, J. L., Lichstein, J. W., Reich, P. B. & Pacala, S. W. Global leaf trait relationships: mass, area, and the leaf economics spectrum. Science 340, 741–744 (2013).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  20. Lloyd, J., Bloomfield, K., Domingues, T. F. & Farquhar, G. D. Photosynthetically relevant foliar traits correlating better on a mass vs an area basis: of ecophysiological relevance or just a case of mathematical imperatives and statistical quicksand? New Phytol. 199, 311–321 (2013).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  21. Lin, D. M. et al. Relationships between rhizosphere microbial communities, soil abiotic properties and root trait variation within a pine species. J. Ecol. 112, 1275–1286 (2024).

    Article 
    CAS 

    Google Scholar 

  22. Roumet et al. Root structure–function relationships in 74 species: evidence of a root economics spectrum related to carbon economy. New Phytol. 210, 815–826 (2016).

    Article 
    PubMed 

    Google Scholar 

  23. Han, M. & Zhu, B. Linking root respiration to chemistry and morphology across species. Glob. Change Biol. 27, 190–201 (2021).

    Article 
    CAS 

    Google Scholar 

  24. Brown, J. H., Gillooly, J. F., Allen, A. P., Savage, V. M. & West, G. B. Toward a metabolic theory of ecology. Ecology 85, 1771–1789 (2004).

    Article 

    Google Scholar 

  25. Michaletz, S. T., Cheng, D. L., Kerkhoff, A. J. & Enquist, B. J. Convergence of terrestrial plant production across global climate gradients. Nature 512, 39–43 (2014).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  26. Michaletz, S. T. & Garen, J. C. Hotter is not (always) better: embracing unimodal scaling of biological rates with temperature. Ecol. Lett. 27, e14381 (2024).

    Article 
    PubMed 

    Google Scholar 

  27. De Sisto, M. L., MacDougall, A. H., Mengis, N. & Antoniello, S. Modelling the terrestrial nitrogen and phosphorus cycle in the UVic ESCM. Geosci. Model Dev. 16, 4113–4136 (2023).

    Article 

    Google Scholar 

  28. Bartlett, M. K., Scoffoni, C. & Sack, L. The determinants of leaf turgor loss point and prediction of drought tolerance of species and biomes: a global meta-analysis. Ecol. Lett. 15, 393–405 (2012).

    Article 
    PubMed 

    Google Scholar 

  29. Sibly, R. M., Brown, J. H. & Kodric-Brown, A. Metabolic Ecology: A Scaling Approach (John Wiley & Sons, 2012).

  30. Vaieretti, M. V., Diaz, S., Vile, D. & Garnier, E. Two measurement methods of leaf dry matter content produce similar results in a broad range of species. Ann. Bot. 99, 955–958 (2007).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar 

  31. Reich, P. B. The world-wide ‘fast–slow’ plant economics spectrum: a traits manifesto. J. Ecol. 102, 275–301 (2014).

    Article 

    Google Scholar 

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

    Article 
    CAS 
    PubMed 

    Google Scholar 

  33. Laughlin, D. C. Plant Strategies: The Demographic Consequences of Functional Traits in Changing Environments (Oxford Univ. Press, 2023).

  34. Beccari, E. & Carmona, C. P. Aboveground and belowground sizes are aligned in the unified spectrum of plant form and function. Nat. Commun. 15, 9199 (2024).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 

  35. West, G. B., Brown, J. H. & Enquist, B. J. A general model for the origin of allometric scaling laws in biology. Science 276, 122–126 (1997).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  36. West, G. B., Brown, J. H. & Enquist, B. J. A general model for the structure and allometry of plant vascular systems. Nature 400, 664–667 (1999).

    Article 
    CAS 

    Google Scholar 

  37. West, G. B., Brown, J. H. & Enquist, B. J. The fourth dimension of life: fractal geometry and allometric scaling of organisms. Science 284, 1677–1679 (1999).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  38. Birouste, M., Zamora-Ledezma, E., Bossard, C., Pérez-Ramos, I. M. & Roumet, C. Measurement of fine root tissue density: a comparison of three methods reveals the potential of root dry matter content. Plant Soil 374, 299–313 (2014).

    Article 
    CAS 

    Google Scholar 

  39. 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 

  40. Zhang, Y. et al. The origin of bi-dimensionality in plant root traits. Trends Ecol. Evol. 39, 77–78 (2024).

    Article 

    Google Scholar 

  41. Kattge, J. & Sandel, B. TRY plant trait database—enhanced coverage and open access. Glob. Change Biol. 26, 5343 (2020).

    Article 

    Google Scholar 

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

    Article 

    Google Scholar 

  43. Kindt, R. WorldFlora: An R package for exact and fuzzy matching of plant names against the World Flora Online Taxonomic Backbone data. Appl. Plant. Sci. 8,e11388 (2020).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar 

  44. Stekhoven, D. J. & Buhlmann, P. MissForest—non-parametric missing value imputation for mixed-type data. Bioinformatics 28, 112–118 (2012).

    Article 
    CAS 
    PubMed 

    Google Scholar 

  45. Jin, Y. & Qian H. V.PhyloMaker2: An updated and enlarged R package that can generate very large phylogenies for vascular plants. Plant Divers. 44, 335–339 (2022).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar 

  46. Legendre, P. lmodel2: Model II regression. R package v.1.7-3. CRAN https://CRAN.R-project.org/package=lmodel2 (2018).

  47. Lembrechts, J. J. et al. Global maps of soil temperature. Glob. Change Biol. 28, 3110–3144 (2022).

    Article 
    CAS 

    Google Scholar 

  48. Fox, J. & Weisberg, S. An R Companion to Applied Regression (Sage Publications, 2019).

  49. Baty, F. et al. The toolbox for nonlinear regression in R: the package nlstools. J. Stat. Softw. 66, 1–21 (2015).

    Article 

    Google Scholar 

  50. Kuhn, M. Building predictive models in R using the caret package. J. Stat. Softw. 28, 1–26 (2008).

    Article 

    Google Scholar 

  51. Padfield, D., O’Sullivan, H. & Pawar, S. rTPC and nls.multstart: A new pipeline to fit thermal performance curves in R. Methods Ecol. Evol. 12, 1138–1143 (2021).

    Article 

    Google Scholar 

  52. Denelle, P., Weigelt, P. & Kreft, H. GIFT—An R package to access the Global Inventory of Floras and Traits. Methods Ecol. Evol. 14, 2738–2748 (2023).

    Article 

    Google Scholar 

  53. Zomer, R. J., Xu, J. C. & Trabucco, A. Version 3 of the global aridity index and potential evapotranspiration database. Sci. Data 9, 409 (2022).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar 

  54. William, R. psych: Procedures for psychological, psychometric, and personality research v.2.5.3. CRAN https://CRAN.R-project.org/package=psych. (2025).

  55. Dinno, A. paran: Horn’s test of principal components/factors. R package v.1.5.3. CRAN https://CRAN.R-project.org/package=paran (2024).

  56. Duong, T. ks: Kernel smoothing. R package v.1.14.3. CRAN https://CRAN.R-project.org/package=ks (2024).

  57. Carmona, C. P. TPD: methods for measuring functional diversity based on Trait Probability Density. R package version 1.1.0. CRAN https://CRAN.R-project.org/package=TPD (2019).

  58. R Core Team R: A Language and Environment for Statistical Computing (R Foundation for Statistical Computing, 2024).

  59. van den Hoogen, J., Lembrechts, J., SoilTemp, Nijs, I. & Lenoir, J. Global Soil Bioclimatic variables at 30 arc second resolution (Version 1) [Data set]. Zenodo https://doi.org/10.5281/zenodo.4558732 (2021).

  60. Zomer, R. & Trabucco, A. Global Aridity Index and Potential Evapotranspiration (ET0) Database: Version 6. figshare https://doi.org/10.6084/m9.figshare.7504448.v6 (2022).

Download references

Acknowledgements

This study was supported by the Young Talent Program from Guangdong Province (grant no. 77010-42150005), the Start-up Grant (no. 77010-12255006) and the Fundamental Research Funds for the Central Universities (grant no. 77010-13130003) from Sun Yat-sen University. C.C. was supported by the National Natural Science Foundation of China (grant no. 32330064). C.P.C. was supported by the Estonian Research Council (grant no. PRG2142) and the Spanish Ministry of Science, Innovation and Universities through the ATRAE Program 2024 (grant no. ATR2024-154934). J.P. and J.S. were supported by the Spanish government grant nos. PID2022-140808NB-I00 and PID2023-153125NB-I00 funded by the MICIU/AEI/10.13039/501100011033 and FEDER, EU.

Author information

Authors and Affiliations

Authors

Contributions

H.H. and H.L. conceived the study. H.L., J.T. and Y.X. collected the trait data and performed a literature search. H.L. performed the analyses. H.L., H.H., C.P.C., S.N., I.J.W., Y.Z., J.P., J.S., Z.W., L.D., J.W., R.L. and C.C. discussed the design and methods and interpreted the results. H.L. and H.H. wrote the paper with contributions from all authors.

Corresponding author

Correspondence to
Heng Huang.

Ethics declarations

Competing interests

The authors declare no competing interests.

Peer review

Peer review information

Nature Plants thanks Benjamin M. Delory, Justus Hennecke and the other, anonymous, reviewer(s) for their contribution to the peer review of this work.

Additional information

Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Extended data

Extended Data Fig. 1 The quantitative effects of root water content on root traits across growth forms.

The nonlinear relationships of RRcor (a), SRAcor (b), SRLcor (c), RDcor (d), RTD (e) with RWC for non-woody (blue) and woody species (orange) are shown, respectively. The parameter values were obtained from the non-linear fitting regression results based on our theoretical framework (equations (5), (6) and (8)–(10)).

Extended Data Fig. 2 The quantitative effects of root water content on root traits across climate zones.

The nonlinear relationships of SRAcor (a), SRLcor (b), RDcor (c), RTD (d) with RWC for dry (defined as areas with aridity index (AI) ≤ 0.65; red points and lines) and humid climate zones (defined as areas with AI > 0.65; green points and lines) are shown, respectively. The parameter values were obtained from the non-linear fitting regression results based on our theoretical framework (equations (5), (6) and (8)–(10)).

Extended Data Fig. 3 Correlations of root traits with mean annual precipitation.

The scaling relationships of RR (a, nmol g−1 s−1), SRA (b, cm2 g−1), SRL (c, m g−1), RD (d, mm), RTD (e, g cm−3), and RWC (f, g g−1) with MAP across the pooled data are shown, respectively. The OLS regression lines (black lines) are shown.

Extended Data Fig. 4 Correlations of root traits with mean annual precipitation.

The scaling relationships of RR (a, nmol g−1 s−1), SRA (b, cm2 g−1), SRL (c, m g−1), RD (d, mm), RTD (e, g cm−3), and RWC (f, g g−1) with mean annual aridity index (AI) are shown across the pooled data, respectively. The OLS regression lines (black lines) are shown.

Extended Data Fig. 5 Pairwise correlations of all traits in the imputed dataset including 1657 species (blue) and complete dataset (red) including 150 species.

Scatterplots indicate the log10-transformed trait-trait relationships. The blue and red lines in the scatterplots represent the SMA regression fitting results. The correlation coefficients are shown in the upper-right triangle. The diagonals represent the probability density distributions of each trait.

Extended Data Fig. 6 The scaling relationships among root traits.

a, The scaling relationship of RR (nmol g−1 s−1) with SRA (cm2 g−1) is presented. b, The scaling relationship of SRL (m g−1) with RD (mm) are shown. c and d, The scaling relationships of RR (nmol g−1 s−1) with SRL (m g−1) and RD (mm) are shown, respectively. The SMA regression lines (black lines) are shown.

Supplementary information

Supplementary Information

Supplementary Figs. 1–10, Tables 1–10, Methods, Results and Data Sources.

Reporting Summary

Rights and permissions

Springer Nature or its licensor (e.g. a society or other partner) holds exclusive rights to this article under a publishing agreement with the author(s) or other rightsholder(s); author self-archiving of the accepted manuscript version of this article is solely governed by the terms of such publishing agreement and applicable law.

Reprints and permissions

About this article

Cite this article

Li, H., Carmona, C.P., Niu, S. et al. The overlooked role of root water content in the root economics space.
Nat. Plants (2026). https://doi.org/10.1038/s41477-026-02232-9

Download citation

  • Received:

  • Accepted:

  • Published:

  • Version of record:

  • DOI: https://doi.org/10.1038/s41477-026-02232-9


Source: Ecology - nature.com

Dynamic deepwater invertebrate populations challenge the concept of oxygen-rich reference conditions for European lakes

Global patterns of commodity-driven deforestation and associated carbon emissions