Skip to main content

Thank you for visiting nature.com. You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser (or turn off compatibility mode in Internet Explorer). In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript.

  • Article
  • Published:

The worldwide leaf economics spectrum

Abstract

Bringing together leaf trait data spanning 2,548 species and 175 sites we describe, for the first time at global scale, a universal spectrum of leaf economics consisting of key chemical, structural and physiological properties. The spectrum runs from quick to slow return on investments of nutrients and dry mass in leaves, and operates largely independently of growth form, plant functional type or biome. Categories along the spectrum would, in general, describe leaf economic variation at the global scale better than plant functional types, because functional types overlap substantially in their leaf traits. Overall, modulation of leaf traits and trait relationships by climate is surprisingly modest, although some striking and significant patterns can be seen. Reliable quantification of the leaf economics spectrum and its interaction with climate will prove valuable for modelling nutrient fluxes and vegetation boundaries under changing land-use and climate.

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

Access options

Rent or buy this article

Prices vary by article type

from$1.95

to$39.95

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

Figure 1: Mean annual rainfall (MAR) and mean annual temperature (MAT).
Figure 2: Three-way trait relationships among the six leaf traits with reference to LMA, one of the key traits in the leaf economics spectrum.
Figure 3: LMA as a function of MAT and MAR at the study sites (data for 2,370 species from 163 sites; rainfall and LMA are log10-scaled).
Figure 4: LL as a function of LMA and MAR (all axes are log10-scaled).

Similar content being viewed by others

References

  1. Bloom, A. J., Chapin, F. S. & Mooney, H. A. Resource limitation in plants—an economic analogy. Annu. Rev. Ecol. Syst. 16, 363–392 (1985)

    Article  Google Scholar 

  2. Orians, G. H. & Solbrig, O. T. A cost-income model of leaves and roots with special reference to arid and semiarid area. Am. Nat. 111, 677–690 (1977)

    Article  Google Scholar 

  3. Givnish, T. J. (ed.) On the Economy of Plant Form and Function (Cambridge Univ. Press, Cambridge/New York, 1986)

  4. Niinemets, U. Global-scale climatic controls of leaf dry mass per area, density, and thickness in trees and shrubs. Ecology 82, 453–469 (2001)

    Article  Google Scholar 

  5. Niinemets, U. Components of leaf dry mass per area—thickness and density—alter leaf photosynthetic capacity in reverse directions in woody plants. New Phytol. 144, 35–47 (1999)

    Article  Google Scholar 

  6. Reich, P. B., Walters, M. B. & Ellsworth, D. S. From tropics to tundra: global convergence in plant functioning. Proc. Natl Acad. Sci. USA 94, 13730–13734 (1997)

    Article  ADS  CAS  Google Scholar 

  7. Reich, P. B. et al. Generality of leaf trait relationships: a test across six biomes. Ecology 80, 1955–1969 (1999)

    Article  Google Scholar 

  8. Field, C. & Mooney, H. A. in On the Economy of Plant Form and Function (ed. Givnish, T. J.) 25–55 (Cambridge Univ. Press, Cambridge, 1986)

    Google Scholar 

  9. Reich, P. B. et al. Relationships of leaf dark respiration to leaf nitrogen, specific leaf area and leaf life-span—a test across biomes and functional groups. Oecologia 114, 471–482 (1998)

    Article  ADS  Google Scholar 

  10. Wright, I. J., Reich, P. B. & Westoby, M. Strategy-shifts in leaf physiology, structure and nutrient content between species of high and low rainfall, and high and low nutrient habitats. Funct. Ecol. 15, 423–434 (2001)

    Article  Google Scholar 

  11. Grime, J. P. et al. Integrated screening validates primary axes of specialisation in plants. Oikos 79, 259–281 (1997)

    Article  Google Scholar 

  12. Whittaker, R. Communities and Ecosystems 2nd edn, 167 (MacMillan, New York, 1975)

    Google Scholar 

  13. Lambers, H., Chapin, F. S. & Pons, T. L. Plant Physiological Ecology (Springer, New York, 1998)

    Book  Google Scholar 

  14. Parkhurst, D. F. Diffusion of CO2 and other gases inside leaves. New Phytol. 126, 449–479 (1994)

    Article  CAS  Google Scholar 

  15. Evans, J. R. & Loreto, F. in Photosynthesis: Physiology and Metabolism (eds Leegood, R. C., Sharkey, T. D. & von Caemmerer, S.) 321–351 (Kluwer Academic, Dordrecht, 2000)

    Book  Google Scholar 

  16. Cannell, M. G. R. & Thornley, J. H. M. Modelling the components of plant respiration: Some guiding principles. Ann. Bot. 85, 45–54 (2000)

    Article  CAS  Google Scholar 

  17. Wright, I. J. & Westoby, M. Leaves at low versus high rainfall: coordination of structure, lifespan and physiology. New Phytol. 155, 403–416 (2002)

    Article  Google Scholar 

  18. Coley, P. D. Herbivory and defensive characteristics of tree species in a lowland tropical forest. Ecol. Monogr. 53, 209–233 (1983)

    Article  Google Scholar 

  19. Wright, I. J., Reich, P. B. & Westoby, M. Least-cost input mixtures of water and nitrogen for photosynthesis. Am. Nat. 161, 98–111 (2003)

    PubMed  Google Scholar 

  20. Farquhar, G. D., Buckley, T. N. & Miller, J. M. Optimal stomatal control in relation to leaf area and nitrogen content. Silva Fennica 36, 625–637 (2002)

    Article  Google Scholar 

  21. Reich, P. B., Walters, M. B. & Ellsworth, D. S. Leaf life-span in relation to leaf, plant, and stand characteristics among diverse ecosystems. Ecol. Monogr. 62, 365–392 (1992)

    Article  Google Scholar 

  22. Lambers, H. & Poorter, H. Inherent variation in growth rate between higher plants: a search for ecological causes and consequences. Adv. Ecol. Res. 23, 187–261 (1992)

    Article  CAS  Google Scholar 

  23. Wright, I. J. & Westoby, M. Nutrient concentration, resorption and lifespan: leaf traits of Australian sclerophyll species. Funct. Ecol. 17, 10–19 (2003)

    Article  Google Scholar 

  24. Sokal, R. R. & Rohlf, F. J. Biometry: the Principles and Practice of Statistics in Biological Research (W. H. Freeman, New York, 1995)

    MATH  Google Scholar 

  25. Koerselman, W. & Meuleman, A. F. M. The vegetation N/P ratio–a new tool to detect the nature of nutrient limitation. J. Appl. Ecol. 33, 1441–1450 (1996)

    Article  Google Scholar 

  26. Sterner, R. W. & Elser, J. J. Ecological Stoichiometry. The Biology of Elements from Molecules to the Biosphere (Princeton Univ. Press, Princeton/Oxford, 2002)

    Google Scholar 

  27. Westoby, M., Warton, D. & Reich, P. B. The time value of leaf area. Am. Nat. 155, 649–656 (2000)

    Article  Google Scholar 

  28. Grime, J. P. in Exploitation of Environmental Heterogeneity in Plants (eds Caldwell, M. M. & Pearcy, R. W.) 1–18 (Academic, San Diego, 1994)

    Book  Google Scholar 

  29. Harper, J. L. The value of a leaf. Oecologia 80, 53–58 (1989)

    Article  ADS  CAS  Google Scholar 

  30. Walters, M. B. & Reich, P. B. Seed size, nitrogen supply, and growth rate affect tree seedling survival in deep shade. Ecology 81, 1887–1901 (2000)

    Article  Google Scholar 

  31. Reich, P. B. & Walters, M. B. Photosynthesis-nitrogen relations in Amazonian tree species. 2. Variation in nitrogen vis-a-vis specific leaf area influences mass-based and area-based expressions. Oecologia 97, 73–81 (1994)

    Article  ADS  CAS  Google Scholar 

  32. Reich, P. B., Ellsworth, D. S. & Walters, M. B. Leaf structure (specific leaf area) modulates photosynthesis-nitrogen relations: evidence from within and across species and functional groups. Funct. Ecol. 12, 948–958 (1998)

    Article  Google Scholar 

  33. Schulze, E.-D., Kelliher, F. M., Körner, C., Lloyd, J. & Leuning, R. Relationships among maximum stomatal conductance, ecosystem surface conductance, carbon assimilation rate, and plant nitrogen nutrition: a global scaling exercise. Annu. Rev. Ecol. Syst. 25, 629–660 (1994)

    Article  Google Scholar 

  34. Peterson, A. G. CMEAL participants. Reconciling the apparent difference between mass- and area-based expressions of the photosynthesis-nitrogen relationship. Oecologia 118, 114–150 (1999)

    ADS  Google Scholar 

  35. Green, D. S. & Kruger, E. L. Light-mediated constraints on leaf function correlate with leaf structure among deciduous and evergreen tree species. Tree Physiol. 21, 1341–1346 (2001)

    Article  CAS  Google Scholar 

  36. Poorter, H. & Evans, J. R. Photosynthetic nitrogen-use efficiency of species that differ inherently in specific leaf area. Oecologia 116, 26–37 (1998)

    Article  ADS  Google Scholar 

  37. Hikosaka, K., Hanba, Y. T., Hirose, T. & Terashima, I. Photosynthetic nitrogen-use efficiency in leaves of woody and herbaceous species. Funct. Ecol. 12, 896–905 (1998)

    Article  Google Scholar 

  38. Maximov, N. A. The Plant in Relation to Water. A Study of the Physiological Basis of Drought Resistance (Allen & Unwin, London, 1929)

    Google Scholar 

  39. Schulze, E.-D. et al. Carbon and nitrogen isotope discrimination and nitrogen nutrition of trees along a rainfall gradient in northern Australia. Aust. J. Plant Physiol. 25, 413–425 (1998)

    Google Scholar 

  40. Fonseca, C. R., Overton, J. M., Collins, B. & Westoby, M. Shifts in trait combinations along rainfall and phosphorus gradients. J. Ecol. 88, 964–977 (2000)

    Article  Google Scholar 

  41. Villar, R. & Merino, J. Comparison of leaf construction costs in woody species with differing leaf life-spans in contrasting ecosystems. New Phytol. 151, 213–226 (2001)

    Article  Google Scholar 

  42. Oleksyn, J., Reich, P. B., Zytkowiak, R., Karolewski, P. & Tjoelker, M. G. Nutrient conservation increases with latitude of origin in European Pinus sylvestris populations. Oecologia 136, 220–235 (2003)

    Article  ADS  CAS  Google Scholar 

  43. Ackerly, D. D. & Bazzaz, F. A. Leaf dynamics, self-shading and carbon gain in seedlings of a tropical pioneer tree. Oecologia 101, 289–298 (1995)

    Article  ADS  CAS  Google Scholar 

  44. Bonan, G. B., Levis, S., Kergoat, L. & Oleson, K. W. Landscapes as patches of plant functional types: an integrating concept for climate and ecosystem models. Glob. Biogeochem. Cycles 16, doi:10.1029/2000GB001360 (2002)

  45. Moorcroft, P. R., Hurtt, G. C. & Pacala, S. W. A method for scaling vegetation dynamics: The ecosystem demography model (ED). Ecol. Monogr. 71, 557–585 (2001)

    Article  Google Scholar 

  46. Körner, C. Alpine Plant Life: Functional Plant Ecology of High Mountain Ecosystems Ch. 3 (Springer, Berlin, 1999)

    Book  Google Scholar 

  47. New, M., Hulme, M. & Jones, P. Representing twentieth-century space-time climate variability. Part I: Development of a 1961–90 mean monthly terrestrial climatology. J. Clim. 12, 829–856 (1999)

    Article  ADS  Google Scholar 

  48. Choudhury, B. J. Global pattern of potential evaporation calculated from the Penman-Monteith equation using satellite and assimilated data. Remote Sens. Environ. 61, 64–81 (1997)

    Article  ADS  Google Scholar 

  49. Barry, R. G. Mountain Weather and Climate 24–29 (Methuen, London/New York, 1981)

    Google Scholar 

  50. Campbell, G. S. & Norman, J. M. An Introduction to Environmental Biophysics (Springer, New York, 1998)

    Book  Google Scholar 

Download references

Acknowledgements

All authors contributed (mostly unpublished) data and intellectual input to the project. The first three authors took the lead in the organization, analysis and writing-up of this work, and contributed 8% of the data, with 223 species from 11 sites. We also thank the many other researchers who provided additional information about their study sites and published data.

Author information

Authors and Affiliations

Authors

Corresponding author

Correspondence to Ian J. Wright.

Ethics declarations

Competing interests

The authors declare that they have no competing financial interests.

Supplementary information

Supplementary Information

This consists of: 1. a list of published literature sources; 2. further details of bivariate trait relationships allowing formulation of predictive regression equations; 3. details of multiple regression analyses mentioned in the text; 4. PCA loadings for the area-based 5 trait analysis. (DOC 320 kb)

Supplementary Data

Glopnet dataset (XLS 747 kb)

Rights and permissions

Reprints and permissions

About this article

Cite this article

Wright, I., Reich, P., Westoby, M. et al. The worldwide leaf economics spectrum. Nature 428, 821–827 (2004). https://doi.org/10.1038/nature02403

Download citation

  • Received:

  • Accepted:

  • Issue Date:

  • DOI: https://doi.org/10.1038/nature02403

This article is cited by

Comments

By submitting a comment you agree to abide by our Terms and Community Guidelines. If you find something abusive or that does not comply with our terms or guidelines please flag it as inappropriate.

Search

Quick links

Nature Briefing

Sign up for the Nature Briefing newsletter — what matters in science, free to your inbox daily.

Get the most important science stories of the day, free in your inbox. Sign up for Nature Briefing