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Global leaf trait estimates biased due to plasticity in the shade


The study of leaf functional trait relationships, the so-called leaf economics spectrum1,2, is based on the assumption of high-light conditions (as experienced by sunlit leaves). Owing to the exponential decrease of light availability through canopies, however, the vast majority of the world's vegetation exists in at least partial shade. Plant functional traits vary in direct dependence of light availability3, with different traits varying to different degrees, sometimes in conflict with expectations from the economic spectrum3. This means that the derived trait relationships of the global leaf economic spectrum are probably dependent on the extent to which observed data in existing large-scale plant databases represent high-light conditions. Here, using an extensive worldwide database of within-canopy gradients of key physiological, structural and chemical traits3, along with three different global trait databases4,5, we show that: (1) accounting for light-driven trait plasticity can reveal novel trait relationships, particularly for highly plastic traits (for example, the relationship between net assimilation rate per area (Aa) and leaf mass per area (LMA)); and (2) a large proportion of leaf traits in current global plant databases reported as measured in full sun were probably measured in the shade. The results show that even though the majority of leaves exist in the shade, along with a large proportion of observations, our current understanding is too focused on conditions in the sun.

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Figure 1: Comparison of trait values from different databases.
Figure 2: The effective shading by trait and plant functional type.
Figure 3: The effect of plasticity on the leaf economic spectrum.


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T.F.K. acknowledges the financial support from the Laboratory Directed Research and Development (LDRD) fund under the auspices of Department of Energy, Biological and Environmental Research Office of Science at Lawrence Berkeley National Laboratory. Ü.N. acknowledges the European Regional Development Fund (Centre of Excellence EcolChange) and the Estonian Ministry of Science and Education (institutional grant IUT-8-3). The authors acknowledge useful feedback from W. Han on an earlier version of the manuscript.

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T.F.K. and Ü.N. conceived the analysis. T.F.K. performed the analysis and both authors participated in drafting the manuscript.

Corresponding author

Correspondence to Trevor F. Keenan.

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The authors declare no competing financial interests.

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Keenan, T., Niinemets, Ü. Global leaf trait estimates biased due to plasticity in the shade. Nature Plants 3, 16201 (2017).

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