Leaf nitrogen from first principles : field evidence for adaptive variation with climate

Nitrogen content per unit leaf area (Nₐᵣₑₐ) is a key variable in plant functional ecology and biogeochemistry. Nₐᵣₑₐ comprises a structural component, which scales with leaf mass per area (LMA), and a metabolic component, which scales with Rubisco capacity. The co-ordination hypothesis, as implemented in LPJ and related global vegetation models, predicts that Rubisco capacity should be directly proportional to irradiance but should decrease with increases in cᵢ : cₐ and temperature because the amount of Rubisco required to achieve a given assimilation rate declines with increases in both. We tested these predictions using LMA, leaf δ¹³C, and leaf N measurements on complete species assemblages sampled at sites on a north–south transect from tropical to temperate Australia. Partial effects of mean canopy irradiance, mean annual temperature, and cᵢ : cₐ (from δ¹³C) on Nₐᵣₑₐ were all significant and their directions and magnitudes were in line with predictions. Over 80 % of the variance in community-mean (ln) Nₐᵣₑₐ was accounted for by these predictors plus LMA. Moreover, Nₐᵣₑₐ could be decomposed into two components, one proportional to LMA (slightly steeper in N-fixers), and the other to Rubisco capacity as predicted by the co-ordination hypothesis. Trait gradient analysis revealed cᵢ : cₐ to be perfectly plastic, while species turnover contributed about half the variation in LMA and Nₐᵣₑₐ. Interest has surged in methods to predict continuous leaf-trait variation from environmental factors, in order to improve ecosystem models. Coupled carbon–nitrogen models require a method to predict Nₐᵣₑₐ that is more realistic than the widespread assumptions that Nₐᵣₑₐ is proportional to photosynthetic capacity, and/or that Nₐᵣₑₐ (and photosynthetic capacity) are determined by N supply from the soil. Our results indicate that Nₐᵣₑₐ has a useful degree of predictability, from a combination of LMA and cᵢ : cₐ – themselves in part environmentally determined – with Rubisco activity, as predicted from local growing conditions. This finding is consistent with a "plant-centred" approach to modelling, emphasizing the adaptive regulation of traits. Models that account for biodiversity will also need to partition community-level trait variation into components due to phenotypic plasticity and/or genotypic differentiation within species vs. progressive species replacement, along environmental gradients. Our analysis suggests that variation in Nₐᵣₑₐ is about evenly split between these two modes.15 page(s