Variation in leaf and twig CO2 flux as a function of plant size: a comparison of seedlings, saplings and trees.

Rates of tissue-level function have been hypothesized to decline as trees grow older and larger, but relevant evidence to assess such changes remains limited, especially across a wide range of sizes from saplings to large trees. We measured functional traits of leaves and twigs of three cold-temperate deciduous tree species in Minnesota, USA, to assess how these vary with tree height. Individuals ranging from 0.13 to 20 m in height were sampled in both relatively open and closed canopy environments to minimize light differences as a potential driver of size-related differences in leaf and twig properties. We hypothesized that (H1) gas-exchange rates, tissue N concentration and leaf mass per unit area (LMA) would vary with tree size in a pattern reflecting declining function in taller trees, yet maintaining (H2) bivariate trait relations, common among species as characterized by the leaf economics spectrum. Taking these two ideas together yielded a third, integrated hypothesis that (H3) nitrogen (N) content and gas-exchange rates should decrease monotonically with tree size and LMA should increase. We observed increasing LMA and decreasing leaf and twig Rd with increasing size, which matched predictions from H1 and H3. However, opposite to our predictions, leaf and twig N generally increased with size, and thus had inverse relations with respiration, rather than the predicted positive relations. Two exceptions were area-based leaf N of Prunus serotina Ehrh. in gaps and mass-based leaf N of Quercus ellipsoidalis E. J. Hill in gaps, both of which showed qualitatively hump-shaped patterns. Finally, we observed hump-shaped relationships between photosynthetic capacity and tree height, not mirroring any of the other traits, except in the two cases highlighted above. Bivariate trait relations were weak intra-specifically, but were generally significant and positive for area-based traits using the pooled dataset. Results suggest that different traits vary with tree size in different ways that are not consistent with a universal shift towards a lower 'return on investment' strategy. Instead, species traits vary with size in patterns that likely reflect complex variation in water, light, nitrogen and carbon availability, storage and use.