Strong overestimation of water-use efficiency responses to rising CO in tree-ring studies.

The carbon isotope ratio (δ C) in tree rings is commonly used to derive estimates of the assimilation-to-stomatal conductance rate of trees, that is, intrinsic water-use efficiency (iWUE). Recent studies have observed increased iWUE in response to rising atmospheric CO concentrations (C ), in many different species, genera and biomes. However, increasing rates of iWUE vary widely from one study to another, likely because numerous covarying factors are involved. Here, we quantified changes in iWUE of two widely distributed boreal conifers using tree samples from a forest inventory network that were collected across a wide range of growing conditions (assessed using the site index, SI), developmental stages and stand histories. Using tree-ring isotopes analysis, we assessed the magnitude of increase in iWUE after accounting for the effects of tree size, stand age, nitrogen deposition, climate and SI. We also estimated how growth conditions have modulated tree physiological responses to rising C . We found that increases in tree size and stand age greatly influenced iWUE. The effect of C on iWUE was strongly reduced after accounting for these two variables. iWUE increased in response to C , mostly in trees growing on fertile stands, whereas iWUE remained almost unchanged on poor sites. Our results suggest that past studies could have overestimated the CO effect on iWUE, potentially leading to biased inferences about the future net carbon balance of the boreal forest. We also observed that this CO effect is weakening, which could affect the future capacity of trees to resist and recover from drought episodes.