Explaining global increases in water use efficiency: why have we overestimated responses to rising atmospheric CO(2) in natural forest ecosystems?

Abstract

The analysis of tree-ring carbon isotope composition (δ(13)C) has been widely used to estimate spatio-temporal variations in intrinsic water use efficiency (iWUE) of tree species. Numerous studies have reported widespread increases in iWUE coinciding with rising atmospheric CO(2) over the past century. While this could represent a coherent global response, the fact that increases of similar magnitude were observed across biomes with no apparent effect on tree growth raises the question of whether iWUE calculations reflect actual physiological responses to elevated CO(2) levels.Here we use Monte Carlo simulations to test if an artifact of calculation could explain observed increases in iWUE. We show that highly significant positive relationships between iWUE and CO(2) occur even when simulated data (randomized δ(13)C values spanning the observed range) are used in place of actual tree-ring δ(13)C measurements. From simulated data sets we calculated non-physiological changes in iWUE from 1900 to present and across a 4000 m altitudinal range. This generated results strikingly similar to those reported in recent studies encompassing 22 species from tropical, subtropical, temperate, boreal and mediterranean ecosystems. Only 6 of 49 surveyed case studies showed increases in iWUE significantly higher than predicted from random values.Our results reveal that increases in iWUE estimated from tree-ring δ(13)C occur independently of changes in (13)C discrimination that characterize physiological responses to elevated CO(2). Due to a correlation with CO(2) concentration, which is used as an independent factor in the iWUE calculation, any tree-ring δ(13)C data set would inevitably generate increasing iWUE over time. Therefore, although consistent, previously reported trends in iWUE do not necessarily reflect a coherent global response to rising atmospheric CO(2). We discuss the significance of these findings and suggest ways to distinguish real from artificial responses in future studies.