The Earth’s carbon and hydrologic cycles are intimately coupled by gas exchange through plant stomata. However, uncertainties in the magnitude and consequences of the physiological responses of plants to elevated CO2 in natural environments hinders modelling of terrestrial water cycling and carbon storage. Here we use annually resolved long-term 13C tree-ring measurements across a European forest network to reconstruct the physiologically driven response of intercellular CO2 (Ci) caused by atmospheric CO2 (Ca) trends. When removing meteorological signals from the 13C measurements, we find that trees across Europe regulated gas exchange so that for one ppmv atmospheric CO2 increase, Ci increased by 0.76 ppmv, most consistent with moderate control towards a constant Ci=Ca ratio. This response corresponds to twentieth-century intrinsic water-use efficiency (iWUE) increases of 14 ±10 and 22 ± 6% at broadleaf and coniferous sites, respectively. An ensemble of process-based global vegetation models shows similar CO2 effects on iWUE trends. Yet, when operating these models with climate drivers reintroduced, despite decreased stomatal opening, 5%increases in European forest transpiration are calculated over the twentieth century.This counterintuitive result arises from lengthened growing seasons, enhanced evaporative demand in a warming climate, and increased leaf area, which together oppose effects of CO2-induced stomatal closure. Our study questions changes to the hydrological cycle, such as reductions in transpiration and air humidity, hypothesized to result from plant responses to anthropogenic emissions

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The increasing carbon dioxide (CO 2 ) concentration in the atmosphere in combination with climatic changes throughout the last century are likely to have had a profound effect on the physiology of trees: altering the carbon and water fluxes passing through the stomatal pores.However, the magnitude and spatial patterns of such changes in natural forests remain highly uncertain. Here, stable carbon isotope ratios from a network of 35 tree-ring sites located Central Europe, a region where summer soil-water availability decreased over the last century.We were able to demonstrate that the combined effects of increasing CO 2 and climate change leading to soil drying have resulted in an accelerated increase of iWUE. These findings will help to reduce uncertainties in the land surface schemes of global climate models, where vegetation-climate feedbacks are currently still poorly constrained by observational data. 4