USGS2016
Rising atmospheric [CO 2 ], c a , is expected to affect stomatal regulation of leaf gas-exchange of woody plants, thus influencing energy fluxes as well as carbon (C), water, and nutrient cycling of forests. Researchers have proposed various strategies for stomatal regulation of leaf gas-exchange that include maintaining a constant leaf internal [CO 2 ], c i , a constant drawdown in CO 2 ( c a − c i ), and a constant c i / c a . These strategies can result in drastically different consequences for leaf gas-exchange. The accuracy of Earth systems models depends in part on assumptions about generalizable patterns in leaf gas-exchange responses to varying c a . The concept of optimal stomatal behavior, exemplified by woody plants shifting along a continuum of these strategies, provides a unifying framework for understanding leaf gas-exchange responses to c a . To assess leaf gas-exchange regulation strategies, we analyzed patterns in c i inferred from studies reporting C stable isotope ratios (δ 13 C) or photosynthetic discrimination (∆) in woody angiosperms and gymnosperms that grew across a range of c a spanning at least 100 ppm. Our results suggest that much of the c a -induced changes in c i / c a occurred across c a spanning 200 to 400 ppm. These patterns imply that c a − c i will eventually approach a constant level at high c a because assimilation rates will reach a maximum and stomatal conductance of each species should be constrained to some minimum level. These analyses are not consistent with canalization toward any single strategy, particularly maintaining a constant c i . Rather, the results are consistent with the existence of a broadly conserved pattern of stomatal optimization in woody angiosperms and gymnosperms. This results in trees being profligate water users at low c a , when additional water loss is small for each unit of C gain, and increasingly water-conservative at high c a , when photosystems are saturated and water loss is large for each unit C gain.