Stomatal opening ratio mediates trait coordinating network adaptation to environmental gradients

A trait coordination network is constructed through intercorrelations of functional traits, which reflect trait-based adaptive strategies. However, little is known about how these networks change across spatial scales, and what drivers and mechanisms mediate this change. This study bridges that gap by integrating functional traits related to plant carbon gain and water economy into the coordination network of Siberian elm (Ulmus pumila), a eurybiont that survives along a 3800 km environmental gradient from humid forest to arid desert. Our results demonstrated that both stomatal density and stomatal size reached a physiological threshold at which adjustments in these traits were not sufficient to cope with the increased environmental stress. Network analysis further revealed that the mechanism for overcoming this threshold, the stomatal opening ratio, g_ratio , was represented by the highest values for centrality across different spatial scales, and therefore mediated the changes in the trait coordination network along environmental gradients. The mediating roles manifested as creating the highest maximum theoretical stomatal conductance (g_smax ) but lowest possible g_ratio for pathogen defense in humid regions, while maintaining the g_ratio 'sweet spot' (c. 20% in this region) for highest water use efficiency in semihumid regions, and having the lowest g_smax and highest g_ratio for gas exchange and leaf cooling in arid regions. These results suggested that the stomatal traits related to control of stomatal movement play fundamental roles in balancing gas exchange, leaf cooling, embolism resistance and pathogen defense. These insights will allow more accurate model parameterization for different regions, and therefore better predictions of species' responses to global change.