Stomatal conductance and not stomatal density determines the long-term reduction in leaf transpiration of poplar in elevated CO2

Using a free-air CO2 enrichment (FACE) experiment, poplar trees (Populus · euramericana clone I214) were exposed to either ambient or elevated [CO2] from planting, for a 5-year period during canopy development, closure, coppice and re-growth. In each year, measurements were taken of stomatal density (SD, number mm2) and stomatal index (SI, the proportion of epidermal cells forming stomata). In year 5, measurements were also taken of leaf stomatal conductance (gs, lmol m2 s1), photosynthetic CO2 fixation (A, mmol m2 s1), instantaneous water-use efficiency (A/E) and the ratio of intercellular to atmospheric CO2 (Ci:Ca). Elevated [CO2] caused reductions in SI in the first year, and in SD in the first 2 years, when the canopy was largely open. In following years, when the canopy had closed, elevated [CO2] had no detectable effects on stomatal numbers or index. In contrast, even after 5 years of exposure to elevated [CO2], gs was reduced, A/E was stimulated, and Ci:Ca was reduced relative to ambient [CO2]. These outcomes from the long-term realistic field conditions of this forest FACE experiment suggest that stomatal numbers (SD and SI) had no role in determining the improved instantaneous leaf-level efficiency of water use under elevated [CO2]. We propose that altered cuticular development during canopy closure may partially explain the changing response of stomata to elevated [CO2], although the mechanism for this remains obscure.

Towards an improved and more flexible representation of water stress in coupled photosynthesis–stomatal conductance models

Coupled photosynthesis–stomatal conductance (A–gs) models are commonly used in ecosystem models to represent the exchange rate of CO2 and H2O between vegetation and the atmosphere. The ways these models account for water stress differ greatly among modelling schemes. This study provides insight into the impact of contrasting model configurations of water stress on the simulated leaf-level values of net photosynthesis (A), stomatal conductance (gs), the functional relationship among them and their ratio, the intrinsic water use efficiency (A/gs), as soil dries. A simple, yet versatile, normalized soil moisture dependent function was used to account for the effects of water stress on gs, on mesophyll conductance (gm) and on the biochemical capacity. Model output was compared to leaf-level values obtained from the literature. The sensitivity analyses emphasized the necessity to combine both stomatal and non-stomatal limitations of A in coupled A–gs models to accurately capture the observed functional relationships A vs. gs and A/gsvs. gs in response to drought. Accounting for water stress in coupled A–gs models by imposing either stomatal or biochemical limitations of A, as commonly practiced in most ecosystem models, failed to reproduce the observed functional relationship between key leaf gas exchange attributes. A quantitative limitation analysis revealed that the general pattern of C3 photosynthetic response to water stress may be well represented in coupled A–gs models by imposing the highest limitation strength to gm, then to gs and finally to the biochemical capacity.