The effects of soil and atmospheric drought on photosynthesis and stomatal control of gas exchange in three coniferous species

Abstract
The responses of steady‐state CO2 assimilation rate (A), transpiration rate (E), and stomatal conductance (gs) to changes in leaf‐to‐air vapour pressure difference (δW) on one hand and to increasing soil drought on the other hand were examined in 2‐year‐old seedlings of Pseudotsuga menziesii, Pseudotsuga macrocarpa and Cedrus atlantica. Analysing the data through A vs intercellular CO2 molar fraction (ci) graphs, we could determine stomatal and mesophyll contributions to changes in A as δW or soil drought were increased. Increasing soil drought affected gs and mesophyll photosynthesis independently, since clearly distinct predawn leaf water potential (ψp) regions appeared in which either stomatal or mesophyll effects prevailed for explaining the changes in A. The two Pseudotsuga species exhibited a large ψP range (between ca ‐0.8 and ‐1.5 to ‐1.9 MPa) in which only stomata were responsible for the decrease in A. A dramatic decline in mesophyll photosynthesis was noticed starting from values as high as ‐1.2 MPa (C. atlantica), ‐1.5 MPa (P. macrocarpa) and ‐1.9 MPa (P. menziesii). Increasing ΔW at high soil water content led to a sharp decline in A primarily due to an alteration of mesophyll photosynthesis. Stomatal conductance for CO2 diffusion was affected in a lesser extent and in close correlation with the changes in mesophyll photosynthesis, which could suggest the existence of a functional linkage between mesophyll photosynthesis and stomata. Surprisingly, the drought resistant P. macrocarpa exhibited the least conservative water use efficiency in response to the two types of drought. In this species drought adaptation seems to be mainly due to its high root growth and soil prospection ability.