This paper reports the results of two experiments undertaken to investigate the influence of season and soil drying on stomatal responses to leaf-to-air vapour pressure differences. We examined the response of stomatal conductance to increasing leaf-to-air vapour pressure difference, in the wet and dry seasons, of five tropical tree species. We also examined leaves of these species for anatomical differences to determine whether this could explain differences in stomatal sensitivity to leaf-to-air vapour pressure differences. Finally, we conducted a split-root experiment with one of those species to look for interactions between xylem abscisic acid concentration, predawn water potential, leaf area to root mass ratio and stomatal responses to leaf-to-air vapour pressure differences. Stomatal conductance declined linearly with increasing leaf-to-air vapour pressure difference in all species. Leaves that expanded in the ‘dry’ season were more sensitive to leaf-to-air vapour pressure differences than those that had expanded in the ‘wet’ season. The value of leaf-to-air vapour pressure difference where 50% of extrapolated maximum stomatal conductance would occur was 5.5 kPa for wet season but only 3.4 kPa for dry season leaves. In the wet season, transpiration rate increased with increasing leaf-to-air vapour pressure difference in most example species. However, in the dry season, transpiration was constant as leaf-to-air vapour pressure differences increased in most cases. There were significant changes in the proportion of cell wall exposed to air space in leaves, between wet and dry seasons, in three of four species examined. In the split-root experiment, a very mild water stress increased stomatal sensitivity to leaf-to-air vapour pressure differences, and stomatal conductivity declined linearly with decreasing predawn water potential. However, levels of ABA in the xylem did not change, and stomatal sensitivity to exogenous ABA did not change. The ratio of leaf area to root mass declined during water stress and was correlated to changes in stomatal sensitivity to leaf-to-air vapour pressure differences.

BackgroundStomata respond to vapour pressure deficit (D) – when D increases, stomata begin to close. Closure is the result of a decline in guard cell turgor, but the link between D and turgor is poorly understood. We describe a model for stomatal responses to increasing D based upon cellular water relations. The model also incorporates impacts of increasing levels of water stress upon stomatal responses to increasing D.ResultsThe model successfully mimics the three phases of stomatal responses to D and also reproduces the impact of increasing plant water deficit upon stomatal responses to increasing D. As water stress developed, stomata regulated transpiration at ever decreasing values of D. Thus, stomatal sensitivity to D increased with increasing water stress. Predictions from the model concerning the impact of changes in cuticular transpiration upon stomatal responses to increasing D are shown to conform to experimental data.Sensitivity analyses of stomatal responses to various parameters of the model show that leaf thickness, the fraction of leaf volume that is air-space, and the fraction of mesophyll cell wall in contact with air have little impact upon behaviour of the model. In contrast, changes in cuticular conductance and membrane hydraulic conductivity have significant impacts upon model behaviour.ConclusionCuticular transpiration is an important feature of stomatal responses to D and is the cause of the 3 phase response to D. Feed-forward behaviour of stomata does not explain stomatal responses to D as feedback, involving water loss from guard cells, can explain these responses.