Co-ordination of physiological and morphological responses of stomata to elevated [CO2] in vascular plants

Plant stomata display a wide range of short-term behavioural and long-term morphological responses to atmospheric carbon dioxide concentration ([CO2]). The diversity of responses suggests that plants may have different strategies for controlling gas exchange, yet it is not known whether these strategies are co-ordinated in some way. Here, we test the hypothesis that there is co-ordination of physiological (via aperture change) and morphological (via stomatal density change) control of gas exchange by plants. We examined the response of stomatal conductance (Gs) to instantaneous changes in external [CO2] (Ca) in an evolutionary cross-section of vascular plants grown in atmospheres of elevated [CO2] (1,500 ppm) and sub-ambient [O2] (13.0 %) compared to control conditions (380 ppm CO2, 20.9 % O2). We found that active control of stomatal aperture to [CO2] above current ambient levels was not restricted to angiosperms, occurring in the gymnosperms Lepidozamia peroffskyana and Nageia nagi. The angiosperm species analysed appeared to possess a greater respiratory demand for stomatal movement than gymnosperm species displaying active stomatal control. Those species with little or no control of stomatal aperture (termed passive) to Ca were more likely to exhibit a reduction in stomatal density than species with active stomatal control when grown in atmospheres of elevated [CO2]. The relationship between the degree of stomatal aperture control to Ca above ambient and the extent of any reduction in stomatal density may suggest the co-ordination of physiological and morphological responses of stomata to [CO2] in the optimisation of water use efficiency. This trade-off between stomatal control strategies may have developed due to selective pressures exerted by the costs associated with passive and active stomatal control.