Plant stomata display a wide range of short-term behavioural and long-term morphological responses to atmospheric carbon dioxide concentration ([CO(2)]). 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 (G(s)) to instantaneous changes in external [CO(2)] (C(a)) in an evolutionary cross-section of vascular plants grown in atmospheres of elevated [CO(2)] (1,500 ppm) and sub-ambient [O(2)] (13.0 %) compared to control conditions (380 ppm CO(2), 20.9 % O(2)). We found that active control of stomatal aperture to [CO(2)] 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 C(a) were more likely to exhibit a reduction in stomatal density than species with active stomatal control when grown in atmospheres of elevated [CO(2)]. The relationship between the degree of stomatal aperture control to C(a) above ambient and the extent of any reduction in stomatal density may suggest the co-ordination of physiological and morphological responses of stomata to [CO(2)] 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.

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One strategy for plants to optimize stomatal function is to open and close their stomata quickly in response to environmental signals. It is generally assumed that small stomata can alter aperture faster than large stomata. We tested the hypothesis that species with small stomata close faster than species with larger stomata in response to darkness by comparing rate of stomatal closure across an evolutionary range of species including ferns, cycads, conifers, and angiosperms under controlled ambient conditions (380 ppm CO2; 20.9% O2). The two species with fastest half-closure time and the two species with slowest half-closure time had large stomata while the remaining three species had small stomata, implying that closing rate was not correlated with stomatal size in these species. Neither was response time correlated with stomatal density, phylogeny, functional group, or life strategy. Our results suggest that past atmospheric CO2 concentration during time of taxa diversification may influence stomatal response time. We show that species which last diversified under low or declining atmospheric CO2 concentration close stomata faster than species that last diversified in a high CO2 world. Low atmospheric [CO2] during taxa diversification may have placed a selection pressure on plants to accelerate stomatal closing to maintain adequate internal CO2 and optimize water use efficiency.

BackgroundPlant growth chambers provide a controlled environment to analyse the effects of environmental parameters (light, temperature, atmospheric gas composition etc.) on plant function. However, it has been shown that a ‘chamber effect’ may exist whereby results observed are not due to an experimental treatment but to inconspicuous differences in supposedly identical chambers. In this study, Vicia faba L. ‘Aquadulce Claudia’ (broad bean) plants were grown in eight walk-in chambers to establish if a chamber effect existed, and if so, what plant traits are best for detecting such an effect. A range of techniques were used to measure differences between chamber plants, including chlorophyll fluorescence measurements, gas exchange analysis, biomass, reproductive yield, anatomical traits and leaf stable carbon isotopes.Results and discussionFour of the eight chambers exhibited a chamber effect. In particular, we identified two types of chamber effect which we term ‘resolvable’ or ‘unresolved’; a resolvable chamber effect is caused by malfunctioning components of a chamber and an unresolved chamber effect is caused by unknown factors that can only be mitigated by appropriate experimental design and sufficient replication. Not all measured plant traits were able to detect a chamber effect and no single trait was capable of detecting all chamber effects. Fresh weight and flower count detected a chamber effect in three chambers, stable carbon isotopes (δ13C) and net rate CO2 assimilation (An) identified a chamber effect in two chambers, stomatal conductance (gs) and total performance index detected an effect only in one chamber.Conclusion(1) Chamber effects can be adequately detected by fresh weight measurements and flower counts on Vicia faba plants. These methods were the most effective in terms of detection and most efficient in terms of time. (2) δ13C, gs and An measurements help distinguish between resolvable and unresolved chamber effects. (3) Unresolved chamber effects require experimental unit replication while resolvable chamber effects require investigation, repair and retesting in advance of initiating further experiments.

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