Internal CO2 Measured Directly in Leaves

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Abstract
Observations of nonuniform photosynthesis across leaves cast doubt on internal CO2 partial pressures (pi) calculated on the assumption of uniformity and can lead to incorrect conclusions about the stomatal control of photosynthesis. The problem can be avoided by measuring pi directly because the assumptions of uniformity are not necessary. We therefore developed a method that allowed pi to be measured continuously in situ for days at a time under growth conditions and used it to investigate intact leaves of sunflower (Helianthus annuus L.), soybean (Glycine max L. Merr.), and bush bean (Phaseolus vulgaris L.) subjected to high or low leaf water potentials (ψw) or high concentrations of abscisic acid (ABA). The leaves maintained a relatively constant differential (Δp) between ambient CO2 and measured pi throughout the light period when water was supplied. When water was withheld, ψw decreased and the stomata began to close, but measured pi increased until the leaf reached a ψw of −1.76 (bush bean), −2.12 (sunflower) or −3.10 (soybean) megapascals, at which point Δp = 0. The increasing pi indicated that stomata did not inhibit CO2 uptake and a Δp of zero indicated that CO2 uptake became zero despite the high availability of CO2 inside the leaf. In contrast, when sunflower leaves at high ψw were treated with ABA, pi did not increase and instead decreased rapidly and steadily for up to 8 hours even as ψw increased, as expected if ABA treatment primarily affected stomatal conductance. The accumulating CO2 at low ψw and contrasting response to ABA indicates that photosynthetic biochemistry limited photosynthesis at low ψw but not at high ABA.