Photocontrol of the Functional Coupling between Photosynthesis and Stomatal Conductance in the Intact Leaf

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Abstract
The photocontrol of the functional coupling between photosynthesis and stomatal conductance in the leaf was investigated in gas exchange experiments using monochromatic light provided by lasers. Net photosynthesis and stomatal conductance were measured in attached leaves of Malva parviflora L. as a function of photon irradiance at 457.9 and 640.0 nanometers. Photosynthetic rates and quantum yields of photosynthesis were higher under red light than under blue, on an absorbed or incident basis. Stomatal conductance was higher under blue than under red light at all intensities. Based on a calculated apparent photon efficiency of conductance, blue and red light had similar effects on conductance at intensities higher than 0.02 millimoles per square meter per second, but blue light was several-fold more efficient at very low photon irradiances. Red light had no effect on conductance at photon irradiances below 0.02 millimoles per square meter per second. These observations support the hypothesis that stomatal conductance is modulated by two photosystems: a blue light-dependent one, driving stomatal opening at low light intensities and a photosynthetically active radiation (PAR)-dependent one operating at higher irradiances. When low intensity blue light was used to illuminate a leaf already irradiated with high intensity, 640 nanometers light, the leaf exhibited substantial increases in stomatal conductance. Net photosynthesis changed only slightly. Additional far-red light increased net photosynthesis without affecting stomatal conductance. These observations indicate that under conditions where the PAR-dependent system is driven by high intensity red light, the blue light-dependent system has an additive effect on stomatal conductance. The wavelength dependence of photosynthesis and stomatal conductance demonstrates that these processes are not obligatorily coupled and can be controlled by light, independent of prevailing levels of intercellular CO2. The blue light-dependent system in the guard cells may function as a specific light sensor while the PAR-dependent system supplies a CO2-modulated energy source providing functional coupling between the guard cells and the photosynthesizing mesophyll.