Lateral Diffusion of CO2 in Leaves Is Not Sufficient to Support Photosynthesis

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Abstract
Lateral diffusion of CO2 was investigated in photosynthesizing leaves with different anatomy by gas exchange and chlorophyll a fluorescence imaging using grease to block stomata. When one-half of the leaf surface of the heterobaric species Helianthus annuus was covered by 4-mm-diameter patches of grease, the response of net CO2 assimilation rate (A) to intercellular CO2 concentration (Ci) indicated that higher ambient CO2 concentrations (Ca) caused only limited lateral diffusion into the greased areas. When single 4-mm patches were applied to leaves of heterobaric Phaseolus vulgaris and homobaric Commelina communis, chlorophyll a fluorescence images showed dramatic declines in the quantum efficiency of photosystem II electron transport (measured as Fq′/Fm′) across the patch, demonstrating that lateral CO2 diffusion could not support A. The Fq′/Fm′ values were used to compute images of Ci across patches, and their dependence on Ca was assessed. At high Ca, the patch effect was less in C. communis than P. vulgaris. A finite-volume porous-medium model for assimilation rate and lateral CO2 diffusion was developed to analyze the patch images. The model estimated that the effective lateral CO2 diffusion coefficients inside C. communis and P. vulgaris leaves were 22% and 12% of that for free air, respectively. We conclude that, in the light, lateral CO2 diffusion cannot support appreciable photosynthesis over distances of more than approximately 0.3 mm in normal leaves, irrespective of the presence or absence of bundle sheath extensions, because of the CO2 assimilation by cells along the diffusion pathway.