Stomatal Metabolism: Carbon Dioxide Fixation in Attached and Detached Epidermis of Commelina

Rates of accumulation of radioactivity and the nature of 14CO2 fixation products were measured in mesophyll, attached epidermis and detached epidermis of Commelina cyanea and C. communis. In the illuminated detached epidermis of C. cyanea, most of the fixation products were malate and aspartate (in almost equal proportions), with small amounts of sugars, sugar phosphates, serine, glycine, alanine and TCA cycle intermediates. In that of C. communis there was a smaller proportion of aspartate and a higher proportion of sugars, glutamate and tricarboxylic acids. The much higher rates of accumulation of labelled fixation products in attached epidermis of C. cyanea can in part be attributed to glycine, serine and alanine, which appear to be imported from the mesophyll very shortly after the leaf is first exposed to 14CO2. Over longer periods of time, labelled sugars contributed an appreciable and increasing proportion. In C. communis, after 15-30 min, most of the difference between attached and detached epidermis was attributable to the presence of labelled sugars. The fixation pattern in the mesophyll of these species was typical of C*3-type photosynthesis. Autoradiographs of detached epidermis showed that the label was predominantly in stomata while those of attached epidermis showed more label in stomata than elsewhere after 1 min and they were uniformly labelled after 30 min. These findings suggest that metabolites are translocated from the mesophyll to the epidermis fairly readily. There is probably flow in the reverse direction as well as gaseous exchange of 14C between these tissues.