Growth, structure, stomatal responses and secondary metabolites of birch seedlings (Betula pendula) under elevated UV-B radiation in the field

Three-year-old birch (Betula pendula Roth.) seedlings were exposed, in the field, to supplemental levels of UV-B radiation. Control seedlings were exposed to ambient levels of UV radiation, using arrays of unenergized lamps. A control for UV-A radiation was also included in the experiment. Enhanced UV-B radiation had no significant effects on height growth, and shoot and root biomass of birch seedlings. Leaf expansion rate increased transiently in the middle of the growing period in enhanced UV-B- and UV-A-exposed plants; however, final leaf size and relative growth rate remained unaffected. Leaf thickness and spongy intercellular spaces were increased in UV-B-exposed seedlings along with increased density of glandular trichomes. At the ultrastructural level, enhanced UV-B increased the number of cytoplasmic lipid bodies, and abnormal membrane whorls were found. Both enhanced UV-B and UV-A radiation induced swelling of chloroplast thylakoids. Stomatal density and conductance were significantly increased by elevated UV-B radiation. UV-A radiation increased the length and width of stomata, whereas UV-B radiation had only a marginal effect on stomatal size. UV-A and enhanced UV-B radiation attenuated the appearance of necrotic spots in autumn, probably caused by the fungus Pyrenopeziza betulicola, suggesting a direct harmful effect of UV on pathogens or reduced susceptibility to pathogens in UV-exposed seedlings. Secondary metabolite analysis showed increases in (+)-catechin, quercetin, cinnamic acid derivative, apigenin and pentagalloylglucose in birch leaves under enhanced UV-B radiation. Negative correlations between apigenin, and particularly quercetin concentrations and lipid peroxidation levels indicated an antioxidant role of secondary metabolites in birch leaves exposed to UV-B radiation.