Ozone exposure over two growing seasons alters root-to-shoot ratio and chemical composition of birch (Betula pendula Roth)

Physiological and chemical responses of 17 birch (Betula pendula Roth) clones to 1.5-1.7 x ambient ozone were studied in an open-field experiment over two growing seasons. The saplings were studied for growth, foliar visible injuries, net photosynthesis, stomatal conductance, and chlorophyll, carotenoid, Rubisco, total soluble protein, macronutrient and phenolic concentrations in leaves. Elevated ozone resulted in growth enhancement, changes in shoot-to-root (s/r) ratio, visible foliar injuries, reduced stomatal conductance, lower late-season net photosynthesis, foliar nutrient imbalance, changes in phenolic composition, and reductions in pigment, Rubisco and soluble protein contents indicating accelerated leaf senescence. Majority of clones responded to ozone by changing C allocation towards roots, by stomatal closure (reduced ozone uptake), and by investment in low-cost foliar antioxidants to avoid and tolerate ozone stress. A third of clones, showing increased s/r ratio, relied on inducible efficient high-cost antioxidants, and enhanced leaf production to compensate ozone-caused decline in leaf-level net photosynthesis. However, the best ozone tolerance was found in two s/r ratio-unaffected clones showing a high constitutive amount of total phenolics, investment in low-cost antioxidants and N distribution to leaves, and lower stomatal conductance under ozone stress. The results highlight the importance of phenolic compounds in ozone defence mechanisms in the birch population. Depending on the genotype, ozone detoxification was improved by an increase in either efficient high-cost or less efficient low-cost antioxidative phenolics, with close connections to whole-plant physiology.