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Injury caused by low O3 concentrations (0, 0.05, 0.075, 0.1 μl 1-1) was analyzed in the epidermis and mesophyll of fully developed birch leaves by gas exchange experiments and low-temperature SEM: (I) after leaf formation in O3-free and ozonated air, and (II) after transferring control plants into ozonated air. In control leaves, autumnal senescence also was studied in O3-free air (III). As O3 concentration increased, leaves of (I) stayed reduced in size, but showed increased specific weight and stomatal density. The declining photosynthetic capacity, quantum yield and carboxylation efficiency lowered the light saturation of CO2 uptake and the water-use efficiency (WUE). Carbon gain was less limited by the reduced stomatal conductance than by the declining ability of CO2 fixation in the mesophyll. The changes in gas exchange were related to the O3 dose and were mediated by narrowed stomatal pores (overriding the increase in stomatal density) and by progressive collapse of mesophyll cells. The air space in the mesophyll increased, preceded by exudate formation on cell walls. Ozonated leaves, which had developed in O3-free air (II), displayed a similar but more rapid decline than the leaves from (I). In senescent leaves (III), CO2 uptake showed a similar decrease as in leaves with O3 injury but no changes in mesophyll structure and WUE. The nitrogen concentration declined only in senescent leaves in parallel with the rate of CO2 uptake. A thorough understanding of O3 injury and natural senescence requires combined structural and functional analyses of leaves.

Impairment of gas exchange and structure in birch leaves (Betula pendula) caused by low ozone concentrations