The interactive effects of ozone and light on leaf structure, carbon dioxide uptake and short-term carbon allocation of sugar maple (Acer saccharum Marsh.) seedlings were examined using gas exchange measurements and 14 Cmacroautoradiographic techniques. Two-year-old sugar maple seedlings were fumigated from budbreak for 5 months with ambient or 3 × ambient ozone in open-top chambers, receiving either 35% (high light) or 15% (low light) of full sunlight. Ozone accelerated leaf senescence, and reduced net photosynthesis, 14 CO 2 uptake and stomatal conductance, with the effects being most pronounced under low light. The proportion of intercellular space increased in leaves of seedlings grown under elevated ozone and low light, possibly enhancing the susceptibility of mesophyll cells to ozone by increasing the cumulative dose per mesophyll cell. Indeed, damage to spongy mesophyll cells in the elevated ozone × low light treatment was especially frequent. 14 C macroautoradioraphy revealed heterogeneous uptake of 14 CO 2 in well defined areole regions, suggesting patchy stomatal behaviour in all treatments. However, in seedlings grown under elevated ozone and low light, the highest 14 CO 2 uptake occurred along larger veins, while interveinal regions exhibited little or no uptake. Although visible symptoms of ozone injury were not apparent in these seedlings, the cellular damage, reduced photosynthetic rates and reduced whole-leaf chlorophyll levels corroborate the visual scaling of whole-plant senescence, suggesting that the ozone × low light treatment accelerated senescence or senescence-like injury in sugar maple.
Key-words:Acer saccharum Marsh; leaf morphology; ozone; photosynthesis; shade adaptation; stomatal patchiness; sugar maple.
INTRODUCTIONSugar maple (Acer saccharum Marsh.) is a slow-growing, shade-tolerant species that is widely distributed in eastern North America where elevated ozone concentrations occur, and forests may currently be undergoing widespread decline. Because of its high phytotoxicity, ozone probably has the most negative impact of any air pollutant on tree vigour and growth (Reich 1987). Once in the leaf, ozone reacts with water to form oxygen free radicals, H 2 O 2 and hydroxyl ions (Heath 1980;Mehlhorn, Tabner & Wellburn 1990), oxidizes proteins and membrane phospholipids, increases membrane permeability (Mehlhorn et al. 1986) and directly inhibits photosynthesis (Heath 1994). Stromal reactions of photosynthesis are particularly sensitive to ozone, with ozone affecting the quantity and activity of rubisco (Pell, Eckardt & Enyedi 1992). Plant responses to ozone are proportional to ozone uptake, which is a function of atmospheric concentration and stomatal conductance (Laisk, Kull & Moldau 1989). Plants can reduce ozone uptake and minimize internal damage by closing stomata or reducing the size of the stomatal opening, resulting in a decline in stomatal conductance. However, this response may vary within and between species, with stage of leaf development, and be dependent upon atm...