Birch cuttings (Betula pendula Roth) were grown in a sand-culture system with two concentrations (0·05, HF and 0·005%, LF) of fertilizer containing macronutrients and micronutrients, and were exposed to 90/40 nl l−1 O3 (day/night) and <3 nl l−1 O3 (control) for one growing season in the field fumigation chambers at Birmensdorf (Switzerland). Leaves of different ages were analysed for gas exchange, contents of chlorophyll, protein, and for metabolites as well as enzyme activities of carbohydrate metabolism.

Ozone reduced net photosynthesis and chlorophyll contents in mature leaves of both fertilization treatments, whereas that of protein was only reduced in high-fertilized plants (HF). However, net photosynthesis, chlorophyll, and protein increased in young leaves of low-fertilized plants (LF). The effects of ozone on enzyme activities of carbohydrate metabolism were most pronounced in leaves of LF plants. Specific activities of the sucrose-cleaving enzymes, sucrose synthase and alkaline invertase, were induced, whereas acid invertase was unchanged. Extractable activity of sucrose phosphate synthase, which is a key enzyme of sucrose synthesis, was reduced. Levels of fructose 2,6-bisphosphate, an inhibitor of sucrose synthesis, were increased in leaves of O3/LF plants, but reduced in O3/HF plants. In addition, activities of enzymes involved in starch metabolism, ADP-glucose pyrophosphorylase and starch phosphorylase, were lowered in ozone-treated samples and the ratio of ATP[ratio ]ADP was increased.

It is concluded that chronic ozone exposure leads to an inhibition of sucrose synthesis and favours sucrose degradation. This effect is modulated by the nutrient status of the plants, indicating higher O3 tolerance in HF plants. Furthermore, as the metabolic responses in the ozone-treated samples resemble very closely those observed under end-product inhibition of photosynthesis, we assume that the O3 effect is mainly due to reduced photosynthate export.

Cloned cuttings of Betula pendula Roth were grown in field fumigation chambers at Birmensdorf throughout one growing season in filtered air with either <3 (control) or 90/40 nl l−1 O3 (day/night; ozone generated from pure oxygen). Each ozone regime was split into high and low soil nutrient regimes by watering plants with either a 0·05% or a 0·005% solution of a fertilizer which contained macronutrients and micronutrients.

Fertilization had a strong effect on plant growth, enzyme activities and the expression of ozone-induced effects at the biochemical level. The activities of PEPC and Rubisco were enhanced about threefold in the plants with high fertilization (HF). Significant effects of ozone were in most cases found only in the older leaves of the plants with low fertilization (LF). There, sucrose, glucose and fructose levels were enhanced. In both fertilization treatments, the number of starch granules along the minor veins was increased. These ozone effects point to a decreased or inhibited phloem loading. The increased PEPC activity and the enhanced malate levels in the ozone-exposed plants might be the result of a redirection of carbon flow from sucrose synthesis and translocation towards anapleurotic processes, which can feed detoxification and repair of ozone injury as indicated by enhanced respiration. These findings agree well with the observed effects of ozone in lowering the root[ratio ]shoot biomass ratio. Although there was a marked reduction in the O3/LF plants, O3/HF plants showed no significant response. Inositol was decreased under ozone exposure in both fertilizer treatments, contrasting with the pattern for carbohydrates.

These results demonstrate the role of fertilization as an important modifier of ozone-induced effects at the plant biochemical level. Well fertilized plants appear to cope better with the impact of ozone on metabolism.