This paper presents a mathematical model which enables the semi-quantification of ozone (O3) detoxification, based upon the direct reaction of the pollutant with ascorbate (ASC) located in the aqueous matrix associated with the cell wall (i.e. the apoplast). The model describes the uptake of ozone into the leaf and its direct reaction with ASC, taking into consideration the regeneration of dehydroascorbic acid in the cytosol, the rate of replenishment of cell wall ASC and the distribution of ASC between sub-cellular compartments based upon the permeability of biomembranes to the neutral species, ascorbic acid and the pH of various sub-cellular compartments. The importance of various physico-chemical characteristics (e.g. stomatal conductance, mesophyll cell wall thickness and tortuosity, chloroplast volume, apoplast pH, ASC:O3 reaction stoichiometry) in mediating the flux of ozone to the plasmalemma is analysed. Model simulations, supported by experimental observations, suggest that the ASC concentration in the leaf apoplast is high enough to scavenge a significant proportion of the O3 taken up into the leaf interior, under environmentally relevant conditions. However, there is considerable variation between taxa in the potential degree of protection afforded by apoplastic ASC, emphasizing the need for an improved understanding of the reaction chemistry of O3 in the cell wall.

Individual tillers of S23 ryegrass, in three paddocks set-stocked with sheep at widely differing grazing pressures, were marked and studied from 11 July to 19 August, 1966, and observations were made on the frequency and severity of grazing. The uprooting of tillers, and the destruction of others, were major sources of loss of records. It was found that an acceptable measure of the frequency of defoliation could be made by recording three times a week. In a subsidiary experiment it was shown that repeated handling did not affect the DM yield, the number of tillers or the length of green leaf (G.L.L.) of single plants. Despite the extreme differences in grazing pressure, there were no significant differences between paddocks in the daily increase in the G.L.L. of tillers which were not grazed during a short period of intensive recording. In all three paddocks the sheep tended to graze the tillers with the greatest G.L.L. at any one time, £ind there was evidence of patchy grazing only at the lowest grazing pressure. An increase in grazing pressure resulted in increases in both the frequency and severity of defoliation. Comparisons were made between the product of the frequency X severity of defoliation of individual tillers, and the stocking rate, grazing pressure and herbage consumption on the three paddocks.

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Two cultivars of spring wheat {Triticum aestivum L. cvs. Alexandria and Hanno) and three cultivars of winter wheat (cvs. Riband, Mercia and Haven) were grown at two concentrations of CO2 [ambient (355 pmol mol"^) and elevated (708 |.imol mol"^)] under two O3 regimes [clean air (< 5 nmol mol"' O3) and polluted air (15 nmol mol"' O3 at night rising to a midday maximum of 75 nmol mol""^)l in a phytotron at the University of Newcast!e-upon-Tyne. Between the two-leaf stage and anthesis, measurements of leaf gas-exchange, non-structural carbohydrate content, visible O3 damage, growth, dry matter partitioning, yield components and root development were made in order to examine responses to elevated CO2 and/or O3.Growth at elevated CO2 resulted in a sustained increase in the rate of CO2 assimilation, but after roughly 6 weeks' exposure there was evidence of a slight decline in the photosynthetic rate (c.-15%) measured under growth conditions which was most pronounced in the winter cultivars. Enhanced rates of CO2 assimilation were accompanied by a decrease in stomatal conductance which improved the instantaneous water use efficiency of individual leaves. CO2 enrichment stimulated shoot and root growth to an equivalent extent, and increased tillering and yield components, however, non-structural carbohydrates still accumulated in source leaves. In contrast, long-term exposure to O3 resulted in a decreased CO2 assimilation rate (c. -13%), partial stomatal closure, and the accumulation of fructan and starch in leaves in the light. These effects were manifested in decreased rates of shoot and root growth, with root growth more severely affected than shoot growth.In the combined treatment growth of 03-treated plants was enhanced by elevated CO2, but there was little evidence that CO2 enrichment afforded additional protection against O3 damage. The reduction in growth induced by O3 at elevated CO2 was similar to that induced by O3 at ambient CO2 despite additive effects of the individual gases on stomatal conductance that would be expected to reduce the O3 flux by 20%, and also CO2-induced increases in the provision of substrates for detoxification and repair processes. These observations suggest that CO2 enrichment may render plants more susceptible to O3 damage at the cellular level. Possible mechanisms are discussed.

The aim of this work was to examine the correspondence between apoplastic\symplastic antioxidant status and previously reported plant age-related shifts in the ozone (O $ ) resistance of Plantago major L. Seed-grown plants were fumigated in duplicate controlled environment chambers with charcoal\Purafil2-filtered air (CFA) or CFA plus 70 nmol mol −" O $ for 7 h d −" over a 42 d period. Measurements of stomatal conductance and antioxidants were made after 14, 28 and 42 d fumigation, on leaves at an equivalent stage of development (youngest fully expanded leaf, measured c. 9 d after emergence). Ozone exposure resulted in a similar decline in stomatal conductance across plant ages, indicating that increases in O $ resistance with plant age were mediated through changes in the tolerance of leaf tissue rather than enhanced pollutant exclusion. Leaf apoplastic washing fluid was found to contain ' unspecific ' peroxidase, ascorbate peroxidase, superoxide dismutase and ascorbate, but not glutathione and the enzymes required to facilitate the regeneration of ascorbate from its oxidized forms. A weak induction in the activity of certain symplastic antioxidants was found after 14 d O $ fumigation, despite a lack of visible symptoms of injury, but shifts in symplastic antioxidant enzyme activity were not consistent with previously observed increases in resistance to O $ with plant age. By contrast, changes in ' unspecific ' peroxidase activity and in the small pool of ascorbate in the leaf apoplast were found to accompany age-related shifts in O $ resistance. It is concluded that constituents of the leaf apoplast may constitute a potentially important front line defence against O $ .