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Patterns of environmental change in the biosphere include concurrent and sequential combinations of increasing ultraviolet (UV-B) and ozone (O(3)) at increasing carbon dioxide (CO(2)) levels; long-term changes are resulting mainly from stratospheric O(3) depletion, greater tropospheric O(3) photochemical synthesis, and increasing CO(2) emissions. Effects of selected combinations were evaluated in tomato (Lycopersicon esculentum cv. New Yorker) seedlings using sequential exposures to enhanced UV-B radiation and O(3) in differential CO(2) concentrations. Ambient (7.2 kJ m(-2 )day(-1)) or enhanced (13.1 kJ m(-2) day(-1)) UV-B fluences and ambient (380 microl l(-1)) or elevated (600 microl l(-1)) CO(2) were imposed for 19 days before exposure to 3-day simulated O(3) episodes with peak concentrations of 0.00, 0.08, 0.16 or 0.24 microl l(-1) O(3) in ambient or elevated CO(2). CO(2) enrichment increased dry mass, leaf area, specific leaf weight, chlorophyll concentration and UV-absorbing compounds per unit leaf area. Exposure to enhanced UV-B increased leaf chlorophyll and UV-absorbing compounds but decreased leaf area and root/shoot ratio. O(3) exposure generally inhibited growth and leaf photosynthesis and did not affect UV-absorbing compounds. The highest dose of O(3) eliminated the stimulating effect of CO(2) enrichment after ambient UV-B pre-exposure on leaf photosynthesis. Pre-exposure to enhanced UV-B mitigated O(3) damage to leaf photosynthesis at elevated CO(2).

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We investigated how salicylic acid (SA) enhances H2O2 and the relative significance of SA-enhanced H2O2 in Arabidopsis thaliana. SA treatments enhanced H2O2 production, lipid peroxidation, and oxidative damage to proteins, and resulted in the formation of chlorophyll and carotene isomers. SA-enhanced H2O2 levels were related to increased activities of Cu,Zn-superoxide dismutase and were independent of changes in catalase and ascorbate peroxidase activities. Prolonging SA treatments inactivated catalase and ascorbate peroxidase and resulted in phytotoxic symptoms, suggesting that inactivation of H2O2-degrading enzymes serves as an indicator of hypersensitive cell death. Treatment of leaves with H2O2 alone failed to invoke SA-mediated events. Although leaves treated with H2O2 accumulated in vivo H2O2 by 2-fold compared with leaves treated with SA, the damage to membranes and proteins was significantly less, indicating that SA can cause greater damage than H2O2. However, pretreatment of leaves with dimethylthiourea, a trap for H2O2, reduced SA-induced lipid peroxidation, indicating that SA requires H2O2 to initiate oxidative damage. The relative significance of the interaction among SA, H2O2, and H2O2-metabolizing enzymes with oxidative damage and cell death is discussed.

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Ultraviolet-B (UVB; 280-320 nm) radiation is a small but biologically significant portion of the solar spectrum reaching the earth's surface. Research interests have been fostered because UVB has been increasing in recent years due to depletion of stratospheric ozone. Ultraviolet-B that penetrates into plant tissue may damage important cellular macromolecules. Although there has been considerable research on the effects of UVB on plants, the influence of the level of photosynthetically active radiation (PAR; 400-700 nm) on effects of UVB requires further definition as a prelude to studies of UVB sensitivity and defense mechanisms. Arabidopsis thaliana wild-type ecotype Landsberg erecta (LER), which is relatively insensitive to UVB, and the relatively sensitive LER-based mutant transparent testa-5 (tt5), were grown under 100 or 250 mumol m-2 s-1 PAR and then exposed to (zero) or 7 kJ m-2 day-1 UVBBE under these PAR levels. Plants exposed to UVB had reduced dry weight and leaf area and higher levels of UV-absorbing compounds in leaf tissue. The level of PAR did influence the effects of UVB, with the higher level of PAR prior to UVB exposure reducing sensitivity of LER to UVB. In contrast to other studies, higher PAR supplied simultaneously with UVB increased rather than decreased sensitivity of both genotypes to UVB. These results demonstrate the importance of controlling and comparing PAR levels when undertaking studies of UVB sensitivity, as effects of UVB on plants are influenced by the PAR levels plants are growing under prior to and during exposure to UVB.

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Earlier studies with Arabidopsis thaliana exposed to ultraviolet B (UV-B) and ozone (O3) have indicated the differential responses of superoxide dismutase and glutathione reductase. In this study, we have investigated whether A. thaliana genotype Landsberg erecta and its flavonoid-deficient mutant transparent testa (tt5) is capable of metabolizing UV-B- and O3-induced activated oxygen species by invoking similar antioxidant enzymes. UV-B exposure preferentially enhanced guaiacol-peroxidases, ascorbate peroxidase, and peroxidases specific to coniferyl alcohol and modified the substrate affinity of ascorbate peroxidase. O3 exposure enhanced superoxide dismutase, peroxidases, glutathione reductase, and ascorbate peroxidase to a similar degree and modified the substrate affinity of both glutathione reductase and ascorbate peroxidase. Both UV-B and O3 exposure enhanced similar Cu,Zn-superoxide dismutase isoforms. New isoforms of peroxidases and ascorbate peroxidase were synthesized in tt5 plants irradiated with UV-B. UV-B radiation, in contrast to O3, enhanced the activated oxygen species by increasing membrane-localized NADPH-oxidase activity and decreasing catalase activities. These results collectively suggest that (a) UV-B exposure preferentially induces peroxidase-related enzymes, whereas O3 exposure invokes the enzymes of superoxide dismutase/ascorbate-glutathione cycle, and (b) in contrast to O3, UV-B exposure generated activated oxygen species by increasing NADPH-oxidase activity.

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O3-induced changes in growth, oxidative damage to protein, and specific activities of certain antioxidant enzymes were investigated in wheat plants (Triticum aestivum L. cv Roblin) grown under ambient or high CO2. High CO2 enhanced shoot biomass of wheat plants, whereas O3 exposure decreased shoot biomass. The shoot biomass was relatively unaffected in plants grown under a combination of high CO2 and O3. O3 exposure under ambient CO2 decreased photosynthetic pigments, soluble proteins, and ribulose-1,5-bisphosphate carboxylase/oxygenase protein and enhanced oxidative damage to proteins, but these effects were not observed in plants exposed to O3 under high CO2. O3 exposure initially enhanced the specific activities of superoxide dismutase, peroxidase, glutathione reductase, and ascorbate peroxidase irrespective of growth in ambient or high CO2. However, the specific activities decreased in plants with prolonged exposure to O3 under ambient CO2 but not in plants exposed to O3 under high CO2. Native gels revealed preferential changes in the isoform composition of superoxide dismutase, peroxidases, and ascorbate peroxidase of plants grown under a combination of high CO2 and O3. Furthermore, growth under high CO2 and O3 led to the synthesis of one new isoform of glutathione reductase. This could explain why plants grown under a combination of high CO2 and O3 are capable of resisting O3-induced damage to growth and proteins compared to plants exposed to O3 under ambient CO2.

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The impact of sequential exposure to ozone (O3) and UVB (290-320 nm) was studied using two genotypes of Arabidopsis thaliana differing in UVB sensitivity. The negative impact of UVB on dry matter production and photosynthetic pigments was absent in the ecotype Landsberg erecta (LER), while the negative impact of UVB was more pronounced when LER plants pre-exposed to O3 were irradiated with UVB. However, the growth of tt5 plants (a mutant virtually incapable of synthesizing flavonoids) was significantly affected by the UVB exposures, while the impact of UVB was significantly counteracted when tt5 plants pre-exposed to O3 were irradiated with UVB. These results suggest that pre-exposure to O3 decreased sensitivity of tt5 but increased sensitivity of LER to UVB. Concentrations of UV-absorptive compounds were almost the same in plants exposed to UVB alone or sequentially to O3 and UVB. Exposures of LER and tt5 to UVB enhanced both ascorbic acid and glutathione as well as their redox state compared to control plants. Pre-exposure to O3 enhanced the total ascorbic acid and glutathione as well as the redox state of ascorbate and glutathione in tt5 but decreased the redox state in LER. Irradiation of plants pre-exposed to O3 with UVB enhanced the redox state of ascorbate and glutathione slightly in tt5 but decreased it further in LER. The high redox state of ascorbate and glutathione in tt5 pre-exposed to O3 would have protected plants from UVB and decreased their sensitivity to UVB in spite of their inability to synthesize flavonoids.(ABSTRACT TRUNCATED AT 250 WORDS)

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Phaseolus vulgaris cv. Kinghorn Wax seedlings grown in darkness at 25 degrees C for 7 days with half strength Hoagland's nutrient solution containing no nitrogen, were transferred to lit continuous stirred tank reactors (CSTRs) in atmospheres containing 0 or 0.3 ppm NO(2) and irrigated with a nutrient solution containing 0 or 5 mm nitrate as sole nitrogen source and allowed to grow for a period of up to 5 days in a 14 h photoperiod. Exposure to NO(2) increased total Kjeldahl nitrogen in the leaves. Further, the exposure to NO(2) increased chlorophyll content from day 3 onwards and inhibited the leaf dry weight substantially on days 4 and 5. The primary leaves of the seedlings exposed to 0.3 ppm NO(2) and supplied with nitrate accumulated some nitrite after 5 days of exposure. Some of the seedlings were returned from CSTRs to growth chambers and allowed to grow for a further period of 5 days in a 14 h photoperiod without NO(2). The growth which developed after the NO(2) exposure growth period, as measured by fresh and dry weights of the leaves, was significantly less in NO(2)-exposed plants than in nitrate-grown plants. The experiments demonstrate that the leaves of greening seedlings are able to assimilate NO(2) and that a reduction in leaf dry weight by prolonged NO(2) exposure in the presence of nutrient nitrate can be associated with nitrite accumulation, and that NO(2) has a carry-over effect beyond the duration of NO(2) exposure. It is apparent that NO(2) induces some durable biochemical or cytological aberration in the presence of nutrient nitrate, which adversely affects subsequent leaf growth.

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The response to ozone (O(3)) of greenness, in terms of estimated total chlorophyll concentration (Chl), of leaves at three plant canopy levels was studied in tomato (Lycopersicon esculentum Mill.) over a 10-day period following O(3) exposure. Plants of the cultivars 'New Yorker' and 'Tiny Tim' were grown at 25/15 degrees or 30/15 degrees day/night temperatures in growth chambers and exposed to 0.00, 0.08, 0.16 or 0.24 microl litre(-1) O(3) for 7 h day(-1) for four consecutive days in controlled environment exposure chambers. Measurement of Chl in the top, middle and bottom canopy leaves with a calibrated SPAD-501 leaf greenness meter indicated that the growth temperatures tested did not significantly influence the response of Chl to O(3). Ozone-induced loss of Chl was widespread in the entire foliage canopy, including foliage which did not demonstrate visible injury. In both cultvars the Chl in leaves at all three canopy levels declined as a function of increasing O(3) concentration when measured 2, 4, 6, 8 and 10 days after the exposure period. However, the slopes for leaves in the top and middle canopies decreased with increasing time after exposure. An analysis of this apparent Chl recovery indicated that leaves in the top and middle canopies exposed to 0.16 and 0.24 microl litre(-1) increased in greenness at a rapid rate after the marked initial decline associated with O(3) treatment. The apparent recovery of the top canopy may have reflected the growth of new leaves and their inclusion in the measurements, but this was not the case for the middle canopy for which the same leaves were measured throughout the post-exposure period. Bottom canopy leaves did not demonstrate significant recovery of Chl.

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Tomato (Lycopersicon esculentum Mill.) 'New Yorker' plants were exposed to O(3) to compare leaf diffusive conductance (LDC) before exposure to O(3) with O(3) sorption rates and visible injury ratings. Two plant development stages and four or five leaf growth stages were examined. The LDC varied among leaf growth stages and between plant development stages and leaf surfaces; there was no continuity in the LDC pattern. Sorption rates differed among some leaf growth stages, and between plant development stages in expanding leaves (growth stage 1). For both development stages high sorption rates occurred in fully mature leaves; otherwise little similarity between corresponding leaf growth stages was evident. Total O(3) flux to the leaf was not well predicted by the LDC for water vapour; nor was visible injury well related to total flux. Differential mesophyll processes and leaf surface sorption capabilities may have accounted for some of the inconsistencies observed.

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In the ambient environment, concentrations of air pollutants vary on a diurnal cycle, resulting in various patterns of concurrent and sequential exposures of plants. The response of tomato plants to sequential and concurrent NO2 and O3 exposures was determined using pollutant levels equal to the maximum acceptable levels recommended by the National Ambient Air Quality Objectives of Environment Canada for a 1 h average. The concurrent treatment, 1 h of NO2 + O3, was compared to 1 h of NO20, O3 or control in plants at the 4 to 6 or the 9 to 11 leaf stage. At the 4 to 6 leaf stage, leaf and stem fresh weights were significantly reduced by the NO2 + O3 treatment relative to control, whereas these growth parameters were not reduced relative to control by the single pollutants indicating a coalitive response. Leaf area was significantly smaller as a result of the NO2 + O3 treatment relative to the NO2 treatment. A main effect of O3 was observed on leaf dry weight. The sequential treatments were: NO2 followed by O3 (NO2-O3); O3 followed by NO2 (O3-NO2); NO2 at night followed by O3 during the daytime (NO2(N)-O3(D)). Each gas exposure was 1 h; only plants at the 4 to 6 leaf stage were treated. Only the O3-NO2 treatment significantly reduced leaf area, leaf fresh weight and stem fresh and dry weights relative to control plants. Inconsistencies among treatments occurring at different time periods of the day suggest that time period of exposure should reflect ambient time periods. The coalitive action, and the sequential treatment response, of these pollutants indicated that criteria based on single pollutants may not be adequate to establish air quality objectives when these pollutants occur together.

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The influence of nutrient nitrate level (0-20 millimolar) on the effects of NO(2) (0-0.5 parts per million) on nodulation and in vivo acetylene reduction activity of the roots and on growth and nitrate and Kjeldahl N concentration in shoots was studied in bean (Phaseolus vulgaris L. cv Kinghorn Wax) plants. Exposing 8-day old seedlings for 6 hours each day, for 15 days, to 0.02 to 0.5 parts per million NO(2) decreased total nodule weight at 0 and 1 millimolar nitrate, and nitrogenase (acetylene reduction) activity at all concentrations of nitrate. The pollutant had little effect on root fresh or dry weights. Shoot growth was inhibited by NO(2). The NO(2) exposure increased nitrate concentration in roots only at 20 millimolar nutrient nitrate. Exposure to NO(2) markedly increased Kjeldahl N concentration in roots but generally decreased that in shoots. The experiments demonstrated that nutrient N level and NO(2) concentration act jointly in affecting nodulation and N fixing capability, plant growth and composition, and root/shoot relationships of bean plants.

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The influence of nutrient nitrate level (0-20 millimolar) on the effects of NO(2) (0-0.5 parts per million) on growth, K, photosynthetic pigment, N contents, and the activities of enzymes of N assimilation was studied in bean (Phaseolus vulgaris L. cv Kinghorn Wax) leaves. Exposing 7-day old bean seedlings for 5 days continuously to 0.02 to 0.5 parts per million NO(2) increased plant height, fresh weight, chlorophyll, carotenoid, organic N and nitrate contents, and nitrate reductase and glutamate synthase activities in the leaves of seedlings supplied with no external N. At 20 millimolar nitrate, most of the parameters examined were inhibited except for organic N and nitrate contents and glutamate synthase activity which increased in most cases. Generally, with an increase in NO(2) concentration, the stimulatory effect declined and/or the inhibitory effect increased. A 3-hour exposure of 12-day-old bean seedlings to 0.1 to 2.0 parts per million NO(2) increased nitrate content and nitrate reductase activity at each nutrient nitrate level except for a slight inhibition of enzyme activity during exposure to 2.0 parts per million NO(2) at 20 millimolar nitrate. The experiments demonstrated that the effect of NO(2) is strongly influenced by nutrient N level and that NO(2) is assimilated into organic nitrogenous compounds to serve as a source of N, only to a limited extent.

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A second order rotatable design was used to obtain polynomial equations describing the effects of combinations of sulfur dioxide (SO(2)) and ozone (O(3)) on foliar injury and plant growth. The response surfaces derived from these equations were displayed as contour or isometric (3-dimensional) plots. The contour plots aided in the interpretation of the pollutant interactions and were judged easier to use than the isometric plots. Plants of ;Grand Rapids' lettuce (Lactuca sativa L.), ;Cherry Belle' radish (Raphanus sativus L.), and ;Alsweet' pea (Pisum sativum L.) were grown in a controlled environment chamber and exposed to seven combinations of SO(2) and O(3). Injury was evaluated based on visible chlorosis and necrosis and growth was evaluated as leaf area and dry weight. Covariate measurements were used to increase precision. Radish and pea had greater injury, in general, that did lettuce; all three species were sensitive to O(3), and pea was most sensitive and radish least sensitive to SO(2). Leaf injury responses were relatively more affected by the pollutants than were plant growth responses in radish and pea but not in lettuce. In radish, hypocotyl growth was more sensitive to the pollutants than was leaf growth.

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Experiments were conducted in growth chambers to examine the effect of a mild water stress (-200 kilopascals polyethylene glycol) on frond elongation and water status of the ostrich fern (Matteuccia struthiopteris [L.] Todaro). Measurements were taken for two days, starting one day after the application of polyethylene glycol. Total water potential in control (well-watered) plants was always significantly higher in immature fronds than in mature fronds. The osmotic potential in mature fronds was always significantly lower (about 800 kilopascals) than in immature fronds in both control and stressed plants. In immature fronds, the stress decreased elongation and total water and pressure potentials, while in mature fronds it increased total water and pressure potentials. The decreases in total and pressure potentials in immature fronds were approximately equal to the increases in mature fronds.

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Ten selections of citronella (Cymbopogon nardus [L.] Rendle) were grown at 32/27, 27/21, or 15/10 C day/night temperatures, and plants from three populations of lemongrass (Cymbopogon citratus [D.C.] Stapf from Japan or Sri Lanka and Cymbopogon flexuosus [D.C.] Stapf from India) were grown at 8- or 15-hour photoperiods. Net photosynthetic rates of mature leaves were measured in a controlled environment at 25 C and 260 microeinsteins per meter(2) per second. Rates declined with increasing leaf age, and from the tip to the base of the leaf blade. Rates for citronella leaves grown at 15/10 C were extremely low for all selections. Highest rates of net photosynthesis were recorded for four selections grown at 27/21 C and for two selections grown at 32/27 C. Lemongrass grown at 8-hour photoperiod had higher photosynthetic rates than that grown at 15-hour photoperiod.

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Carbon dioxide compensation values of mature leaves from 10 selections of citronella (Cymbopogon nardus [L.] Rendle) grown at 32/27 or 27/21 C day/night temperatures and three strains of lemongrass (Cymbopogon citratus [D.C.] Stapf. and Cymbopogon flexuosus [D.C.] Stapf.) grown at 8- or 15-hour photoperiods were measured in a controlled environment at 25 C. All leaves had low compensation values but citronella varied from 1.3 to 9.7 mul/liter and lemongrass from 0.7 to 3.5 mul/liter. Lower growing temperature generally resulted in lower compensation values for citronella but there was no consistent photoperiod effect on lemongrass.