Effects of ozone impact on the gas exchange and chlorophyll fluorescence of juvenile birch stems (Betula pendula Roth.)

Effects of ozone impact on gas exchange and chlorophyll fluorescence of juvenile birch (Betula pendula) stems and leaves were investigated. Significant differences in the response of leaves and stems to ozone were found. In leaves, O3 exposure led to a significant decline in photosynthetic rates, whereas stems revealed an increased dark respiration and a concomitant increase in corticular photosynthesis. In contrast to birch leaves, corticular photosynthesis appeared to support the carbon balance of stems or even of the whole-tree under O3 stress. The differences in the ozone-response between leaves and stems were found to be related to ozone uptake rates, and thus to inherent differences in leaf and stem O3 conductance.     

Impact of ozone on the growth of birch (Betula pendula) saplings

Saplings of one half-sib family of birch, Betula pendula, were exposed to three levels of ozone in open-top chambers (OTCs) during two growing seasons 1997-1998. The ozone treatments were non-filtered air (NF, accumulated daylight AOT40 over the two growing seasons of 3.0 l l-1 h), non-filtered air with extra ozone (NF+, accumulated daylight AOT40 of 27.3 l l-1 h) and non-filtered air with additional extra ozone (NF++, accumulated daylight AOT40 of 120 l l-1 h). The birch saplings, including the roots, were harvested after the first and second growing seasons. After the first growing season, the NF++ treatment reduced the total wood biomass by 22%, relative to the NF treatment. There was no further reduction of the total wood biomass in the NF++ treatment after the second growing season. The root biomass was reduced by 30% after the first growing season. The shoot/root ratio, as well as the proportional biomass of leaves, were increased by ozone during both years. The ozone impact on the relative growth rate was estimated to -2% per 10 l l-1 h daylight AOT40 per growing season.     

Responses of two birch (Betula pendula Roth) clones to different ozone profiles with similar AOT40 exposure

Saplings of two clones of European white birch (Betula pendula Roth) were exposed to three different ozone profiles resulting in same AOT40 value of 13–14 ppm h in a chamber experiment. The sensitive clone 5 and the more tolerant clone 2 were growing (1) under filtered air (=control), or (2) were exposed to 70 ppb ozone for 24 h d⁻¹ (=profile 1), (3) to 100 ppb ozone for 12 h d⁻¹ at 8:00–20:00 (=profile 2), or (4) to 200 ppb ozone for 4.5 h d⁻¹ at 9:30–14:00 (=profile 3) for 20 d. The saplings were determined for growth, visible leaf injuries, stomatal conductance, and concentrations of Rubisco, chlorophyll and carotenoids. Growth responses and induction of visible foliar injuries under different ozone profiles were variable, resulting in 4–17% lower dry mass of shoot, 16–46% reduction in stem height increment and 11–43% increase in visible injuries in clone 5, which was accompanied by higher leaf turnover rate under profile 3 indicating compensation growth. In clone 2, ozone-induced responses ranged from slight stimulation in stem height growth to 13% decrease in dry mass of shoot and 2–16% increase in visible injuries. Daytime stomatal conductance rates were lowered by 14–54% in clone 5 and 9–74% in clone 2, depending on profile. The additional power-weighted analyses revealed that high peak concentrations and exposure shape were important for induction of visible injuries in both clones and reduction in stomatal conductance in clone 5, whereas growth reductions were rather related to total cumulative exposure. The results indicate that profile of ozone exposure, night-time stomatal conductance (24 h flux), and recovery time for defence and compensations reactions should not be ignored in plant response and ozone flux modelling.     

Variation in ozone sensitivity among clones of Betula pendula and Betula pubescens

Forty clones of Betula pendula and 6 clones of Betula pubescens, originating from southern and central Finland, were ranked in order of ozone sensitivity according to visible injuries, growth and leaf senescense under low ozone exposure. The plants were fumigated in natural climatic conditions using an open-air exposure system during two growing seasons. Control plants were grown under ambient air, and the elevated-ozone exposures were 1.6x the ambient in 1994 and 1.7x the ambient in 1995. The differences in ozone sensitivity among clones were large. Ozone tolerance was related to thicker leaves and higher stomatal density as compared to sensitive clones. Ultrastructural ozone-induced symptoms were found in chloroplasts of sensitive clones. Increased number of visibly injured leaves on fumigated plants was correlated with reduced leaf formation, foliage area, shoot dry wt and number of stomata, and increased yellowing of leaves. The results suggest that a considerable proportion of birch trees, showing high sensitivity to ozone, are at risk if ambient ozone exposures increase.     

Acquired changes in stomatal characteristics in response to ozone during plant growth and leaf development of bush beans (Phaseolus vulgaris L.) indicate phenotypic plasticity

Bush bean (Phaseolus vulgaris L.) lines 'S156' (O3-sensitive)/'R123' (O3-tolerant) and cultivars 'BBL 290' (O3-sensitive)/'BBL 274' (O3-tolerant) were used to study the effects of O3 on stomatal conductance (gs), density, and aperture size on leaf and pod surfaces with the objective of establishing links between the degree of plant sensitivity to O3 and plasticity of stomatal properties in response to O3. Studies in open-top chambers (OTCs) and in continuously stirred tank reactors (CSTRs) established a clear relationship between plant developmental stages, degrees of O3 sensitivity and gs: while 'S156' had higher gs rates than 'R123' earlier in development, similar differences between 'BBL 290' and 'BBL 274' were observed at later stages. Gs rates on the abaxial leaf surfaces of 'S156' and 'BBL 290', accompanied by low leaf temperatures, were significantly higher than their O3-tolerant counterparts. Exposure to O3 in CSTRs had greater and more consistent impacts on both stomatal densities and aperture sizes of O3-sensitive cultivars. Stomatal densities were highest on the abaxial leaf surfaces of 'S156' and 'BBL 290' at higher O3 concentrations (60 ppb), but the largest aperture sizes were recorded on the adaxial leaf surfaces at moderate O3 concentrations (30 ppb). Exposure to O3 eliminated aperture size differences on the adaxial leaf surfaces between sensitive and tolerant cultivars. Regardless of sensitivity to O3 and treatment regimes, the smallest aperture sizes and highest stomatal densities were found on the abaxial leaf surface. Our studies showed that O3 has the potential to affect stomatal plasticity and confirmed the presence of different control mechanisms for stomatal development on each leaf surface. This appeared to be more evident in O3-sensitive cultivars.     

Visible injury, crown condition, and growth responses of selected Italian forests in relation to ozone exposure

The impact of ozone on forest ecosystems in Italy is monitored within the CONECOFOR programme. Ozone levels are measured in 30 plots using passive samplers. Response parameters used are: crown condition (transparency), BAI (basal area increment), and visible symptoms on spontaneous vegetation. Levels of AOT40 are above the concentration-based critical level of 5 ppmh in all sites, but the evidence of impact on forest vegetation remains limited. Ozone is a predictor of crown transparency residuals in beech sites over two consecutive years, but the variance explained amounts to less than 10%. The relation between BAI reduction and ozone is even less certain. Transparency and BAI are more readily explainable in terms of ecological conditions of the site and climate fluctuations. The interpretation of visible symptoms is doubtful, and is conditioned by the prevailing ecological factors in the areas.     

Foliar injury, leaf gas exchange and biomass responses of black cherry (Prunus serotina Ehrh.) half-sibling families to ozone exposure

Open pollinated families of black cherry seedlings were studied to determine genotypic differences in foliar ozone injury and leaf gas exchange in 1994 and growth response following three growing seasons. An O(3)-sensitive half-sibling family (R-12) and an O(3)-tolerant half-sibling family (MO-7) planted in natural soil were studied along with generic nursery stock (NS) seedlings. Ozone exposure treatments were provided through open top chambers and consisted of 50, 75, and 97% of ambient ozone, and open plots from May 9 to August 26, 1994. Ambient ozone concentrations reached an hourly peak of 88 ppb with 7-hour averages ranging from 39 to 46 ppb. Seedlings in the 50 and 75% of ambient chambers were never exposed to greater than 80 ppb O(3). Visible foliar ozone injury (stipple) was significantly higher for R-12 seedlings than MO-7 seedlings and increased with increasing ozone exposures. For the chamber treatments averaged over all families, there was no significant difference in stomatal conductance and net photosynthetic rates, but there was a significant decrease in root biomass, and a significant decrease in root/shoot ratio between the 50 and 97% of ambient chambers. Stomatal conductance and net photosynthetic rates were significantly different between families with R-12 seedlings generally greater than MO-7 seedlings. The R-12 seedlings had a 7.5 mmol m(-2) increase in ozone uptake compared to MO-7, and at the same cumulative O(3) exposure R-12 exhibited 40.9% stippled leaf area, whereas MO-7 had 9.2% stippled leaf area. Significant differences were observed in stem volume growth and total final biomass between the open-top chambers and open plots. Although R-12 had the most severe foliar ozone injury, this family had significantly greater stem volume growth and total final biomass than MO-7 and NS seedlings. Root:shoot ratio was not significantly different between MO-7 and R-12 seedlings.     

Effect of birch (Betula spp.) and associated rhizoidal bacteria on the degradation of soil polyaromatic hydrocarbons, PAH-induced changes in birch proteome and bacterial community

Two birch clones originating from metal-contaminated sites were exposed for 3 months to soils (sand-peat ratio 1:1 or 4:1) spiked with a mixture of polyaromatic hydrocarbons (PAHs; anthracene, fluoranthene, phenanthrene, pyrene). PAH degradation differed between the two birch clones and also by the soil type. The statistically most significant elimination (p ≤ 0.01), i.e. 88% of total PAHs, was observed in the more sandy soil planted with birch, the clearest positive effect being found with Betula pubescens clone on phenanthrene. PAHs and soil composition had rather small effects on birch protein complement. Three proteins with clonal differences were identified: ferritin-like protein, auxin-induced protein and peroxidase. Differences in planted and non-planted soils were detected in bacterial communities by 16S rRNA T-RFLP, and the overall bacterial community structures were diverse. Even though both represent complex systems, trees and rhizoidal microbes in combination can provide interesting possibilities for bioremediation of PAH-polluted soils.     

Inheritance of ambient ozone insensitivity in common bean (Phaseolus vulgaris L.)

The inheritance of ozone (O(3)) insensitivity in common bean (Phaseolus vulgaris L.) was evaluated using F(2) and F(3) populations under ambient conditions. This study was conducted over two growing seasons (1987, 1988) at Virginia State University, Randolph Research Farm, Petersburg, Virginia. Two populations were obtained by crossing insensitive plant introductions with sensitive commercial cultivars. Ratings on the scale of 1 to 5 (1 = 0 to 20% leaf injury, 2 = 21 to 40%, 3 = 41 to 60%, 4 = 61 to 80%, and 5 > 80%) were made on 160 F(2), F(3) progenies, and parental lines. Population mean injury ratings were recorded and estimates of genotypic, environmental, and phenotypic variances were computed. Estimates of heritability in the broadsense and of genetic advance were calculated for each population using F(2) and family component variance methods. Population means of the F(2) and F(3) progenies were not significantly different from their mid-parent values, suggesting that genetic variance was primarily additive. Broad-sense heritability estimates using F(2) variance method ranged from 51.4 to 70.5% and using family component variance method ranged from 62.1 to 75.6%. In this study, the computed genetic advance values closely parallel those of heritability estimated values. The high heritable nature of insensitivity would indicate that effective levels of insensitivity could be transferred to agronomically superior cultivars in a relatively short time.     

Reduction of stem growth and site dependency of leaf injury in Massachusetts black cherries exhibiting ozone symptoms

Leaf ozone symptoms in natural ecosystems are increasingly reported but ozone effects on tree growth and the mediation of site conditions are still little documented. This study tests two hypotheses: (1) leaf injury in black cherry is associated with decline in radial growth, (2) symptoms are more prevalent on mesic sites. On sites supporting black cherry across Massachusetts, tree growth and leaf ozone injury were surveyed in 1996 using a randomized plot network established in the 1960s. Forty-seven percent of 120 trees sampled for ozone symptoms were symptomatic with generally low levels of injury. Over a 31-year period symptomatic trees had 28% lower stem growth rates than asymptomatic trees. Ozone symptom expression was enhanced in well growing stands on moister, cooler and more elevated sites. Ozone appeared to increase environmental stress and had a more pronounced effect on growth in better growing black cherry stands. This complicates management decisions as thinning increases growth and moisture availability.     

Estimating contribution of wildland fires to ambient ozone levels in National Parks in the Sierra Nevada, California

Data from four continuous ozone and weather monitoring sites operated by the National Park Service in Sierra Nevada, California, are used to develop an ozone forecasting model and to estimate the contribution of wildland fires on ambient ozone levels. The analyses of weather and ozone data pointed to the transport of ozone precursors from the Central Valley as an important source of pollution in these National Parks. Comparisons of forecasted and observed values demonstrated that accurate forecasts of next-day hourly ozone levels may be achieved by using a time series model with historic averages, expected local weather and modeled PM values as explanatory variables. Results on fire smoke influence indicated occurrence of significant increases in average ozone levels with increasing fire activity. The overall effect on diurnal ozone values, however, was small when compared with the amount of variability attributed to sources other than fire.     

Consequences of elevated carbon dioxide and ozone for foliar chemical composition and dynamics in trembling aspen (Populus tremuloides) and paper birch (Betula papyrifera).

Atmospheric chemical composition affects foliar chemical composition, which in turn influences the dynamics of both herbivory and decomposition in ecosystems. We assessed the independent and interactive effects of CO2 and O3 fumigation on foliar chemistry of quaking aspen (Populus tremuloides) and paper birch (Betula papyrifera) at a Free-Air CO2 Enrichment (FACE) facility in northern Wisconsin. Leaf samples were collected at five time periods during a single growing season, and analyzed for nitrogen. starch and condensed tannin concentrations, nitrogen resorption efficiencies (NREs), and C:N ratios. Enriched CO2 reduced foliar nitrogen concentrations in aspen and birch; O3 only marginally reduced nitrogen concentrations. NREs were unaffected by pollution treatment in aspen, declined with 03 exposure in birch, and this decline was ameliorated by enriched CO2. C:N ratios of abscised leaves increased in response to enriched CO2 in both tree species. O3 did not significantly alter C:N ratios in aspen, although values tended to be higher in + CO2 + O3 leaves. For birch, O3 decreased C:N ratios under ambient CO2 and increased C:N ratios under elevated CO2. Thus, under the combined pollutants, the C:N ratios of both aspen and birch leaves were elevated above the averaged responses to the individual and independent trace gas treatments. Starch concentrations were largely unresponsive to CO2 and O3 treatments in aspen. but increased in response to elevated CO2 in birch. Levels of condensed tannins were negligibly affected by CO2 and O3 treatments in aspen, but increased in response to enriched CO2 in birch. Results from this work suggest that changes in foliar chemical composition elicited by enriched CO2 are likely to impact herbivory and decomposition, whereas the effects of O3 are likely to be minor, except in cases where they influence plant response to CO2.     

Assessing plant response to ambient ozone: growth of young apple trees in open-top chambers and corresponding ambient air plots

Open-top chambers (OTCs) and corresponding ambient air plots (AA) were used to assess the impact of ambient ozone on growth of newly planted apple trees at the Montague Field research center in Amherst, MA. Two-year-old apple trees (Malus domestica Borkh 'Rogers Red McIntosh') were planted in the ground in circular plots. Four of the plots were enclosed with OTCs where incoming air was charcoal-filtered (CF); four were enclosed with OTCs where incoming air was not charcoal-filtered (NF) and four were not enclosed, allowing access to ambient air conditions (AA). Conditions in both CF and NF OTCs resulted in increased tree growth and changed incidence of disease and arthropod pests, compared to trees in AA. As a result, we were not able to use the OTC method to assess the impact of ambient ozone on growth of young apple trees in Amherst, MA.     

Shoot growth of mature Fagus sylvatica and Picea abies in relation to ozone

Epidemiological analysis of sequential growth data may be a tool in assessing ozone sensitivity of mature trees. Annual shoot growth of mature Fagus sylvatica in 83 Swiss permanent forest observation plots and of Picea abies in 61 plots was evaluated for 11 and 8 consecutive years, respectively, using branches harvested every 4 years. The data were assessed as annual deviation from average growth and related to fructification, ozone, meteorological parameters, and modelled soil water content using a mixed linear model. In beech, a significant association between ozone and shoot growth was observed which corresponded to a 7.4% growth reduction between 0 and 10 ppm h AOT40 (accumulated ozone over threshold 40). This is in the same order of magnitude as the response observed in experiments with seedlings. No interaction was found between ozone and drought parameters. In Norway spruce, shoot growth was neither associated with ozone nor with drought.     

White birch (Betula papyrifera Marshall) foliar litter decomposition in relation to trace metal atmospheric inputs at metal-contaminated and uncontaminated sites near Sudbury, Ontario and Rouyn-Noranda, Quebec, Canada

Decomposition of white birch (Betula papyrifera Marshall) foliar litter was examined at metal-contaminated and uncontaminated sites established along gradients of soil Cu, Ni, Pb and Zn concentrations near Sudbury, Ontario and Rouyn-Noranda, Quebec. Over an 18-month study period, a significantly lower rate of litter mass loss was observed at the Sudbury contaminated site (S1) than at the uncontaminated site (S2). This result was not duplicated at corresponding sites (RN1, RN2) in Rouyn-Noranda, despite similar levels of soil metal contaminants and atmospheric inputs. Concentrations of metals in litter increased at all sites with time. However, the greatest litter Cu and Ni concentrations were observed at S1 (188 and 192 microg/g, respectively), a result of substantial net gains of these elements from atmospheric inputs. On a per hectare basis, Cu accumulation in litter at S1 approached recommended application rates of Cu as copper sulphate for control of fungal diseases in agricultural operations, indicating that the current rate of Cu smelter emissions in Sudbury may cause the observed impairment of decomposition.     

Effects of ozone on growth, yield and leaf gas exchange rates of four Bangladeshi cultivars of rice (Oryza sativa L.)

To assess the effects of tropospheric O₃ on rice cultivated in Bangladesh, four Bangladeshi cultivars (BR11, BR14, BR28 and BR29) of rice (Oryza sativa L.) were exposed daily to charcoal-filtered air or O₃ at 60 and 100 nl l⁻¹ (10:00-17:00) from 1 July to 28 November 2008. The whole-plant dry mass and grain yield per plant of the four cultivars were significantly reduced by the exposure to O₃. The exposure to O₃ significantly reduced net photosynthetic rate of the 12th and flag leaves of the four cultivars. The sensitivity to O₃ of growth, yield and leaf gas exchange rates was not significantly different among the four cultivars. The present study suggests that the sensitivity to O₃ of yield of the four Bangladeshi rice cultivars is greater than that of American rice cultivars and is similar to that of Japanese rice cultivars and that O₃ may detrimentally affect rice production in Bangladesh.     

Effects of ozone on growth, yield and leaf gas exchange rates of two Bangladeshi cultivars of wheat (Triticum aestivum L.)

To clarify the effects of O₃ on crop plants cultivated in Bangladesh, two Bangladeshi wheat cultivars (Sufi and Bijoy) were grown in plastic boxes filled with Andisol and exposed daily to charcoal-filtered air or O₃ at 60 and 100 nl l⁻¹ (10:00-17:00) from 13 March to 4 June 2008. The whole-plant dry mass and grain yield per plant of the two cultivars at the final harvest were significantly reduced by the exposure to O₃. Although there was no significant effect of O₃ on stomatal diffusive conductance to H₂O of flag leaf, net photosynthetic rate of the leaf was significantly reduced by the exposure to O_3. The sensitivity of growth, yield, yield components and leaf gas exchange rates to O₃ was not significantly different between the two cultivars. The results obtained in the present study suggest that ambient levels of O₃ may detrimentally affect wheat production in Bangladesh.     

Assessment of metal availability to vegetation (Betula pendula) in Pb-Zn ore concentrate residues with different features

In this work, characterisation of several ore concentrate remains from an abandoned Pb-Zn mining factory was performed determining chemical and physical properties such as pH, organic carbon content, particle size distribution, total heavy metal content (Pb, Zn, Cu, As and Cd) as well as mineralogical composition which showed, in most cases, the oxidization of the parent ore material (mostly galena: PbS and sphalerite: ZnS) to more mobile fractions as anglesite (PbSO(4)) and goslarite (ZnSO(4)). Moreover, two operational defined extraction procedures commonly used in soil and sediment studies (first and second steps of BCR procedure and DTPA extraction protocol) were applied in the different mining wastes in order to study Pb and Zn mobility and likely bioavailability to Betula pendula growing on the same mining spoils, which presents lead and zinc contents in leaves over ten times background values.     

Protection of ash (Fraxinus excelsior) trees from ozone injury by ethylenediurea (EDU): roles of biochemical changes and decreased stomatal conductance in enhancement of growth

Treatments with ethylenediurea (EDU) protect plants from ozone foliar injury, but the processes underlying this protection are poorly understood. Adult ash trees (Fraxinus excelsior), with or without foliar ozone symptoms in previous years, were treated with EDU at 450ppm by gravitational trunk infusion in May-September 2005 (32.5ppmh AOT40). At 30-day intervals, shoot growth, gas exchange, chlorophyll a fluorescence, and water potential were determined. In September, several biochemical parameters were measured. The protective influence of EDU was supported by enhancement in the number of leaflets. EDU did not contribute its nitrogen to leaf tissue as a fertiliser, as determined from lack of difference in foliar N between treatments. Both biochemical (increase in ascorbate-peroxidase and ascorbic acid, and decrease in apoplastic hydrogen peroxide) and biophysical (decrease in stomatal conductance) processes regulated EDU action. As total ascorbic acid increased only in the asymptomatic trees, its role in alleviating O(3) effects on leaf growth and visible injury is controversial.     

Modelling stomatal responses of spring wheat (Triticum aestivum L. cv. Turbo) to ozone and different levels of water supply

Spring wheat (Triticum aestivum L. cv. Turbo) was exposed to different levels of ozone and water supply in open-top chambers in 1991. Air was charcoal filtered (CF), non-filtered (NF) and CF plus proportional addition of ambient or twice ambient ozone (CF1, CF2). Seasonal means of O(3), taken over 24 h, were 2.3, 20.6, 17.3, and 34.5 nl litre(-1) for CF, NF, CF1 and CF2 treatments, respectively. A split-plot design was used to obtain two levels of water supply: one-half of the pots was irrigated sufficiently not to show any symptoms of drought stress; the others were exposed to low water supply and received 50% of these amounts. Using a steady-state porometer approximately 800 measurements of stomatal conductance (g(s)) were made on flag leaves from 68 to 106 days after sowing. The measurements yielded only small differences of maximum conductance between the two levels of water supply. Therefore, low water supply did not protect wheat plants against ozone injury via reduced stomatal uptake in this experiment. To describe the effects of environmental variables on the stomatal behaviour, boundary-line analysis and non-linear regression analysis were used. Besides microclimatic parameters, the ozone dose of flag leaves was introduced as an independent variable affecting stomatal aperture. A well-defined boundary line for ozone dose was found, suggesting that increasing ozone dose caused stomatal closure in wheat flag leaves. But at high ozone doses, co-acting senescence seems also responsible for the decrease in stomatal conductance. A multiplicative boundary-line model was used to predict stomatal conductance from combinations of environmental variables. In the test carried out with the measurements of stomatal conductance, the model accounted only for 40% of the variation of g(s). Generalized stomatal response patterns of the herbaceous growth form, the dependence of the variables' age and ozone dose and the lack of an important factor influencing stomatal response (water status of the plant) in the model, are suggested as explanations of the poor results of the test.