Role of climate, crown position, tree age and altitude in calculated ozone flux into needles of Picea abies and Pinus cembra: a synthesis

Ozone (O(3)) flux into Norway spruce (Picea abies) and cembran pine (Pinus cembra) needles was estimated under ambient conditions at six rural sites between 580 and 1950 m a.s.l. We also assessed age-related differences in O(3) flux by examining changes in leaf conductance across the life span of Norway spruce. At the leaf level O(3) flux into the needles was effectively controlled by stomatal conductance and, hence by factors such as temperature, irradiance and humidity, which control stomatal conductance. Seasonal variations in O(3) flux were mainly attributed to the course of the prevailing temperature. During the growing season, however, data have emphasised leaf-air vapour pressure difference as the environmental factor most likely to control stomatal conductance and O(3) flux into the needles. In the sun crown stomatal conductance averaged over the growing season decreased with increasing tree age from 42.0+/-3.5 mmol O(3) m(-2) s(-1) in 17-year-old trees to 7.1+/-1.0 mmol O(3) m(-2) s(-1) in 216-year-old trees, indicating that O(3) concentration in the substomatal cavities is higher in young than in old trees. Independent from tree age stomatal conductance and O(3) flux were approximately 50% lower in shade needles as compared to sun-exposed needles. Stomatal conductance was also greater in the current flush (24+/-5.6 mmol O(3) m(-2) s(-1)) and in 1-year old needles (16+/-4 mmol O(3) m(-2) s(-1)) than in older needle age classes (12+/-1 mmol O(3) m(-2) s(-1), averaged across the four older needle age classes). In trees similar in age (60-65 years old) average O(3) flux into sun needles increased from 0.55+/-0.36 nmol m(-2) s(-1) at the valley floor to 0.9 nmol m(-2) s(-1) in 1950 m a.s.l. Cumulative O(3) uptake during the vegetation period increased from 11.4+/-1.7 mol m(-2) in the valley to 14 mol m(-2) at the alpine timberline. Although stomatal conductance provides the principal limiting factor for O(3) flux, additional field research is necessary in order to improve our understanding concerning the quantitative 'physiological threshold dose' which internally can be active and can have adverse effects of O(3) on forest trees.     

Influence of ozone stress on growth processes, yields and grain quality characteristics among soybean cultivars

Field studies were conducted at USDA Beltsville Agricultural Research Center, Beltsville, Maryland, in 1984 and 1985 using open-top chambers to acquire information on the responses of 12 soybean (Glycine max L. Merr.) cultivars to O3 stress and to examine the interactions between maturity groups and O3 stress. Cultivars representing Groups III, IV, and V were exposed for approximately 3 months to charcoal-filtered air (CF) and nonfiltered air plus 40 nl litre(-1) O3 (NF + O3). Ozone was added 6 h d(-1), 5 d week(-1) for 13 weeks. The CF effectively reduced the accumulative oxidant exposure (AOX) to less than 1.0 microl litre(-1) h and the NF + O3 treatment approximately doubled the ambient AOX (16.7 microl litre(-1) h) to about 30 microl litre(-1) h. The AOX estimates the total O3 exposure above 30 nl litre(-1) during an entire growing season. Plant growth rates and relative growth rates were reduced by 17.0 and 14.4%, respectively, when averaged over cultivars. Based on growth rates, the Group III cultivars were the most affected by O3 stress. Averaged over cultivars, leaf expansion rates, leaf conductance, and transpiration rates were lower in the NF + O3 treatment compared to the CF control; however, wide variation was found with the stomatal results from field observations. Combined over years and cultivars, grain yield was reduced by an average of 12.5% by O3 stress with 3 of 12 cultivars showing significant reductions. Grain protein content was increased by 0.7% by O3 stress, but cultivar differences were equal to the differences caused by the O3 treatments. Grain oil content was unchanged by the O3 treatments. Group IV cultivars showed the greatest decrease in grain yield due to O3 stress. Multiple regression analyses were calculated using the difference between the CF and NF + O3 treatment as a measure of O3 stress. Significant positive relationships were found among net assimilation rates, plant growth rates, relative growth rates, and leaf expansion rates, which suggest that growth analysis characteristics would be useful in addition to yield in air pollution tolerance improvement studies with soybeans.     

Simulations of stomatal conductance and ozone uptake to Norway spruce saplings in open-top chambers

A simulation model was developed to estimate the stomatal conductance and ozone flux to Norway spruce saplings in open-top chambers. The model was parameterized against needle conductance measurements that were made on 4-6-year-old spruce saplings, grown in open-top chambers, in July-September during three different seasons. The spruce saplings were either maintained well watered or subject to a 7-8 week drought period in July-September each year. The simulated conductance showed a good agreement with the measured conductance for the well-watered as well as the drought stress-treated saplings. The simulations were significantly improved when different vapour pressure deficit (VPD) functions were applied for well-watered and drought-stressed spruce saplings. The cumulated ozone uptake which was calculated from the conductance simulations showed less variation between years, compared to the cumulative ozone exposure index AOT40 (accumulated exposure over a threshold of 40 ppb or nl l(-1)) for the corresponding time periods. Measurements in May 1995 demonstrated the occurrence of long-term 'memory-effects' from the drought stress treatments on the conductance. Memory-effects need to be considered when simulation models for stomatal conductance are to be applied to long-lived forest trees under a multiple stress situation.     

Responses of bean (Phaseolus vulgaris L. cv. Pros) to chronic ozone exposure at two levels of atmospheric ammonia

Plants of bean (Phaseolus vulgaris cv. Pros) were exposed to a range of O3 concentrations up to 70 nl litre(-1) for 9 h day(-1) in the presence (45 nl litre(-1)) and absence (21 nl litre(-1)) of enhanced NH3 in 12 open-top chambers. Treatment effects on visible injury, growth and yield were assessed after 49 (intermediate harvest) and 62 days of exposure (final harvest). The proportion of leaves with visible injury at final harvest increased with increasing concentrations of O3. Enhanced NH3 did not cause any symptoms and did not affect injury by O3. The estimated seasonal mean concentration corresponding with 5% injury was circa 23 nl litre(-1) O3. Biomass production and green pod yield decreased with increasing concentrations of O3 and were generally stimulated by enhanced NH3 at both harvests. Adverse effects of O3 on biomass and pod yield did not depend on the NH3 level. Relative yield response to increasing 9-h daily mean O3 concentrations was nonlinear and yield losses of 5 and 10% were calculated to occur at seasonal daytime mean concentrations of 27 and 33 nl litre(-1) O3, respectively. Linear regression showed that the Accumulated exposures Over a Threshold of 30 (AOT30) and 40 nl litre(-1) (AOT40) O3 performed equally well. The estimated accumulated O3 exposures corresponding with a yield loss of 5% were 1600 nl litre(-1) h for AOT30 and 400 nl litre(-1) h for AOT40. The results are discussed in relation to the long-term critical level that is used as a guideline to protect crops against adverse effects by O3.     

Ozone exposure-response relationships for biomass and root/shoot ratio of beech (Fagus sylvatica), ash (Fraxinus excelsior), Norway spruce (Picea abies) and Scots pine (Pinus sylvestris)

Current-year seedlings of beech, ash, Norway spruce and Scots pine were exposed during one growing season to different, but moderate, ozone (O(3)) scenarios representative for Switzerland (50, 85, 100% ambient, 50% ambient+30 nl l(-1)) in open-top chambers (OTCs) and to ambient O(3) concentrations in the field. Biomass significantly decreased with increasing O(3) dose in all species except for spruce. Losses of 25.5% (ash), 17.4% (beech), 9.9% (Scots pine) were found per 10 microl l(-1) h accumulated O(3) exposure over a threshold concentration of 40 nl l(-1) during daylight hours (AOT40). Ratios of root/shoot biomass (RSR) also significantly decreased with increasing AOT40 levels in beech and ash, but not in Norway spruce and Scots pine. The data show that the deciduous species beech and ash were more susceptible to O(3) with respect to RSR and biomass than the coniferous species Norway spruce and Scots pine.     

Photosynthetic and stomatal conductance responses of Norway spruce and beech to ozone, acid mist and frost--a conceptual model

Two-year-old beech and Norway spruce seedlings were exposed to a combination of ozone and acid mist treatments in open-top chambers in Scotland during the months of July through to September 1988. Replicate pairs of chambers received charcoal-filtered air (control), ozone-enriched air (140 nl ozone litre(-1)) or 140 nl ozone litre(-1) plus a synthetic acid mist (pH 2.5) composed of ammonium nitrate and sulphuric acid. Field measurements of assimilation and stomatal conductance were made during August. In addition, measurements of assimilation and conductance were made during September in the laboratory. Light response curves of assimilation and conductance were determined using a GENSTAT nonrectangular hyperbolic model. During February 1988/9 the Norway spruce were subject to a four day warming period at 12 degrees C and the light response of assimilation determined. The same plants were then subject to a 3-h night-time frost of -10 degrees C. The following day the time-course of the recovery of assimilation was determined. It was found that ozone fumigation did not influence the light response of assimilation of beech trees in the field, although stomatal conductance was reduced in the ozone-fumigated trees. The rate of light-saturated assimilation of Norway spruce was increased by ozone fumigation when measured in the field. Measurements of assimilation of Norway spruce made during the winter showed that prior to rewarming there was no difference in the rate of light-saturated assimilation for control and ozone-fumigated trees. However, the ozone plus acid mist-treated trees exhibited a significantly higher rate. The 4-day period of warming to 12 degrees C increased the rate of light-saturated assimilation in all treatments but only the ozone plus acid mist-treated trees showed a significant increase. Following a 3-h frost to -10 degrees C the control trees exhibited a reduction in the rate of light-saturated assimilation (Amax) to 80% of the pre-frost value. In comparison, following the frost, the ozone-fumigated trees showed an Amax of 74% of the pre-frost value. The ozone plus acid mist-treated trees showed an Amax of 64% of the pre-frost trees. The time taken for Amax to attain 50% of the pre-frost value increased from 30 min (control) to 85 min for ozone-fumigated trees to 190 min (ozone plus acid mist). These results are discussed in relation to the impact of mild, short-term frosts, which are known to occur with greater frequency than extreme, more catastrophic frost events. A simple conceptual framework is proposed to explain the variable results obtained in the literature with respect to the impact of ozone upon tree physiology.     

Ozone uptake by an evergreen forest canopy: temporal variation and possible mechanisms

Patterns of ozone concentration ([O(3)]), O(3) deposition velocity (v(d)) and O(3) flux (F(c)) over an evergreen forest canopy are shown in relation to measuring method, physiological activity of the trees, and time of year. The gradient and eddy correlation methods were compared and showed similar diel v(d) patterns. Daytime F(c) was correlated with CO(2) and water vapour fluxes, while no correlation between [O(3)] in the range 10-70 ppb (nl l(-1)) and F(c) was seen in this study. F(c) was primarily driven by stomatal conductance, reactions with surfaces, particles and gases, and not by [O(3)]. On a monthly basis, [O(3)] was always highest in the afternoon while v(d) was typically higher in the morning, resulting in an equal F(c) over the day. Night-time F(c) was more than half of the daytime O(3) flux. The data reveal the importance of emissions of nitric oxide and terpenes as O(3) removal factors in evergreen forest dominated by Norway spruce.     

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.     

Visible and microscopic injury in leaves of five deciduous tree species related to current critical ozone levels

Because the current critical level of ozone (O(3)) for forest trees is based only on one species, the responses of five deciduous tree species were differentiated in a climate chamber experiment. The number of symptomatic leaves per tree was significantly increased, and stomatal conductance was decreased under 50% ambient+30 nl l(-1) O(3) as compared to 'normal' senescence at 50% ambient [O(3)]. Species with a high stomatal conductance did not show earlier or more leaf injury symptoms. The additional 30 nl l(-1) O(3) induced specific pectinaceous cell wall protrusions, phenolic cell wall incrustations, tonoplast vesicles, and inhomogeneous, condensed/precipitated phenolic material in the vacuoles. Due to added O(3), cell senescence was accelerated with increased electron-density of the cytoplasm, and initial chloroplast degeneration. The slow degeneration process started in mesophyll cells, and expanded into epidermal and finally guard cells. Because of the large variance in biomass between individuals and species, the current critical level is supported by the assessment of visible leaf symptoms rather than growth reduction.     

A statistical approach to estimate O3 uptake of ponderosa pine in a mediterranean climate

In highly polluted sites, stomatal behavior is sluggish with respect to light, vapor pressure deficit, and internal CO2 concentration (Ci) and poorly described by existing models. Statistical models were developed to estimate stomatal conductance (gs) of 40-year-old ponderosa pine at three sites differing in pollutant exposure for the purpose of calculating O3 uptake. Gs was estimated using julian day, hour of day, pre-dawn xylem potential and photosynthetic photon flux density (PPFD). The median difference between estimated and observed field gs did not exceed 10 mmol H2O m(-2) s(-1), and estimated gs within 95% confidence intervals. 03 uptake was calculated from hourly estimated gs, hourly O3 concentration, and a constant to correct for the difference in diffusivity between water vapor and 03. The simulation model TREGRO was also used to calculate the cumulative 03 uptake at all three sites. 03 uptake estimated by the statistical model was higher than that simulated by TREGRO because gas exchange rates were proportionally higher. O3 exposure and uptake were significantly correlated (r2>0.92), because O3 exposure and gs were highly correlated in both statistical and simulation models.     

Modelling of stomatal conductance and ozone flux of Norway spruce: comparison with field data

It has been proposed that stomatal flux of ozone would provide a more reliable basis than ozone exposure indices for the assessment of the risk of ozone damage to vegetation across Europe. However, implementation of this approach requires the development of appropriate models which need to be rigorously tested against actual data collected under field conditions. This paper describes such an assessment of the stomatal component of the model described by Emberson et al. (2000. Modelling stomatal ozone flux across Europe. Environmental Pollution 110). Model predictions are compared with field measurements of both stomatal conductance (g(s)) and calculated ozone flux for shoots of mature Norway spruce (Picea abies) growing in the Tyrol Mountains in Austria. The model has been developed to calculate g(s) as a function of leaf phenology and four environmental variables: photosynthetic flux density (PFD), temperature, vapour pressure deficit (VPD) and soil moisture deficit (SMD). The model was run using climate data measured on site, although the SMD component was omitted since the necessary data were not available. The model parameterisation for Norway spruce had previously been collected from the scientific literature and therefore established independently from the measurement study. Overall, strong associations were found between model predictions and measured values of stomatal conductance to ozone (GO(3)) and calculated stomatal ozone flux (FO(3)). Average diurnal profiles of GO(3) and FO(3) showed good agreement between the field data and modelled values except during the morning period of 1990. The diurnal pattern of ozone flux was determined primarily by PFD and VPD, as there was little diurnal variation in ozone concentration. In general, the model predicted instances of high ozone flux satisfactorily, indicating its potential applicability in identifying areas of high ozone risk for this species.     

Response of cellular antioxidants to ozone in wheat flag leaves at different stages of plant development

The effect of ozone (O3) on growth, yield and foliar antioxidants of spring wheat (Triticum aestivum L. cv. Turbo) was investigated in 1990 and 1991 in Braunschweig, Germany. Plants were grown full-season in pots in open-top chambers ventilated with charcoal-filtered (CF) air to which one or two levels of O3 were added. Mean 8 h day(-1) (10.00-18.00 h) O3 concentrations in the CF and CF + O3 treatments were 5.9, 61.2 and 92.5 nl litre(-1) in 1990, and 4.7 and 86.4 nl litre(-1) in 1991. Plants that received the high O3 level showed symptoms of premature senescence of the oldest leaves and yield reductions in both growing seasons. The contents of ascorbate and glutathione and the enzyme activities of ascorbate peroxidase and glutathione reductase were measured in symptomless flag leaves in weekly intervals before and after the beginning of anthesis. Leaf age had a significant effect on all antioxidants investigated. The O3 exposures of about 90 nl litre(-1) increased the activity of ascorbate peroxidase and the concentration of glutathione, but there were no pollutant effects on ascorbate content and glutathione reductase activity. Measurements of the antioxidant levels throughout one day showed no clear indications of diurnal changes in the antioxidative capacity in wheat flag leaves. The results are discussed in relation to the role of antioxidants in O3 detoxification.     

Ozone-induced changes in CO2 assimilation, O2 evolution and chlorophyll a fluorescence transients in barley

Spring barley (Hordeum vulgare cv. Klaxon) plants, 9 days old, were exposed to 0.05, 0.10 or 0.15 microl litre(-1) ozone (O3) for 12 days. Fumigation was administered for 7 h between 9.00 h and 16.00 h each day. Using conventional IRGA equipment, the carbon dioxide exchange rate (CER) was shown to decrease with increasing concentration of O3 during the exposure period, falling to 60% of the control value at the highest O3 concentration. Transpiration rates and stomatal conductance showed similar trends. Light saturation curves, obtained using a leaf disc oxygen electrode, demonstrated that O3-treated leaves had lower apparent quantum yields (QY) and generally lower rates of O2 evolution at saturating light and CO2 levels. Oscillations in chlorophyll a fluorescence, normally observed in control plants, could not be detected after O3 treatment and could only be restored to some extent by feeding the phosphate sequestering agent D-mannose to the leaves.     

Whole-plant growth and leaf formation in ozonated hybrid poplar (Populus x euramericana)

Seasonal growth was studied in potted cuttings of hybrid poplar (one clone of Populus x euramericana) either exposed to ozone in filtered air (0 = control, 0.05, 0.10 microl litre(-1)) or in ambient air (mean = 0.03 microl litre(-1)). Only at 0.10 microl litre(-1) was biomass production reduced and related to leaf loss rather than leaf formation, since the latter was similar in all treatments. Stem length at 0.10 microl litre(-1) approached that of the control, whereas starch concentration in the green stem bark tended to be reduced, as were the ratios of stem weight/length and root/shoot biomass. The changes in carbon allocation and biomass production gradually became established during the second half of the growing season. At the altered carbon allocation at 0.10 microl litre(-1), the ratio of whole-plant production/attached foliage area resembled that of the other O(3) regimes. However, the latter ratio was strongly reduced at 0.10 microl litre(-1) when calculated on the basis of the potential foliage area, as compensated for the O(3)-induced leaf loss. Thus the carbon return/cost balance of the totally formed foliage was low, although the relative-growth rate of ozonated plants temporarily reached that of the control. The relation between leaf differentiation under ozonation (lowered stomatal density) and whole-plant production remains uncertain. The plant behavior found is discussed in terms of passive response or acclimatization to O(3) stress.     

Responses to ozone of insects feeding on a crop and a weed species

The influence of ozone on insect herbivore growth and population development was investigated. Fumigation of both pea (Pisum sativum L.) and dock (Rumex obtusifolius L.) at a range of O(3) concentrations between 21-206 nl litre(-1) produced changes in mean relative growth rates of the aphids Acyrthosiphon pisum Harris and Aphis rumicis L. of between 24 and -6% relative to controls. However, there was no evidence of a dose-related response to O(3) fumigation and no clear differences in aphid response when fumigated with the plant on prefumigated or previously unfumigated plant material. It is suggested that this may, in part, be due to the presence of NO contamination during O(3) fumigation. However, the MRGR of dock aphids was found to be greater on new compared to old leaves as well as the increase on the new growth and decrease on the old growth of fumigated plants relative to unfumigated controls. The size of egg batches of the chrysomelid beetle Gastrophysa viridula Degeer were found to be larger, survival and productivity of larvae was higher, and the food consumption lower on R. obtusifolius fumigated with 70 nl litre(-1) O(3) compared with unfumigated controls. This meant that these beetle larvae consumed less leaf area per mg of production on fumigated leaves probably because of their better nutritional quality and/or reduced leaf defences. However, the rate of development of larvae was similar on fumigated and control plants.     

Effects of elevated CO2 concentration on the polyamine levels of field-grown soybean at three O3 regimes

Effects of increased ozone (O3) and carbon dioxide (CO2) on polyamine levels were determined in soybean (Glycine max L. Merr. cv. Clark) grown in open-top field chambers. The chamber treatments consisted of three O3 regimes equal to charcoal filtered (CF), non-filtered (NF), and non-filtered plus 40 nl litre(-1) O3 and CO2 treatments equal to 350, 400 and 500 microl litre(-1) for a total of nine treatments. Leaf samples were taken at three different times during the growing season. Examination of growth and physiological characteristics, such as photosynthesis, stomatal resistance, and shoot weight, revealed that increasing CO2 ameliorated the deleterious effects of increased O3. Results from the initial harvest, at the pre-flowering growth stage (23 days of treatment), showed that increasing O3 at ambient CO2 caused increases in putrescine (Put) and spermidine (Spd) of up to six-fold. These effects were lessened with increased CO2. Elevated CO2 increased polyamines in plants treated with CF air, but had no effect in the presence of ambient or enhanced O3 levels. Leaves harvested during peak flowering (37 days of treatment) showed O3-induced increases in Put and Spd at ambient CO2 concentrations. However, increased CO2 levels inhibited this response by blocking the O3-induced polyamine increase. Leaves harvested during the pod fill stage (57 days of treatment) showed no significant O3 or CO2 effects on polyamine levels. Our results demonstrate that current ambient O3 levels induce the accumulation of Put and Spd early in the growing season and that further increases in O3 could result in even greater polyamine increases. These results are consistent with a possible antiozonant function for polyamines. The ability of increased CO2 to protect soybeans from O3 damage, however, does not appear to involve polyamine accumulation.     

Effects of elevated O3 and CO2 on chlorophyll fluorescence and gas exchange in Scots pine during the third growing season

Naturally regenerated, 30-year-old Scots pines (Pinus Sylvestris L.) were grown in open-top chambers and exposed in situ to doubled ambient O(3), doubled ambient CO(2) and a combination of elevated O(3) and CO(2) from 15 April to 15 September for three growing seasons (1994-1996). To examine the effects of O(3) and/or CO(2) on photosynthesis, chlorophyll a fluorescence and gas exchange were measured simultaneously. Doubled ambient O(3) significantly decreased the rates of photosynthesis at all levels of photon flux density. This was related mainly to a significant decrease in the photochemical efficiency of photosystem II (PS II) and the rate of whole electron transport, rather than to a decrease in stomatal conductance. When measurements were made at doubled ambient concentration of CO(2) (700 micromol mol(-1)), doubled ambient CO(2) treatment did not lead to a significant change in the intrinsic capacity of photosynthesis, as manifested by no changes in PS II, the rate of electron transport, the maximal rate of photosynthesis and the apparent quantum yield of CO(2) assimilation. However, elevated CO(2) increased the sensitivity of stomatal conductance to light and decreased maximal stomatal conductance. When O(3) and CO(2) were combined, the O(3)-induced decrease in photosynthesis rate was reduced significantly by a high concentration of CO(2). This may be partly related to the decrease in stomatal conductance induced by the high concentration of CO(2). The complete mechanism behind this interaction is, however, still unclear.     

Ozone induced leaf loss and decreased leaf production of European Holly (Ilex aquifolium L.) over multiple seasons

European Holly (Ilex aquifolium L.) was used to study the impact of one short (28 day) ozone fumigation episode on leaf production, leaf loss and stomatal conductance (g(s)), in order to explore potential longer term effects over 3 growing seasons. Young I. aquifolium plants received an episode of either charcoal-filtered air or charcoal-filtered air with 70 nl l(-1) O(3) added for 7 h d(-1) over a 28 day period from June 15th 1996, then placed into ambient environment, Stoke-on-Trent, U.K. Data were collected per leaf cohort over the next three growing seasons. Ozone exposure significantly increased leaf loss and stomatal conductance and reduced leaf production over all subsequent seasons. Impact of the initial ozone stress was still detected in leaves that had no direct experimental ozone exposure. This study has shown the potential of ozone to introduce long-term phenological perturbations into ecosystems by influencing productivity over a number of seasons.     

Joint action of ozone and hydrogen fluoride on foliar senescence in maize

Maize (Zea mays, L.) plants were exposed intermittently to O(3), HF or both pollutants and the progression of foliar senescence was followed by measuring chlorophyll loss, membrane breakdown and changes in stomatal conductance. At concentrations insufficient to cause foliar symptoms (0.06 microl O(3) litre(-1) and 1.0 microg Fm(-3)), exposures to HF had little or no effect, whereas O(3) exposures accelerated the rate of senescence. The rapid rate of senescence produced by O(3) was moderated if the plants were also exposed to HF. Topical application of 6-benzyladenine (BA) prior to pollutant exposures delayed senescence in all plants and completely prevented the O(3)-induced acceleration of senescence.     

Dynamics of biomass partitioning in field-grown radish varieties, treated with ethylenediurea

During the growing season of 1990, five staggered crops of radish (Raphanus sativus L.) were grown in the field, using the cultivars 'Cherry Belle', 'Red Prince', and 'Red Devil B'. Half of the plants received a soil drench (100 ml plant(-1); 100 mg litre(-1) of ethylenediurea (EDU) once, early in plant development. Destructive harvests were carried out at 2-day intervals during vegetative development. Non-linear growth kinetics, derived from Richards' function, were fitted to the dry weight data of the total plant, main organs (shoot and hypocotyl) and to the dry weight ratio between below-ground and above-ground organs. Estimating the parameters of these non-linear functions and testing their differences between EDU-treated and untreated plants unveiled biologically meaningful information on the impact of different levels of ambient ozone (O(3)) during the growth periods. The modified function which was applied to the data of biomass partitioning between the major plant parts was more powerful in detecting transient alterations in assimilate allocation compared to the growth dynamics of individual plant organs. At low levels of O(3), biomass partitioning towards the below-ground sink organs was slightly delayed and finally restricted in EDU-treated plants. When ambient O(3) reached moderate levels, which did not cause visible foliar injury, assimilate partitioning between organs was only insignificantly altered during early growth when EDU-treatments were compared. As growth progressed, however, less assimilates were allocated towards the hypocotyl and roots in the plants not protected by EDU. This pattern was similar in all cultivars tested, but was smallest in 'Cherry Belle', which is known to be sensitive to O(3) with respect to foliar injury. During the 15- to 19-day periods of rapid growth, the O(3)-exposure >80 nl litre(-1) ranged from 0.015 to 0.209 microl litre(-1) O(3) h, which corresponds to 7 h d(-1) mean values between 40 and 50 nl litre(-1) O(3), confirming that ambient ozone did not exceed a moderate level in this study.