The Greenhouse effect: impacts of ultraviolet-B (UV-B) radiation, carbon dioxide (CO2), and ozone (O3) on vegetation

There is a fast growing and an extremely serious international scientific, public and political concern regarding man's influence on the global climate. The decrease in stratospheric ozone (O3) and the consequent possible increase in ultraviolet-B (UV-B) is a critical issue. In addition, tropospheric concentrations of 'greenhouse gases' such as carbon dioxide (CO2), nitrous oxide (N2O) and methane (CH4) are increasing. These phenomena, coupled with man's use of chlorofluorocarbons (CFCs), chlorocarbons (CCs), and organo-bromines (OBs) are considered to result in the modification of the earth's O3 column and altered interactions between the stratosphere and the troposphere. A result of such interactions could be the global warming. As opposed to these processes, tropospheric O3 concentrations appear to be increasing in some parts of the world (e.g. North America). Such tropospheric increases in O3 and particulate matter may offset any predicted increases in UV-B at those locations. Presently most general circulation models (GCMs) used to predict climate change are one- or two-dimensional models. Application of satisfactory three-dimensional models is limited by the available computer power. Recent studies on radiative cloud forcing show that clouds may have an excess cooling effect to compensate for a doubling of global CO2 concentrations. There is a great deal of geographic patchiness or variability in climate. Use of global level average values fails to account for this variability. For example, in North America: 1. there may be a decrease in the stratospheric O3 column (1-3%); however, there appears to be an increase in tropospheric O3 concentrations (1-2%/year) to compensate up to 20-30% loss in the total O3 column; 2. there appears to be an increase in tropospheric CO2, N2O and CH4 at the rate of roughly 0.8%, 0.3% and 1-2%, respectively, per year; 3. there is a decrease in erythemal UV-B; and 4. there is a cooling of tropospheric air temperature due to radiative cloud forcing. The effects of UV-B, CO2 and O3 on plants have been studied under growth chamber, greenhouse and field conditions. Few studies, if any, have examined the joint effects of more than one variable on plant response. There are methodological problems associated with many of these experiments. Thus, while results obtained from these studies can assist in our understanding, they must be viewed with caution in the context of the real world and predictions into the future. Biomass responses of plants to enhanced UV-B can be negative (adverse effect); positive (stimulatory effect) or no effect (tolerant). Sensitivity rankings have been developed for both crop and tree species. However, such rankings for UV-B do not consider dose-response curves. There are inconsistencies between the results obtained under controlled conditions versus field observations. Some of these inconsistencies appear due to the differences in responses between cultivars and varieties of a given plant species; and differences in the experimental methodology and protocol used. Nevertheless, based on the available literature, listings of sensitive crop and native plant species to UV-B are provided. Historically, plant biologists have studied the effects of CO2 on plants for many decades. Experiments have been performed under growth chamber, greenhouse and field conditions. Evidence is presented for various plant species in the form of relative yield increases due to CO2 enrichment. Sensitivity rankings (biomass response) are agein provided for crops and native plant species. However, most publications on the numerical analysis of cause-effect relationships do not consider sensitivity analysis of the mode used. Ozone is considered to be the most phytotoxic regional scale air pollutant. In the pre-occupation of loss in the O3 column, any increases in tropospheric O3 concentrations may be undermined relative to vegetation effects. As with the other stress factors, the effects of O3 have been studied both under controlled and field conditions. Thboth under controlled and field conditions. The numerical explanation of cause-effect relationships of O3 is a much debated subject at the present time. Much of the controversy is directed toward the definition of the highly stochastic, O3 exposure dynamics in time and space. Nevertheless, sensitivity rankings (biomass response) are provided for crops and native vegetation. The joint effects of UV-B, CO2 and O3 are poorly understood. Based on the literature of plant response to individual stress factors and chemical and physical climatology of North America, we conclude that nine different crops may be sensitive to the joint effects: three grain and six vegetable crops (sorghum, oat, rice, pea, bean, potato, lettuce, cucumber and tomato). In North America, we consider Ponderosa and loblolly pines as vulnerable among tree species. This conclusion should be moderated by the fact that there are few, if any, data on hardwood species. In conclusion there is much concern for global climate change and its possible effects on vegetation. While this is necessary, such a concern and any predictions must be tempered by the lack of sufficient knowledge. Experiments must be designed on an integrated and realistic basis to answer the question more definitively. This would require very close co-operation and communication among scientists from multiple disciplines. Decision makers must realize this need.     

Stratospheric ozone depletion, UV-B radiation and crop disease

Ultraviolet-B radiation (UV-B: 290-315 nm) is expected to increase as the result of stratospheric ozone depletion. Within the environmental range, UV-B effects on host plants appear to be largely a function of photomorphogenic responses, while effects on fungal pathogens may include both photomorphogenesis and damage. The effects of increased UV-B on plant-pathogen interactions has been studied in only a few pathosystems, and have used a wide range of techniques, making generalisations difficult. Increased UV-B after inoculation tends to reduce disease, perhaps due to direct damage to the pathogen, although responses vary markedly between and within pathogen species. Using Septoria tritici infection of wheat as a model system, it is suggested that even in a species that is inherently sensitive to UV-B, the effects of ozone depletion in the field are likely to be small compared with the effects of variation in UV-B due to season and varying cloud. Increased UV-B before inoculation causes a range of effects in different systems, but an increase in subsequent disease is a common response, perhaps due to changes in host surface properties or chemical composition. Although it seems unlikely that most crop diseases will be greatly affected by stratospheric ozone depletion within the limits currently expected, the lack of a detailed understanding of the mechanisms by which UV-B influences plant-pathogen interactions in most pathosystems is a significant limit to such predictions.     

The impact of UV-B radiation and ozone on terrestrial vegetation

Although terrestrial vegetation has been exposed to UV-B radiation and ozone over the course of evolutionary history, it is essential to view the effects on vegetation of changing levels of these factors in the context of other features of climate change, such as increasing CO(2) levels and changes in temperature and precipitation patterns. Much of our understanding of the impacts of increased UV-B and ozone levels has come from studies of the effects of each individual factor. While such information may be relevant to a wider understanding of the roles that these factors may play in climate change, experience has shown that the interactions of environmental stresses on vegetation are rarely predictable. A further limitation on the applicability of such information results from the methodologies used for exposing plants to either factor. Much of our information comes from growth chamber, greenhouse or field studies using experimental protocols that made little or no provision for the stochastic nature of the changes in UV-B and ozone levels at the earth's surface, and hence excluded the roles of repair mechanisms. As a result, our knowledge of dose-response relationships under true field conditions is both limited and fragmentary, given the wide range of sensitivities among species and cultivars. Adverse effects of increased levels of either factor on vegetation are qualitatively well established, but the quantitative relationships are far from clear. In both cases, sensitivity varies with stage of plant development. At the population and community levels, differential responses of species to either factor has been shown to result in changes in competitiveness and community structure. At the mechanistic level, ozone generally inhibits photosynthetic gas exchange under both controlled and field conditions, and although UV-B is also inhibitory in some species under controlled conditions, others appear to be indifferent, particularly in the field. Both factors affect metabolism; a common response is increased secondary metabolism leading to the accumulation of phenolic compounds that, in the case of UV-B, offer the leaf cell some protection from radiation. Virtually no information is available about the effects of simultaneous or sequential exposures. Since both increased surface UV-B and ozone exposures have spatial and temporal components, it is important to evaluate the different scenarios that may occur, bearing in mind that elevated daytime ozone levels will attenuate the UV-B reaching the surface to some extent. The experimentation needed to acquire unequivocal effects data that are relevant to field situations must therefore be carried out using technologies and protocols that focus on quantification of the interactions of UV-B and ozone themselves and their interactions with other environmental factors.     

Assessment of the impact of rising carbon dioxide and other potential climate changes on vegetation

The projected doubling of current levels of atmospheric carbon dioxide concentration ([CO(2)]) during the next century along with increases in other radiatively active gases have led to predictions of increases in global air temperature and shifts in precipitation patterns. Additionally, stratospheric ozone depletion may result in increased ultraviolet-B (UV-B) radiation incident at the Earth's surface in some areas. Since these changes in the Earth's atmosphere may have profound effects on vegetation, the objectives of this paper are to summarize some of the recent research on plant responses to [CO(2)], temperature and UV-B radiation. Elevated [CO(2)] increases photosynthesis and usually results in increased biomass, and seed yield. The magnitude of these increases and the specific photosynthetic response depends on the plant species, and are strongly influenced by other environmental factors including temperature, light level, and the availability of water and nutrients. While elevated [CO(2)] reduces transpiration and increases photosynthetic water-use efficiency, increasing air temperature can result in greater water use, accelerated plant developmental rate, and shortened growth duration. Experiments on UV-B radiation exposure have demonstrated a wide range of photobiological responses among plants with decreases in photosynthesis and plant growth among more sensitive species. Although a few studies have addressed the interactive effects of [CO(2)] and temperature on plants, information on the effects of UV-B radiation at elevated [CO(2)] is scarce. Since [CO(2)], temperature and UV-B radiation may increase concurrently, more research is needed to determine plant responses to the interactive effects of these environmental variables.     

Mycorrhizal mediation of plant response to atmospheric change: Air quality concepts and research considerations

The term 'global climate change' encompasses many physical and chemical changes in the atmosphere that have been induced by anthropogenic pollutants. Increases in concentrations of CO2 and CH4 enhance the 'greenhouse effect' of the atmosphere and may contribute to changes in temperature and precipitation patterns at the earth's surface. Nitrogen oxides and SO2 are phytotoxic and also react with other pollutants to produce other phytotoxins in the troposphere such as O3 and acidic substances. However, release of chlorofluorocarbons into the atmosphere may cause depletion of stratospheric O3, increasing the transmittance of ultraviolet-B (UV-B) radiation to the earth's surface. Increased intensities of UV-B could affect plants and enhance photochemical reactions that generate some phytotoxic pollutants. The role of mycorrhizae in plant responses to such stresses has received little attention. Although plans for several research programs have acknowledged the importance of drought tolerance and soil fertility in plant responses to atmospheric stresses, mycorrhizae are rarely targeted to receive specific investigation. Most vascular land plants form mycorrhizae, so the role of mycorrhizae in mediating plant responses to atmospheric change may be an important consideration in predicting effects of atmospheric changes on plants in managed and natural ecosystems.     

Interactive effects of elevated ozone and carbon dioxide on growth and yield of leaf rust-infected versus non-infected wheat

Spring wheat (Triticum aestivum L. cv. Turbo) was grown from seedling emergence to maturity (129 days) in chambers simulating the physical climate and ozone pollution of a field site in Northern Germany from 1 April to 31 July with a mean 1-h daily maximum of 61.5-62.4 nl l(-1) ozone compared to a constant low level of 21.5-22.8 nl l(-1) ozone. The two ozone levels were combined with either a current (374.1-380.2 microl l(-1)) or enriched (610.6-615.0 microl l(-1)) CO(2) atmosphere. Additionally, a leaf rust epidemic (Puccinia recondita f. sp. tritici) was induced at tillering stage by repeated re-inoculations with the inoculum formed on the plants. Leaf rust disease was strongly inhibited by ozone, but largely unaffected by elevated CO(2). Ozone damage on leaves was strongly affected by CO(2) and infection. On infected plants, ozone lesions appeared 2-4 weeks earlier and were up to fourfold more severe compared to non-infected plants. Elevated CO(2) did not delay the onset of ozone lesions but it significantly reduced the severity of leaf damage. It also enhanced the photosynthetic rate of flag leaves and increased the water use efficiency, biomass formation and grain yield. The relative increases in growth and yield induced by CO(2) were much larger on ozone-stressed than on non-stressed plants. Both ozone and fungal infection reduced biomass formation, number of grains per plant, thousand grain weight and grain yield; however, adverse effects of leaf rust infection were more severe. Elevated CO(2) largely equalized the negative effects of ozone on the photosynthetic rate, growth and yield parameters, but was not capable of compensating for the detrimental effects of fungal infection. The data imply that the impact of ozone in the field cannot be estimated without considering the predisposing effects deriving from fungal infections and the compensating effects deriving from elevated CO(2).     

Depletion of stratospheric ozone over the Antarctic and Arctic: responses of plants of polar terrestrial ecosystems to enhanced UV-B, an overview

Depletion of stratospheric ozone over the Antarctic has been re-occurring yearly since 1974, leading to enhanced UV-B radiation. Arctic ozone depletion has been observed since 1990. Ozone recovery has been predicted by 2050, but no signs of recovery occur. Here we review responses of polar plants to experimentally varied UV-B through supplementation or exclusion. In supplementation studies comparing ambient and above ambient UV-B, no effect on growth occurred. UV-B-induced DNA damage, as measured in polar bryophytes, is repaired overnight by photoreactivation. With UV exclusion, growth at near ambient may be less than at below ambient UV-B levels, which relates to the UV response curve of polar plants. UV-B screening foils also alter PAR, humidity, and temperature and interactions of UV with environmental factors may occur. Plant phenolics induced by solar UV-B, as in pollen, spores and lignin, may serve as a climate proxy for past UV. Since the Antarctic and Arctic terrestrial ecosystems differ essentially, (e.g. higher species diversity and more trophic interactions in the Arctic), generalization of polar plant responses to UV-B needs caution.     

Interactive effects of O3 exposure on California black oak (Quercus kelloggii Newb.) seedlings with and without N amendment

We examined the short-term separate and combined effects of simulated nitrogen (N) deposition (fertilization) and ozone (O(3)) exposure on California black oak seedlings (Quercus kelloggii Newb.), an ecologically important tree of the San Bernardino Mountains downwind of Los Angeles. Realistic concentrations of O(3) were found to cause statistically and biologically significant negative effects on plant health, including lowered photosynthetic ability, lowered water use efficiency, and increased leaf chlorosis and necrosis. When subjected to abrupt changes in light levels, O(3)-exposed plants showed both a slower and smaller response than O(3)-free plants. Fertilized plants exhibited a significantly greater pre- to post-treatment decline in A at saturated [CO(2)] and a significantly lower level of post-treatment chlorosis than unfertilized plants. Fertilization tended to reduce plant sensitivity to O(3).     

The effect of reciprocal treatments with ozone and ultraviolet-B radiation on photosynthesis and growth of perennial grass Elymus athericus

The impact on plant growth of the simultaneously changing factors of the global climate, rising tropospheric O3 concentrations and increasing UV-B radiation fluxes, has been tested in a combined glasshouse and growth chamber experiment. The saltmarsh grass species Elymus athericus was sequentially fumigated for two weeks with O3 and for another two weeks irradiated with UV-B (vv). Exposure to elevated UV-B did not negatively affect photosynthesis or plant growth. Fumigation with O3 had a depressing effect on net photosynthesis, the number and biomass of flowers, the number of leaves and the number of shoots. O3-induced damage only was observed in plants which had been fumigated during the last two weeks of the experiment. Since interactive responses were not observed, results suggest different primary target sites for O3 and UV-B within the plant.     

Additive and antagonistic effects of ozone and salinity on the growth, ion contents and gas exchange of five varieties of rice (Oryza sativa L.)

Five varieties of rice (Oryza sativa L.) of varying salinity resistance were grown in non-saline and in saline conditions, with and without a repeated exposure to ozone at a concentration of 83 nmol mol(-1) giving an AOT40 (cumulative exposure above 40 nmol mol(-1)) of 3600 nmol mol(-1) h. Salinity caused a substantial reduction in shoot and root dry weight in all varieties, but the effect on root growth was proportionately less than on shoot growth. Ozone reduced root dry weight but the treatment used did not significantly affect shoot dry weight. Both salinity and ozone reduced plant height. The potassium concentration in the leaves of all five varieties was reduced by salinity, and by ozone in both saline and non-saline treatments. Ozone reduced the sodium concentration in plants grown at 50 mM NaCl but had no effect upon the chloride concentration. Carbon dioxide assimilation, transpiration and stomatal conductance were all reduced by salinity and by ozone and there was close quantitative similarity between the effects of ozone and/or salinity upon assimilation, stomatal conductance and transpiration. There were some antagonistic effects but there were additive effects of salinity and of ozone on root dry weight, plant height, shoot potassium concentration, photosynthesis, transpiration and stomatal conductance. The possible basis of the additive effects of salinity and ozone on gas exchange and mineral uptake are discussed.     

Growth and crown architecture of two aspen genotypes exposed to interacting ozone and carbon dioxide.

To study the impact of ozone (O3) and O3 plus CO2 on aspen growth, we planted two trembling aspen clones, differing in sensitivity to O3 in the ground in open-top chambers and exposed them to different concentrations of O3 and O3 plus CO, for 98 days. Ozone exposure (58 to 97 microl l(-1)-h. total exposure) decreased growth and modified crown architecture of both aspen clones. Ozone exposure decreased leaf, stem, branch, and root dry weight particularly in the O3 sensitive clone (clone 259). The addition of CO2 (150 microl l(-1) over ambient) to the O3 exposure counteracted the negative impact of O3 only in the O3 tolerant clone (clone 216). Ozone had relatively little effect on allometric ratios such as, shoot/root ratio, leaf weight ratio, or root weight ratio. In both clones, however, O3 decreased the shoot dry weight, shoot length ratio and shoot diameter. This decrease in wood strength caused both current terminals and long shoots to droop and increased the branch angle of termination. These results show that aspen growth is highly sensitive to O3 and that O3 can also significantly affect crown architecture. Aspen plants with drooping terminals and lateral branches would be at a competitive disadvantage in dense stands with limited light.     

Physiological effects of ozone on cultivars of watermelon (Citrullus lanatus) and muskmelon (Cucumis melo) widely grown in Spain

Two cultivars of watermelon (Citrullus lanatus) and muskmelon (Cucumis melo), which are widely grown in Spain, were exposed to ozone (70 nl litre(-1), 6 h d(-1)) for 21 days. Ozone sensitivity was assessed by recording the extent of visible injury, changes in fast-fluorescence kinetics, the relative-growth rate (R) of root (RR) and shoot (RS), and effects on the number of flowers produced per plant. Leaf gas exchange was measured in order to provide some indication of the factors underlying differential response to ozone. After 9-10 days of fumigation, all the cultivars developed typical visible symptoms of zone injury on the older leaves. However, significant (P < 0.05) changes in fast-fluorescence kinetics were detected prior to the development of visible foliar injury, indicating that detectable effects of ozone on primary photochemical processes supersede the appearance of visible symptoms. In both muskmelon and watermelon, there was a marked reduction in the rate of CO(2) assimilation as a result of exposure to ozone, and this was accompanied by a parallel decrease in stomatal conductance. Mean plant-relative-growth rate (R) was markedly (P < 0.01) suppressed by ozone in the two cultivars of watermelon, but there were no significant effects on R in muskmelon. Ozone reduced root growth relative to the shoot in three out of four cultivars-an effect that may be of considerable ecological significance. Moreover, exposure to ozone reduced flower production in both muskmelon and watermelon, which indicated effects on yield. There was no correlation between a variety of methods used to assess ozone sensitivity and visible injury, and reasons for this are discussed. This observation draws clear attention to the dangers in ranking plants for ozone sensitivity purely on the basis of visible symptoms. It is concluded from this study that ozone-insensitive genotypes should be identified and considered for planting in the major areas of melon production concentrated on the Mediterranean coast of Spain.     

Effects of pre-exposure to ultraviolet-B radiation on responses of tomato (Lycopersicon esculentum cv. New Yorker) to ozone in ambient and elevated carbon dioxide

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).     

Atmosphere and agriculture in the new millennium

Effects of changing climate (CO(2), O(3), aerosols, UV-B radiation, temperature and precipitation) on crops are predominantly based on univariate studies. Limited bivariate studies suggest rising CO(2) levels would be beneficial to crops but may be offset by adverse O(3) effects. Elevated UV-B and ambient crop yields are difficult to project due also to limited research. Climate warming concerns, using average daily temperatures may be less important than the effects of rising nocturnal temperatures on crop growth. Traditional approaches of examining air pollutant-induced visible foliar injury or the effects of single air pollutants on crop productivity need to be redirected to the analysis of integrated holistic systems. In that context, present and future agriculture in India and the USA are compared.     

Impact of fungicides on active oxygen species and antioxidant enzymes in spring barley (Hordeum vulgare L.) exposed to ozone.

Two modern fungicides, a strobilurin, azoxystrobin (AZO), and a triazole, epoxiconazole (EPO), applied as foliar spray on spring barley (Hordeum vulgare L. cv. Scarlett) 3 days prior to fumigation with injurious doses of ozone (150-250 ppb; 5 days; 7 h/day) induced a 50-60% protection against ozone injury on leaves. Fungicide treatments of barley plants at growth stage (GS) 32 significantly increased the total leaf soluble protein content. Additionally, activities of the antioxidative enzymes superoxide dismutase (SOD), catalase (CAT), ascorbate-peroxidase (APX) and glutathione reductase (GR) were increased by both fungicides at maximal rates of 16, 75, 51 and 144%, respectively. Guiacol-peroxidase (POX) activity was elevated by 50-110% only in AZO treated plants, while this effect was lacking after treatments with EPO. This coincided with elevated levels of hydrogen peroxide (H2O2) only in EPO and not in AZO treated plants. The enhancement of the plant antioxidative system by the two fungicides significantly and considerably reduced the level of superoxide (O2*-) in leaves. Fumigation of barley plants for 4 days with non-injurious ozone doses (120-150 ppb, 7 h/day) markedly and immediately stimulated O2*- accumulation in leaves, while H2O2 was increased only after the third day of fumigation. Therefore, O2*- itself or as precursor of even more toxic oxyradicals appears to be more indicative for ozone-induced leaf damage than H2O2. Ozone also induced significant increases in the activity of antioxidant enzymes (SOD, POX and CAT) after 2 days of fumigation in fungicide untreated plants, while after 4 days of fumigation these enzymes declined to a level lower than in unfumigated plants, due to the oxidative degradation of leaf proteins. This is the first report demonstrating the marked enhancement of plant antioxidative enzymes and the enhanced scavenging of potentially harmful O2*- by fungicides as a mechanism of protecting plants against noxious oxidative stress from the environment. The antioxidant effect of modern fungicides widely used in intense cereal production in many countries represents an important factor when evaluating potential air pollution effects in agriculture.     

Responses of spring wheat (Triticum aestivum L.) to ozone produced by either electric discharge and dry air or by UV-lamps and ambient air

The aim of the present study was to examine if ozone produced similar effects on spring wheat growth with and without small amounts of nitrogen oxides. Two methods were used to produce ozone: the first method consisted of dry pressurized air fed to an electric discharge generator generating the byproducts, N2O5 and N2O, the second method consisted of ambient air fed to UV-lamps. Two spring wheat cultivars (Triticum aestivum L. cvs Minaret and Eridano) were exposed in small open-top chambers to charcoal-filtered air, non-filtered ambient air, and non-filtered ambient air with the addition of ozone for 8 h (0900 to 1700 h) daily, for five weeks. Plants were harvested every week. The growth of Minaret was shown to be more sensitive to O3 than that of Eridano. Leaf senescence increased with increasing ozone level in both cultivars. The total above-ground biomass dry weight decreased with increasing ozone concentration in Minaret, but not in Eridano. The Minaret plants reacted with more damaged leaf dry weight and inhibition of growth when O3 was produced by UV-lamps than when O3 was produced by air fed to an electric discharge generator. This could be explained by more nitrogen content per plant but not by increased nitrogen concentration in plant tissue in plants exposed to increased O3 and small amounts of incidental nitrogen oxides.     

Effects of salinity and ozone,individually and in combination,on the growth and ion contents of two chickpea (Cicer arietinum L.) varieties

The aim of this study was to determine the effects of ozone and salinity, singly and in combination, on the growth and ion contents of two chickpea (Cicer arietinum L.) varieties. Chickpea plants were grown in non-saline and saline conditions, with and without a repeated exposure to ozone. Salinity at a concentration of 30 mM NaCl caused a substantial reduction in plant height, number of leaves and the dry weights of the leaves, stems and roots. Biomass allocation to the leaves increased, predominantly at the expense of the roots. Ozone at a concentration of 85 nmol mol(-1) for 6 h per day for 25 days reduced plant height and dry weights but had no effect on leaf number. The results show substantial effects of salinity and ozone on chickpea growth and ion concentrations. When ozonated plants are grown in the presence of salinity, further reductions in growth occur.     

Dose-response studies with the antiozonant ethylenediurea (EDU), applied as a soil drench to two growth substrates, on greenhouse-grown varieties of Phaseolus vulgaris L

To study plant growth and yield effects of the antiozonant ethylenediurea (EDU), which is frequently used for ozone crop loss assessments, dose-response studies were carried out with potted bean plants under greenhouse conditions in winter and spring. Two cultivars of Phaseolus vulgaris L., differing in sensitivity to ozone (O(3)), were grown in unfiltered air on a sandy loam rich in organic matter and on a vermiculite-clay mixture. Four treatments of EDU at concentrations from 300 to 800 mg liter(-1) were given as a soil drench during plant development. Foliar symptoms of EDU phytoxicity were observed at all doses, and plant biomass, particularly pod dry weight, was considerably reduced to increasing doses of EDU. Primary and first trifoliate leaf weight in EDU-treated plants increased as did the number of buds, indicating an extension of vegetative growth and a delay of reproductive processes. 'BBL 290' beans, which are O(3)-sensitive, were injured by EDU more than the O(3)-tolerant 'BBL 274'. The phytotoxic effects of EDU were more pronounced in the synthetic growth substrate than in field soil. In a second experiment, EDU was applied in concentrations from 100 to 400 mg liter(-1) to 'BBL 290' plants, exposed to filtered air or simulated levels of O(3) pollution. In field soil, plant growth and biomass partitioning in filtered air was only slightly altered by EDU, although leaf injury due to EDU occurred. In the vermiculite-clay mix, the biomass of most plant organs, particularly that of roots, was linearly reduced with increasing EDU doses. O(3) did not cause any alteration in plant biomass in field soil-grown and EDU-treated plants. Ozone leaf injury, which affected 67% of primary leaf area in non-treated plants, was completely suppressed by EDU doses as low as 100 mg liter(-1). This indicates that low concentrations of EDU, which do not affect plant growth in field soil, provide sufficient protection from O(3) injury. The need for careful EDU dose-response studies prior to field assessments is emphasized.     

Ozone episodes in Southern Lower Saxony (FRG) and their impact on the susceptibility of cereals to fungal pathogens

Spring wheat (Triticum aestivum L.) and spring barley (Hordeum vulgare L.) plants were exposed to simulated ozone (O(3)) episodes (7 h day(-1) for 7 days) at maximum concentrations of 120, 180 and 240 microg m(-3) O(3), in comparison to a charcoal-filtered air control. Fumigations were conducted in four closed chambers placed in a climate room. Exposures took place prior to inoculation of the plants with six different facultative leaf pathogens. On wheat, significant enhancement of leaf attack by Septoria nodorum Berk. and S. tritici Rob. ex Desm. appeared, particularly on the older leaves and at the highest level of O(3). The same was true for Gerlachia nivalis W. Gams et E. Müll/Fusarium culmorum (W.F.Sm.) Sacc. on wheat and net blotch (Drechslera teres (Sacc.) Shoem.) or G. nivalis leaf spots on barley. Disease development was promoted both on leaves with and without visible injury following exposure to O(3). Sporulation of the two Septoria species increased at 120 and 180 microg m(-3) O(3); however, it was reduced to the level of the control, if 240 microg m(-3) were applied. No significant effects of predisposition were observed with Bipolaris sorokiniana (Sacc.) Shoem. (syn. Helminthosporium sativum Pamm., King et Bakke), the causal agent of spot blotch, neither on wheat nor on barley. Doses and peak concentrations applied in the experiments were in good agreement with measurements of ambient ozone in Southern Lower Saxony, FRG. Six years' ozone data (1984-1989) revealed the annual occurrence of between 3 and 11 ozone episodes with potentially harmful effects on cereals (three or more consecutive 'ozone days' with 8-h means above 80 microg m(-3)). The frequency of ozone episodes followed by weather periods favourable for infections by facultative pathogens was higher in years with low O(3) pollution than in ozone-rich years, and varied between one and five cases per season. The number of ozone days during the main growing season of cereals (1 April until 31 August) varied from 25 in 1984 to 98 in 1989. However, only 7.9% of ozone days during the 6 years examined were concurrent with weather conditions suitable for fungal infections. It is concluded that the majority of leaf infections in the field happens under low-level concentrations of photooxidants.