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.     

UV-B radiation and acclimation in timberline plants

Research has shown that some plants respond to enhanced UV-B radiation by producing smaller and thicker leaves, by increasing the thickness of epidermis and concentration of UV-B absorbing compounds of their surface layers and activation of the antioxidant defence system. The response of high-altitude plants to UV-B radiation in controlled conditions is often less pronounced compared to low-altitude plants, which shows that the alpine timberline plants are adapted to UV-B. These plants may have a simultaneous co-tolerance for several stress factors: acclimation or adaptation to the harsh climate can also increase tolerance to UV-B radiation, and vice versa. On the other hand, alpine timberline plants of northern latitudes may be less protected against increasing UV-B radiation than plants from more southern latitudes and higher elevations due to harsh conditions and weaker preadaptation resulting from lower UV-B radiation exposure. It is evident that more long-term experimental field research is needed in order to study the interaction of climate, soil and UV-B irradiance on the timberline plants.     

Ground-based measurements and modeling of solar UV-B radiation in Lhasa,Tibet

We present results from measurements by a NILU-UV Irradiance Meter of solar UV-B radiation in Lhasa, Tibet for the period from 7 July 1999 to 17 November 2002. The measured maximum UV-B dose rate was 1.50 W/m² during the winter, and 3.96 W/m², during the summer. The clear-sky values were below 3.00 W/m². Comparisons between measured and calculated UV-B dose rates on clear-sky days in Lhasa show good agreement. Comparisons of UV-B radiation levels in Lhasa (Tibet), Oslo (Norway), and Dar-Es-Salaam (Tanzania) show that the UV-B dose rates during the summer in Lhasa are higher than the maximum value in Dar-Es-Salaam, which is at the sea level in the equatorial region, and 60% higher than in Oslo, which is at the sea level but 60 degrees North. We conclude that the UV-B dose rates during the summer on the Tibetan plateau are among the highest levels in habituated regions of the world. Maximum measured daily-integrated UV-B doses in Lhasa range from about 10 kJ/m² in the winter to about 65 kJ/m² in the summer.     

Growth-stage dependent crop yield response to ozone exposure

Data from four crop yield-loss field trials were examined to determine if analysis using an imposed phenological weighting function based on seasonal growth stage would provide a more accurate indication of impact of ozone exposure. Alfalfa (Medicago sativa L. cv. Moapa 69), dry bean (Phaseolus vulgaris L. cv. California Dark Red kidney), fresh market and processing tomato (Lycopersicon esculentum Mill. cv. 6718 VF and VF-145-B7879, respectively) were grown at 9-11 ambient field plots within southern California comprising an ambient gradient of ozone. The growing season for each crop was artificially divided into 'quarters' composed of equal numbers of whole days and roughly corresponding to specific growth stages. Ozone exposure was calculated for each of these 'quarters' and regressed against final crop yield using 163 different exposure statistics. Weighting functions were developed using reciprocal residual mean square (1/RMS) or percentage of the best 100 exposure statistics of the 163 tested (TOP100) for each of the quarters. The third quarter of the alfalfa season was clearly most responsive to ozone as measured by both of the weighting functions. Third quarter ozone was also weighted highest by both weighting functions for dry bean. Fresh market and processing tomato were each influenced the greatest by second quartero zone as demonstrated by both weighting functions. The occurrence of ozone during physiologically important events (flowering and initial fruit set in second quarter for tomato; pod development in third quarter for dry bean) appeared to influence the yield of these crops the greatest. Growth-stage-dependent phenological weighting of pollutant exposure may result in more effective predictions of levels of ozone exposure resulting in yield reductions.     

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.     

Ambient ozone and crop loss: establishing a cause-effect relationship

This paper provides the results of a retrospective mathematical analysis of the US NCLAN (National Crop Loss Assessment Network) open-top chamber data. Some 77% of the 73 crop harvests examined, showed no statistically significant yield differences between NF (non-filtered open-top chamber) and AA (chamberless, ambient air) treatments (no easily discernable chamber effects on yield). However, among these cases only seven acceptable examples showed statistically significant yield reductions in NF compared to the CF (charcoal filtered open-top chamber) treatment. An examination of the combined or cumulative hourly ambient O3 frequency distribution for cases with yield loss in NF compared to a similar match of cases without yield loss showed that the mean, median and the various percentiles were all higher (>/= 3 X) in the former in contrast to the latter scenario. The combined frequency distribution of hourly O3 concentrations for the cases with yield loss in NF were clearly separated from the corresponding distribution with no yield loss, at O3 concentrations > 49 ppb. Univariate linear regressions between various O3 exposure parameters and per cent yield losses in NF showed that the cumulative frequency of occurrence of O3 concentrations between 50 and 87 ppb was the best predictor (adjusted R2 = 0.712 and p = 0.011). This analysis also showed that the frequency distribution of hourly concentrations up to 87 ppb O3 represented a critical point, since the addition of the frequency distributions of > 87 ppb O3 did not improve the R2 values. In fact as the frequency of hourly O3 concentrations included in the regression approached 50-100 ppb, the R2 value decreased substantially and the p value increased inversely. Further, univariate linear regressions between the frequencies of occurrence of various O3 concentrations between 50 and 90 ppb and: (a) cases with no yield difference in NF and (b) cases with yield increase in NF compared to the CF treatment (positive effect) provided no meaningful statistical relationship (adjusted R2 = 0.000) in either category. These results support the basis that additional evaluation of the frequency of occurrence of hourly O3] concentrations between 50 and 87 ppb for cases with the yield reductions could provide a meaningful ambient O3 standard, objective or guideline for vegetation.     

Nocturnal depletion of photochemical ozone at a rural site

Nocturnal depletion of ozone during a period of photochemical pollution has been observed at a rural site by continuous ground-level measurements and the determination of vertical profiles of ozone concentration. An acoustic sounder was used to provide continuous information upon the vertical temperature structure of the atmosphere. The results are interpreted in terms of recent theories of long-range transport of photochemical ozone and the chemical and physical processes causing ozone depletion.     

Ground-level ozone in China: distribution and effects on crop yields

Rapid economic development and an increasing demand for food in China have drawn attention to the role of ozone at pollution levels on crop yields. Some assessments of ozone effects on crop yields have been carried out in China. Determination of ozone distribution by geographical location and resulting crop loss estimations have been made by Chinese investigators and others from abroad. It is evident that surface level ozone levels in China exceed critical levels for occurrence of crop losses. Current levels of information from ozone dose/response studies are limited. Given the size of China, existing ozone monitoring sites are too few to provide enough data to scale ozone distribution to a national level. There are large uncertainties in the database for ozone effects on crop loss and for ozone distribution. Considerable research needs to be done to allow accurate estimation of crop losses caused by ozone in China.     

Evidence of ozone injury to a crop plant in India

Evidence is presented to show that a serious leaf spot disease of potato which appeared each year in the Punjab since 1978 is primarily due to ozone:

• 1.(i) The symptoms of the leaf spot were similar to the ozone stipple of potato reported in the U.S.A.
• 2.(ii) Activated charcoal and ethylenediurea effectively controlled the spots.
• 3.(iii) Elevated ozone in the atmosphere was detected with the bioindicators Nicotiana tabacum var. Bel-W3 and potato variety Cherokee. This is the first report of ozone injury to a crop plant in India.

     

Tethered balloon-based soundings of ozone,aerosols, and solar radiation near Mexico City during MIRAGE-MEX

A tethered balloon sampling system was used to measure vertical profiles of ozone, particles, and solar radiation in the atmospheric boundary layer on the northern edge of Mexico City, in March 2006 as part of the Megacity Impact on Regional and Global Environment-Mexico experiment. Several commercial sensors, designed for surface applications, were deployed on a tethered balloon platform.Profiles indicate that for these 3 scalars the boundary layer (surface up to 700 m) was well mixed in the period 10:00–16:00 LST. Good agreement was observed for median surface and balloon ozone and particle number concentrations. For most profiles, the surface deposition of ozone was not significant compared to median profile concentrations. Particle number concentration (0.3, 0.5, 1.0 and 5.0 μm) also showed little variation with attitude. Radiatprofiles showed a monotonic increase in diffuse radiation from the maximum altitude of profiles to the surface. Consequently, it was inferred that surface measurements of these likely were representative of lower boundary layer values during this time period.     

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.     

Climate change: potential effects of increased atmospheric carbon dioxide (CO2), ozone (O3), and ultraviolet-B (UV-B) radiation on plant diseases

Continued world population growth results in increased emission of gases from agriculture, combustion of fossil fuels, and industrial processes. This causes changes in the chemical composition of the atmosphere. Evidence is emerging that increased solar ultraviolet-B (UV-B) radiation is reaching the earth's atmosphere, due to stratospheric ozone depletion. Carbon dioxide (CO(2)), ozone (O(3)) and UV-B are individual climate change factors that have direct biological effects on plants. Such effects may directly or indirectly affect the incidence and severity of plant diseases, caused by biotic agents. Carbon dioxide may increase plant canopy size and density, resulting in a greater biomass of high nutritional quality, combined with a much higher microclimate relative humidity. This would be likely to promote plant diseases such as rusts, powdery mildews, leaf spots and blights. Inoculum potential from greater overwintering crop debris would also be increased. Ozone is likely to have adverse effects on plant growth. Necrotrophic pathogens may colonize plants weakened by O(3) at an accelerated rate, while obligate biotroph infections may be lessened. Ozone is unlikely to have direct adverse effects on fungal pathogens. Ozone effects on plant diseases are host plant mediated. The principal effects of increased UV-B on plant diseases would be via alterations in host plants. Increased flavonoids could lead to increased diseased resistance. Reduced net photosynthesis and premature ripening and senescence could result in a decrease in diseases caused by biotrophs and an increase in those caused by necrotrophs. Microbial plant pathogens are less likely to be adversely affected by CO(2), O(3) and UV-B than are their corresponding host plants. Changes in host plants may result in expectable alterations of disease incidence, depending on host plant growth stages and type of pathogen. Given the importance of plant diseases in world food and fiber production, it is essential to begin studying the effects of increased CO(2), O(3) and UV-B (and other climate change factors) on plant diseases. We know very little about the actual impacts of climate change factors on disease epidemiology. Epidemiologists should be encouraged to consider CO(2), O(3) and UV-B as factors in their field studies.     

Photoinduced toxicity of retene to Daphnia magna under enhanced UV-B radiation

The effects of UV radiation on the acute toxicity of retene (7-isopropyl-1-methylphenanthrene) to Daphnia magna Straus were studied. Dehydroabietic acid (DHAA) from which retene is formed in the vicinity of pulp and paper industry was also studied. Pyrene, anthracene, and phenanthrene were used as model PAH compounds. The time taken for immobilization (ET50) was monitored under biologically effective UV-B dose rates of 240, 365, 565, and 650 mW m(-2) (UV-A and visible light also present). Median effective concentrations (EC50) were determined after a 15-min UV exposure (565 mW m(-2)) followed by 24 h in the dark. Retene (10-320 microg l(-1)) was not acutely toxic in the dark. The induction of phototoxicity was in agreement with the absorption properties of the compounds (absorption peak of retene at around 300 nm). Photoinduced toxicity followed an order pyrene > anthracene > retene. Phenanthrene and DHAA were not acutely phototoxic. Accumulation of the compound in Daphnia before UV exposure was essential. Some changes in the absorption spectra of the compounds were seen after a 5-h UV irradiation (565 mW m(-2)), but none of the irradiated compounds were acutely toxic without further UV exposure. Therefore, the enhanced acute toxicity was primarily due to internal photosensitization reactions rather than photomodification. The dissolved fraction of 25% pulp and paper mill effluent reduced phototoxicity by attenuating UV radiation. The phototoxicity of retene was a function of both the exposure concentration and the UV-B dose rate, but relatively high UV-B dose rates and concentrations were needed for the acute photoinduced toxicity.     

Photo-oxidant chemistry in the polluted boundary layer under changing UV-B radiation

UV-B radiation is a driving factor for the chemistry of the polluted boundary layer. It is involved in the formation of radicals and consequently influences the formation and concentration of photo-oxidants. The 3-D mesoscale photochemical Metphomod model was employed to study the effect of changes in UV-B radiation on the concentration of photo-oxidants in the boundary layer over the Swiss Plateau. The model chemistry is based on the RACM mechanism and a two-stream approximation of radiative transfer. A summer (July) and a late winter (February) episode were simulated. All simulations were replicated with relatively large changes in the prescribed total ozone. The results for an increase in UV-B radiation show increases in PAN, HNO₃, and ozone at noon in NO_x-rich areas and a decrease in NO_x. In NO_x-poor areas in summer the effect on ozone is weak and has a negative sign, the main effect being an increase in H₂O₂. The spatial variability of NO_x concentrations in the Swiss Plateau in the summer case is such that the effect of increased UV-B radiation on ozone is spatially variable. The effect on the ozone production rate in summer is strongest positive at the surface in the NO_x-rich regions in the morning and strongest negative at some altitude above ground in NO_x-poor regions in the early afternoon. In the winter episode, NO_x-rich conditions are found almost everywhere on the Swiss Plateau, the effect of increased UV-B radiation on the ozone production rate is positive all day long and is largest at 300 m above ground at noon. In this case, in contrast to the summer case, the increase in ozone is carried over to the next day. The model results for ozone are in good agreement with results from a case study and a time series analysis of surface ozone measurements. We estimate the effect of day-to-day changes in total ozone on surface ozone peaks to range from 4 to 6 ppb at most.     

Derivation of ozone flux-yield relationships for lettuce: a key horticultural crop

Ozone flux-response relationships were derived for lettuce, employing a multiplicative approach to model the manner in which stomatal conductance is influenced by key environmental variables, using a dataset collected during field experimentation in Crete and yield-response relationships derived from parallel open-top chamber experiments. Regional agronomic practices were adopted throughout. Computed versus measured data revealed that the derived model explained 51% (P<0.001) of the observed variation in stomatal conductance. Concentration-based indices were compared with flux-based indices. Analyses revealed a significant relationship between accumulated stomatal ozone flux and yield employing flux threshold cut-offs up to 4 nmol m(-2) s(-1). Regressions employing very low or zero flux thresholds resulted in the strongest yield-flux relationships (explaining approximately 80% (P<0.05) of the variation in the dataset).     

Modeling the impact of ozone x drought interactions on regional crop yields

The influence of soil moisture stress on crop sensitivity to O3 was evaluated for corn (Zea mays L.), cotton (Gossypium hirsutum L.), soybean (Glycine max L. Merr.), and wheat (Triticum aestivum L.) grown in the United States. This assessment was accomplished by using yield forecasting models to estimate the influence of soil moisture deficits on regional yield and a previously developed model to predict moisture stress x O3 interactions. Reduced crop sensitivity to O3 was predicted for those regions and years for which soil moisture stress reduced yield. The models predicted a drought-induced reduction in crop sensitivity to O3 of approximately 20% for the 1979 to 1983 period; i.e. a hypothetical O3-induced yield reduction of 5% for adequately watered crops would have been reduced to a 4% effect by the 1979 to 1983 distribution of soil moisture deficits. However, predicted drought effects varied between crops, regions, and years. Uncertainties in the model predictions are also discussed.     

Selecting ozone exposure statistics for determining crop yield loss from air pollutants

Numerous ozone exposure statistics were calculated using hourly ozone data from crop yield loss experiments previously conducted for alfalfa, fresh market and processing tomatoes, cotton, and dry beans in an ambient ozone gradient near Los Angeles, California. Exposure statistics examined included peak (maximum daily hourly) and mean concentrations above specific threshold levels, and concentrations during specific time periods of the day. Peak and mean statistics weighted for ozone concentration and time period statistics weighted for hour of the day were also determined. Polynomial regression analysis was used to relate each of 163 ozone statistics to crop yield. Performance of the various statistics was rated by comparing residual mean square (RMS) values. The analyses demonstrated that no single statistic was best for all crop species. Ozone statistics with a threshold level performed well for most crops, but optimum threshold level was dependent upon crop species and varied with the particular statistics calculated. The data indicated that daily hours of exposure above a critical high-concentration threshold related well to crop yield for alfalfa, market tomatoes, and dry beans. The best statistic for cotton yield was an average of all daily peak ozone concentrations. Several different types of ozone statistics performed similarly for processing tomatoes. These analyses suggest that several ozone summary statistics should be examined in assessing the relationship of ambient ozone exposure to crop yield. Where no clear statistical preference is indicated among several statistics, those most biologically relevant should be selected.