Ozone Inhibition of Photosynthesis in Chlorella sorokiniana

Exposure of Chlorella sorokiniana (07-11-05) to ozone inhibits photosynthesis. In this study, the effects of ozone on O₂ evolution and fluorescence yields are used to characterize this inhibition. At an ozone dose of about 3 micromoles delivered to 2 × 10⁹ cells, the photosynthetic rate of the cells is inhibited 50%, as indicated by a decrease in bicarbonate-stimulated O₂ evolution (control rate, 1.4 ± 0.3 × 10⁻¹⁵ moles per cell per minute).     

The influence of rising tropospheric carbon dioxide and ozone on plant productivity

Human activities result in a wide array of pollutants being released to the atmosphere. A number of these pollutants have direct effects on plants, including carbon dioxide (CO₂), which is the substrate for photosynthesis, and ozone (O₃), a damaging oxidant. How plants respond to changes in these atmospheric air pollutants, both directly and indirectly, feeds back on atmospheric composition and climate, global net primary productivity and ecosystem service provisioning. Here we discuss the past, current and future trends in emissions of CO₂ and O₃ and synthesise the current atmospheric CO₂ and O₃ budgets, describing the important role of vegetation in determining the atmospheric burden of those pollutants. While increased atmospheric CO₂ concentration over the past 150 years has been accompanied by greater CO₂ assimilation and storage in terrestrial ecosystems, there is evidence that rising temperatures and increased drought stress may limit the ability of future terrestrial ecosystems to buffer against atmospheric emissions. Long‐term Free Air CO₂ or O₃ Enrichment (FACE) experiments provide critical experimentation about the effects of future CO₂ and O₃ on ecosystems, and highlight the important interactive effects of temperature, nutrients and water supply in determining ecosystem responses to air pollution. Long‐term experimentation in both natural and cropping systems is needed to provide critical empirical data for modelling the effects of air pollutants on plant productivity in the decades to come.     

Ozone,ultraviolet flux and temperature of the paleoatmosphere

Of all tropospheric species, ozone (O₃) comes closest to being naturally present at toxic levels. In addition, O₃ controls the ultraviolet flux reaching the Earth's surface and affects the temperature of the surface and atmosphere. For these reasons, O₃ was an important species of the paleoatmosphere. Surface and atmospheric levels of paleoatmospheric O₃ were calculated using a detailed photochemical model, including the chemistry of the oxygen, nitrogen, and hydrogen species and the effects of vertical transport. Surface and tropospheric O₃, as well as the total O₃ column, were found to maximize for an atmospheric oxygen level of 10^–1 present atmospheric level (PAL). Coupled photochemical/radiative-convective calculations indicate that the radiative effects of O₃ corresponding to an oxygen level of 10^–1 PAL resulted in a globally-averaged surface temperature increase of 4.5 K.Proceedings of the Fourth College Park Colloquium on Chemical Evolution:Limits of Life, University of Maryland, College Park, 18–20 October 1978.     

Long‐term exposure to elevated CO2 and O3 alters aspen foliar chemistry across developmental stages

Anthropogenic activities are altering levels of greenhouse gases to the extent that multiple and diverse ecosystem processes are being affected. Two gases that substantially influence forest health are atmospheric carbon dioxide (CO₂) and tropospheric ozone (O₃). Plant chemistry will play an important role in regulating ecosystem processes in future environments, but little information exists about the longitudinal effects of elevated CO₂ and O₃ on phytochemistry, especially for long‐lived species such as trees. To address this need, we analysed foliar chemical data from two genotypes of trembling aspen, Populus tremuloides, collected over 10 years of exposure to levels of CO₂ and O₃ predicted for the year 2050. Elevated CO₂ and O₃ altered both primary and secondary chemistry, and the magnitude and direction of the responses varied across developmental stages and between aspen genotypes. Our findings suggest that the effects of CO₂ and O₃ on phytochemical traits that influence forest processes will vary over tree developmental stages, highlighting the need to continue long‐term, experimental atmospheric change research.     

Soil carbon dioxide partial pressure and dissolved inorganic carbonate chemistry under elevated carbon dioxide and ozone

Global emissions of atmospheric CO₂ and tropospheric O₃ are rising and expected to impact large areas of the Earths forests. While CO₂ stimulates net primary production, O₃ reduces photosynthesis, altering plant C allocation and reducing ecosystem C storage. The effects of multiple air pollutants can alter belowground C allocation, leading to changes in the partial pressure of CO₂ (pCO₂) in the soil , chemistry of dissolved inorganic carbonate (DIC) and the rate of mineral weathering. As this system represents a linkage between the long- and short-term C cycles and sequestration of atmospheric CO₂, changes in atmospheric chemistry that affect net primary production may alter the fate of C in these ecosystems. To date, little is known about the combined effects of elevated CO₂ and O₃ on the inorganic C cycle in forest systems. Free air CO₂ and O₃ enrichment (FACE) technology was used at the Aspen FACE project in Rhinelander, Wisconsin to understand how elevated atmospheric CO₂ and O₃ interact to alter pCO₂ and DIC concentrations in the soil. Ambient and elevated CO₂ levels were 360±16 and 542±81 l l^–1, respectively; ambient and elevated O₃ levels were 33±14 and 49±24 nl l^–1, respectively. Measured concentrations of soil CO₂ and calculated concentrations of DIC increased over the growing season by 14 and 22%, respectively, under elevated atmospheric CO₂ and were unaffected by elevated tropospheric O₃. The increased concentration of DIC altered inorganic carbonate chemistry by increasing system total alkalinity by 210%, likely due to enhanced chemical weathering. The study also demonstrated the close coupling between the seasonal ¹³C of soil pCO₂ and DIC, as a mixing model showed that new atmospheric CO₂ accounted for approximately 90% of the C leaving the system as DIC. This study illustrates the potential of using stable isotopic techniques and FACE technology to examine long- and short-term ecosystem C sequestration.     

Long-Term Leaf Production Response to Elevated Atmospheric Carbon Dioxide and Tropospheric Ozone

Elevated concentrations of atmospheric CO₂ and tropospheric O₃ will profoundly influence future forest productivity, but our understanding of these influences over the long-term is poor. Leaves are key indicators of productivity and we measured the mass, area, and nitrogen concentration of leaves collected in litter traps from 2002 to 2008 in three young northern temperate forest communities exposed to elevated CO₂ and/or elevated O₃ since 1998. On average, the overall effect of elevated CO₂ (+CO₂ and +CO₂+O₃ versus ambient and +O₃) was to increase leaf mass by 36% whereas the overall effect of elevated O₃ was to decrease leaf mass by 13%, with similar effects on stand leaf area. However, there were important CO₂ × O₃ × year interactions wherein some treatment effects on leaf mass changed dramatically relative to ambient from 2002 to 2008. For example, stimulation by the +CO₂ treatment decreased (from +52 to +25%), whereas the deleterious effects of the +O₃ treatment increased (from −5 to −18%). In comparison, leaf mass in the +CO₂+O₃ treatment was similar to ambient throughout the study. Forest composition influenced these responses: effects of the +O₃ treatment on community-level leaf mass ranged from +2 to −19%. These findings are evidence that community composition, stand development processes, CO₂, and O₃ strongly interact. Changes in leaf nitrogen concentration were inconsistent, but leaf nitrogen mass (g m⁻²) was increased by elevated CO₂ (+30%) and reduced by elevated O₃ (−16%), consistent with observations that nitrogen cycling is accelerated by elevated CO₂ but retarded by elevated O₃.     

Ozone Induced Carbon Dioxide Evolution in Tobacco Callus Cultures

Callus derived from Bel–W3 and Bel–B tobacco plants when exposed to ozone turned brown as a consequence of surface cell destruction. Ozone fumigations above a threshold concentration of 0.10 μl/1 for two hoars caused an increase in the rate of tissue carbon dioxide (CO₂) evolution. The maximum increase in CO₂ evolution was about 65 percent for both the ozone sensitive Bel–W3 and resistant Bel–B callus. However, the ozone dosage required to attain maximum increase in CO₂ evolution was approximately two times greater for the resistant variety. Callus cultures that grew roots were observed to be more resistant to ozone. The addition of the antioxidant N,N'dipnenyl–p–phenylenediamine (DPPD) m the nutrient medium retarded ozone induced CO₂ evolution.     

THE EFFECTS OF OXYGEN,CARBON DIOXIDE,AND PRESSURE ON GROWTH IN CHILOMONAS PARAMECIUM AND TETRAHYMENA GELEII FURGASON

1. The effects of O₂, CO₂, and pressure were studied in two very different species of protozoa, a flagellate, Chilomonas paramecium, grown in acetate-ammonium solution and a ciliate, Tetrahymena geleii, grown in 2 per cent proteose-peptone solution. 2. Chilomonas and Tetrahymena live and reproduce in solutions exposed to a wide range of O₂ concentrations, but Chilomonas is killed at high O₂ tensions in which Tetrahymena grows best. The optimum O₂ concentration for Chilomonas is about 75 mm. pressure but it lives and reproduces in O₂ tensions as low as 0.5 mm. while Tetrahymena fails to grow in concentrations below 10 mm. O₂ pressure. 3. With a constant O₂ tension of 50 mm. pressure, it was found that there is no significant variation in growth in Chilomonas between 50 mm. and 740 mm. total pressure. In Tetrahymena, however, under the same conditions, an optimum total pressure was found at about 500 mm. and growth is comparatively poor at 50 mm. total pressure. 4. Tetrahymena does not live very long in CO₂ tensions over 122 mm., although Chilomonas grows as well at 400 mm. CO₂ as in air at atmospheric pressure (0.2 mm. CO₂). Tetrahymena grows best in an environment minus CO₂, but the optimum for Chilomonas is 100 mm. CO₂ at which pressure an average of 668,600 ± 30,000 organisms per ml. was produced (temperature, 25 ± 1° C.). 5. Chilomonads grown in high CO₂ concentrations (e.g., 122 mm.) produce larger starch granules and more starch than those grown in ordinary air at atmospheric pressure. 6. In solutions exposed to 75 mm. O₂ tension (optimum) and 122 mm. CO₂ plus 540 mm. N₂ pressure, chilomonads contain very little, if any, fat. This phenomenon seems to be due to the action of CO₂ on the mechanisms concerned with fat production. 7. In Tetrahymena exposed to pure O₂, there is very little fat compared to those grown in atmospheric air. This may be due to the greater oxidation of fat in the higher O₂ concentrations. 8. Further evidence is presented in support of the contention that Chilomonas utilizes CO₂ in the production of starch.     

The Regulation of Photosynthesis in Leaves of Field-Grown Spring Wheat (Triticum aestivum L., cv Albis) at Different Levels of Ozone in Ambient Air

Wheat (Triticum aestivum L. cv Albis) was grown in open-top chambers in the field and fumigated daily with charcoal-filtered air (0.015 microliters per liter O₃), nonfiltered air (0.03 microliters per liter O₃), and air enriched with either 0.07 or 0.10 microliters per liter ozone (seasonal 8 hour/day [9 am-5 pm] mean ozone concentration from June 1 until July 10, 1987). Photosynthetic ¹⁴CO₂ uptake was measured in situ. Net photosynthesis, dark respiration, and CO₂ compensation concentration at 2 and 21% O₂ were measured in the laboratory. Leaf segments were freeze-clamped in situ for the determination of the steady state levels of ribulose 1,5-bisphosphate, 3-phosphoglycerate, triose-phosphate, ATP, ADP, AMP, and activity of ribulose, 1,5-bisphosphate carboxylase/oxygenase. Photosynthesis of flag leaves was highest in filtered air and decreased in response to increasing mean ozone concentration. CO₂ compensation concentration and the ratio of dark respiration to net photosynthesis increased with ozone concentration. The decrease in photosynthesis was associated with a decrease in chlorophyll, soluble protein, ribulose bisphosphate carboxylase/oxygenase activity, ribulose bisphosphate, and adenylates. No decrease was found for triose-phosphate and 3-phosphoglycerate. The ratio of ATP to ADP and of triosephosphate to 3-phosphoglycerate were increased suggesting that photosynthesis was limited by pentose phosphate reductive cycle activity. No limitation occurred due to decreased access of CO₂ to photosynthetic cells since the decrease in stomatal conductance with increasing ozone concentration did not account for the decrease in photosynthesis. Ozonestressed leaves showed an increased degree of activation of ribulose bisphosphate carboxylase/oxygenase and a decreased ratio of ribulose bisphosphate to initial activity of ribulose bisphosphate carboxylase/oxygenase. Nevertheless, it is suggested that photosynthesis in ozone stressed leaves is limited by ribulose bisphosphate carboxylation possibly due to an effect of ozone on the catalysis by ribulose bisphosphate carboxylase/oxygenase.     

Effects of Ambient and Acute Partial Pressures of Ozone on Leaf Net CO(2) Assimilation of Field-Grown Vitis vinifera L

Mature, field-grown Vitis vinifera L. grapevines grown in open-top chambers were exposed to either charcoal-filtered air or ambient ozone partial pressures throughout the growing season. Individual leaves also were exposed to ozone partial pressures of 0.2, 0.4, or 0.6 micropascals per pascal for 5 hours. No visual ozone damage was found on leaves exposed to any of the treatments. Chronic exposure to ambient O₃ partial pressures reduced net CO₂ assimilation rate (A) between 5 and 13% at various times throughout the season when compared to the filtered treatment. Exposure of leaves to 0.2 micropascals per pascal O₃ for 5 hours had no significant effect on A; however, A was reduced 84% for leaves exposed to 0.6 micropascals per pascal O₃ when compared to the controls after 5 hours. Intercellular CO₂ partial pressure (c_i) was lower for leaves exposed to 0.2 micropascals per pascal O₃ when compared to the controls, while c_i of the leaves treated with 0.6 micropascals per pascal of 0₃ increased during the fumigation. The long-term effects of ambient O₃ and short-term exposure to acute levels of O₃ reduced grape leaf photosynthesis due to a reduction in both stomatal and mesophyll conductances.     

Interannual climatic variation mediates elevated CO2 and O3 effects on forest growth

We analyzed growth data from model aspen (Populus tremuloides Michx.) forest ecosystems grown in elevated atmospheric carbon dioxide ([CO₂]; 518 μL L^?1) and ozone concentrations ([O₃]; 1.5 × background of 30–40 nL L^?1 during daylight hours) for 7 years using free‐air CO₂ enrichment technology to determine how interannual variability in present‐day climate might affect growth responses to either gas. We also tested whether growth effects of those gasses were sustained over time. Elevated [CO₂] increased tree heights, diameters, and main stem volumes by 11%, 16%, and 20%, respectively, whereas elevated ozone [O₃] decreased them by 11%, 8%, and 29%, respectively. Responses similar to these were found for stand volume and basal area. There were no growth responses to the combination of elevated [CO₂+O₃]. The elevated [CO₂] growth stimulation was found to be decreasing, but relative growth rates varied considerably from year to year. Neither the variation in annual relative growth rates nor the apparent decline in CO₂ growth response could be explained in terms of nitrogen or water limitations. Instead, growth responses to elevated [CO₂] and [O₃] interacted strongly with present‐day interannual variability in climatic conditions. The amount of photosynthetically active radiation and temperature during specific times of the year coinciding with growth phenology explained 20–63% of the annual variation in growth response to elevated [CO₂] and [O₃]. Years with higher photosynthetic photon flux (PPF) during the month of July resulted in more positive growth responses to elevated [CO₂] and more negative growth responses to elevated [O₃]. Mean daily temperatures during the month of October affected growth in a similar fashion the following year. These results indicate that a several‐year trend of increasingly cloudy summers and cool autumns were responsible for the decrease in CO₂ growth response.     

Ecophysiological responses of the larch species in northern Japan to environmental changes as a basis for afforestation

For sustainable use and suitable management of larch plantations, we must clarify the ecophysiological responses of larch species to environmental changes. The physical environment has been changing dramatically, e.g., increase in atmospheric CO₂ concentration ([CO₂]), nitrogen (N) deposition, and atmospheric ozone concentration ([O₃]), and these changes may negatively affect growth of larch species. This review summarizes the previous experimental studies on the ecophysiological responses of larch species to elevated [CO₂], soil acidification, elevated [O₃], and N load. Based on the advanced studies, although elevated [CO₂] will stimulate the productivity of larch, increase of [O₃] and severe soil acidification will reduce it. Increase of N deposition, at least, will not negatively affect larch productivity. Finally, we propose the future direction for investigation to understand the mechanism of the responses of larch species and to predict the associated risk.     

Influence of Global Atmospheric Change on the Feeding Behavior and Growth Performance of a Mammalian Herbivore,Microtus ochrogaster

Global atmospheric change is influencing the quality of plants as a resource for herbivores. We investigated the impacts of elevated carbon dioxide (CO₂) and ozone (O₃) on the phytochemistry of two forbs, Solidago canadensis and Taraxacum officinale, and the subsequent feeding behavior and growth performance of weanling prairie voles (Microtus ochrogaster) feeding on those plants. Plants for the chemical analyses and feeding trials were harvested from the understory of control (ambient air), elevated CO₂ (560 µl CO₂ l⁻¹), and elevated O₃ (ambient × 1.5) rings at the Aspen FACE (Free Air CO₂ Enrichment) site near Rhinelander, Wisconsin. We assigned individual voles to receive plants from only one FACE ring and recorded plant consumption and weanling body mass for seven days. Elevated CO₂ and O₃ altered the foliar chemistry of both forbs, but only female weanling voles on the O₃ diet showed negative responses to these changes. Elevated CO₂ increased the fiber fractions of both plant species, whereas O₃ fumigation elicited strong responses among many phytochemical components, most notably increasing the carbon-to-nitrogen ratio by 40% and decreasing N by 26%. Consumption did not differ between plant species or among fumigation treatments. Male voles were unaffected by the fumigation treatments, whereas female voles grew 36% less than controls when fed O₃-grown plants. These results demonstrate that global atmospheric change has the potential to affect the performance of a mammalian herbivore through changes in plant chemistry.     

Century-Scale Responses of Ecosystem Carbon Storage and Flux to Multiple Environmental Changes in the Southern United States

Terrestrial ecosystems in the southern United States (SUS) have experienced a complex set of changes in climate, atmospheric CO₂ concentration, tropospheric ozone (O₃), nitrogen (N) deposition, and land-use and land-cover change (LULCC) during the past century. Although each of these factors has received attention for its alterations on ecosystem carbon (C) dynamics, their combined effects and relative contributions are still not well understood. By using the Dynamic Land Ecosystem Model (DLEM) in combination with spatially explicit, long-term historical data series on multiple environmental factors, we examined the century-scale responses of ecosystem C storage and flux to multiple environmental changes in the SUS. The results indicated that multiple environmental changes shifted SUS ecosystems from a C source of 1.20?±?0.56?Pg (1?Pg?=?10¹⁵?g) during the period 1895 to 1950, to a C sink of 2.00?±?0.94?Pg during the period 1951 to 2007. Over the entire period spanning 1895–2007, SUS ecosystems were a net C sink of 0.80?±?0.38?Pg. The C sink was primarily due to an increase in the vegetation C pool, whereas the soil C pool decreased during the study period. The spatiotemporal changes of C storage were caused by changes in multiple environmental factors. Among the five factors examined (climate, LULCC, N deposition, atmospheric CO₂, and tropospheric O₃), elevated atmospheric CO₂ concentration was the largest contributor to C sequestration, followed by N deposition. LULCC, climate, and tropospheric O₃ concentration contributed to C losses during the study period. The SUS ecosystem C sink was largely the result of interactive effects among multiple environmental factors, particularly atmospheric N input and atmospheric CO_2.     

Growth and photosynthetic response of <Emphasis Type="Italic">Fagus crenata</Emphasis> seedlings to ozone and/or elevated carbon dioxide

We investigated the effects of ozone (O₃) and/or elevated CO₂ concentration ([CO₂]) on growth and photosynthetic traits of Fagus crenata seedlings. Two-year-old seedlings were grown in four experimental treatments comprising two O₃ treatments (charcoal-filtered air and 100 nmol mol⁻¹ O₃; 6 h/day, 3 days/week) in combination with two CO₂ treatments (350 and 700 μmol mol⁻¹) for 18 weeks in environmental control growth chambers. The four treatments were designated as control, elevated O₃, elevated CO₂, and elevated CO₂ + O₃. Dry matter growth of the seedlings was greater in elevated CO₂ + O₃ than in elevated CO₂. In elevated CO₂ + O₃, a marked increase of second-flush leaves, considered a compensative response to O₃, was observed. The net photosynthetic rate of first-flush leaves in elevated CO₂ + O₃ increased earlier and was maintained for a longer period of time than that in elevated CO₂. Because emergence of second-flush leaves of F. crenata is greatly affected by the amount of assimilation products of first-flush leaves in current year, we consider that an early increase in the net photosynthetic rate of first-flush leaves contributed to the marked increase in second-flush leaf emergence under elevated CO₂ + O₃. These results imply that we must account for changes in compensative capacity with respect to not only morphological traits but also phenological traits and physiological functions such as photosynthesis when evaluating effects of O₃ on F. crenata under elevated [CO₂].     

Distinct responses of soil microbial communities to elevated CO2 and O3 in a soybean agro-ecosystem

The concentrations of atmospheric carbon dioxide (CO₂) and tropospheric ozone (O₃) have been rising due to human activities. However, little is known about how such increases influence soil microbial communities. We hypothesized that elevated CO₂ (eCO₂) and elevated O₃ (eO₃) would significantly affect the functional composition, structure and metabolic potential of soil microbial communities, and that various functional groups would respond to such atmospheric changes differentially. To test these hypotheses, we analyzed 96 soil samples from a soybean free-air CO₂ enrichment (SoyFACE) experimental site using a comprehensive functional gene microarray (GeoChip 3.0). The results showed the overall functional composition and structure of soil microbial communities shifted under eCO₂, eO₃ or eCO₂+eO₃. Key functional genes involved in carbon fixation and degradation, nitrogen fixation, denitrification and methane metabolism were stimulated under eCO₂, whereas those involved in N fixation, denitrification and N mineralization were suppressed under eO₃, resulting in the fact that the abundance of some eO₃-supressed genes was promoted to ambient, or eCO₂-induced levels by the interaction of eCO₂+eO₃. Such effects appeared distinct for each treatment and significantly correlated with soil properties and soybean yield. Overall, our analysis suggests possible mechanisms of microbial responses to global atmospheric change factors through the stimulation of C and N cycling by eCO₂, the inhibition of N functional processes by eO₃ and the interaction by eCO₂ and eO₃. This study provides new insights into our understanding of microbial functional processes in response to global atmospheric change in soybean agro-ecosystems.     

Elevated atmospheric carbon dioxide and ozone alter soybean diseases at SoyFACE

Human driven changes in the Earth's atmospheric composition are likely to alter plant disease in the future. We evaluated the effects of elevated carbon dioxide (CO₂) and ozone (O₃) on three economically important soybean diseases (downy mildew, Septoria brown spot and sudden death syndrome‐SDS) under natural field conditions at the soybean free air concentration enrichment (SoyFACE) facility. Disease incidence and/or severity were quantified from 2005 to 2007 using visual surveys and digital image analysis, and changes were related to microclimatic variability and to structural and chemical changes in soybean host plants. Changes in atmospheric composition altered disease expression, but responses of the three pathosystems varied considerably. Elevated CO₂ alone or in combination with O₃ significantly reduced downy mildew disease severity (measured as area under the disease progress curve‐AUDPC) by 39–66% across the 3 years of the study. In contrast, elevated CO₂ alone or in combination with O₃ significantly increased brown spot severity in all 3 years, but the increase was small in magnitude. When brown spot severity was assessed in relation to differences in canopy height induced by the atmospheric treatments, disease severity increased under combined elevated CO₂ and O₃ treatment in only one of the 3 years. The atmospheric treatments had no effect on the incidence of SDS or brown spot throughout the study. Higher precipitation during the 2006 growing season was associated with increased AUDPC severity across all treatments by 2.7 and 1.4 times for downy mildew and brown spot, respectively, compared with drought conditions in 2005. In the 2 years with similar precipitation, the higher daily temperatures in the late spring of 2007 were associated with increased severity of downy mildew and brown spot. Elevated CO₂ and O₃ induced changes in the soybean canopy density and leaf age likely contributed to the disease expression modifications.     

Projected Carbon Dioxide to Increase Grass Pollen and Allergen Exposure Despite Higher Ozone Levels

One expected effect of climate change on human health is increasing allergic and asthmatic symptoms through changes in pollen biology. Allergic diseases have a large impact on human health globally, with 10–30% of the population affected by allergic rhinitis and more than 300 million affected by asthma. Pollen from grass species, which are highly allergenic and occur worldwide, elicits allergic responses in 20% of the general population and 40% of atopic individuals. Here we examine the effects of elevated levels of two greenhouse gases, carbon dioxide (CO₂), a growth and reproductive stimulator of plants, and ozone (O₃), a repressor, on pollen and allergen production in Timothy grass (Phleum pratense L.). We conducted a fully factorial experiment in which plants were grown at ambient and/or elevated levels of O₃ and CO₂, to simulate present and projected levels of both gases and their potential interactive effects. We captured and counted pollen from flowers in each treatment and assayed for concentrations of the allergen protein, Phl p 5. We found that elevated levels of CO₂ increased the amount of grass pollen produced by ∼50% per flower, regardless of O₃ levels. Elevated O₃ significantly reduced the Phl p 5 content of the pollen but the net effect of rising pollen numbers with elevated CO₂ indicate increased allergen exposure under elevated levels of both greenhouse gases. Using quantitative estimates of increased pollen production and number of flowering plants per treatment, we estimated that airborne grass pollen concentrations will increase in the future up to ∼200%. Due to the widespread existence of grasses and the particular importance of P. pratense in eliciting allergic responses, our findings provide evidence for significant impacts on human health worldwide as a result of future climate change.     

Foliar physiology of yellow-poplar ( Liriodendron tulipifera L.) exposed to O3 and elevated CO2 over five seasons.

The chronic effects of ozone (O₃) alone or combined with elevated carbon dioxide (CO₂) on the foliar physiology of unfertilized field-grown yellow-poplar ( Liriodendron tulipifera L.) seedlings were studied from 1992 to 1996. Within open-top chambers, juvenile trees were exposed to the following: charcoal-filtered air (CF); 1× ambient ozone (1XO₃); 1.5× ambient ozone (1.5XO₃); 1.5× ambient ozone plus 700 ppm carbon dioxide (1.5XO₃+CO₂); or chamberless open-air (OA). Seasonal 24-h mean ambient O₃ concentrations ranged from 32 to 46 ppm over the five seasons. Averaged over 5 years, midseason net photosynthesis at saturating light ( A _sat) was reduced by 14% ( P =0.029) and stomatal conductance ( g _s) was reduced, albeit non-significant, by 13% ( P =0.096) in upper canopy foliage exposed to 1.5XO₃-air relative to CF controls. There were no significant differences over the 5 years in A _sat and g _s between trees grown in 1XO₃- and 1.5XO₃-air. Our results support the hypothesis that the magnitude of O₃ effects on A _sat and g _s decreases as saplings age. When averaged over the five seasons of exposure, total chlorophyll concentration ( chl) was not significantly affected by exposure to elevated O₃; however, in 1.5XO₃+CO₂-air, foliar chl was reduced by 33% relative to all others ( P <0.001). In 1.5XO₃+CO₂-air, A _sat was 1.4–1.9 times higher ( P <0.001) and g _s was 0.7 times lower ( P =0.022) than all others. O₃ uptake in juvenile trees exposed to elevated O₃ plus elevated CO₂ (1.5XO₃+CO₂-air) was most comparable to trees exposed to ambient air (1XO₃) throughout the study. These findings suggest that elevated CO₂ may minimize the negative effects of O₃ by reducing O₃ uptake through decreased stomatal conductance.     

Effects of elevated O3, alone and in combination with elevated CO2, on tree leaf chemistry and insect herbivore performance: a meta-analysis

We reviewed the effects of elevated ozone (O₃), alone and in combination with elevated carbon dioxide (CO₂) on primary and secondary metabolites of trees and performance of insect herbivores by means of meta‐analysis. Our database consisted of 63 studies conducted on 22 species of trees and published between 1990 and 2005. Ozone alone had no overall effect on concentrations of carbohydrates or nutrients, whereas in combination with CO₂, elevated O₃ reduced nutrient concentrations and increased carbohydrate concentrations. In contrast to primary metabolites, concentrations of phenolics and terpenes were significantly increased by 16% and 8%, respectively, in response to elevated O₃. Effects of ozone in combination with elevated CO₂ were weaker than those of ozone alone on phenolics, but stronger than those of ozone alone on terpenes. The magnitude of secondary metabolite responses depended on the type of ozone exposure facility and increased in the following order: indoor growth chamber 3 than gymnosperms, as shifts in concentrations of carbohydrate and phenolics were observed in the former, but not in the latter. Elevated O₃ had positive effects on some indices of insect performance: pupal mass increased and larval development time shortened, but these effects were counteracted by elevated CO₂. Therefore, despite the observed increase in secondary metabolites, elevated O₃ tends to increase tree foliage quality for herbivores, but elevated CO₂ may alleviate these effects. Our meta‐analysis clearly demonstrated that effects of elevated O₃ alone on leaf chemistry and some indices of insect performance differed from those of O₃+CO₂, and therefore, it is important to study effects of several factors of global climate change simultaneously.