Difference in ozone uptake in grassland species between open-top chambers and ambient air

Exposure-response data from open-top chamber (OTC) experiments are often directly applied to ambient air (AA) conditions. Because microclimatic conditions are modified and pollutant uptake by plants may differ (i.e. 'chamber effect'), there is concern about the influence of OTCs on these relationships. In addition, AA concentrations are often measured at a height which differs from canopy height and correction for the concentration gradient (i.e. 'gradient effect') is necessary. To quantify the relative contribution of plant characteristics and microclimatic factors to these effects, ozone uptake by horizontal leaves at the top of the canopy was calculated for plants grown in OTCs or AA by using a resistance analogy model. Data from an OTC experiment in 1996/97 for six species typical of productive grasslands were used. Ozone concentration inside OTCs was set equal to the concentration measured at a height of 3 m above ground (C(z(ref))) or at canopy height (C(0)). The gradient effect resulted in a 16-27% lower average C(0) than C(z(ref)), depending on species. The main determinant of the chamber effect was a systematic difference in leaf-to-air vapour pressure deficit between OTCs and AA which affected stomatal resistance and ozone uptake. In case of monocultures both effects were species-specific. In species mixtures the gradient effect differed between mixing ratios, whereas the chamber effect was species-specific. Because of the inter-specific difference in the chamber effect on ozone uptake, it is concluded that ozone effects on species mixtures differ systematically between OTCs and AA. The data underline that extrapolation of ozone flux-response relationships from OTC experiments must be based on canopy-level ozone concentrations, and that these relationships should be applied only to single species under microclimatic conditions similar to those prevailing in the experiment.     

Effects of ozone on managed pasture: I. Effects of open-top chambers on microclimate, ozone flux, and plant growth

Open-top chambers (OTC) were established in a field of managed pasture, and environmental parameters were recorded inside and outside to study the influence of OTCs on radiation, air temperature (T(air)), saturation vapour pressure deficit (svpd), and soil water content in relationship to plant growth and yield. Canopy development in OTCs supplied with non-filtered air (NF) and in ambient (AA) plots was followed by measuring leaf area index (LAI). The dry matter yield was determined after three growth periods in each of two consecutive seasons. Boundary layer conductance (g(bw)) and wind speed (u) were measured along a vertical profile, and day-time flux were measured along a vertical profile, and day-time flux of O(3) was estimated throughout the experiment on the basis of a mass balance. The vertical profile of u showed values in the range 1-1.2 m s(-1) at the top of the canopy, and maximum g(bw) was 20-25 mm s(-1). Average reduction in global radiation in OTCs was 25%, and volumetric soil water content was reduced by about 5%. Daily mean T(air) was increased by 1.3 degrees C, mean daily maximum svpd by 0.08 kPa, and the temperature sum (degree days with base temperature of +5 degrees C) by 12%. Fluctuations in the difference in daily mean T(air) and svpd during the daytime between OTCs and ambient air were related to canopy structure. Differences were largest after each cut and declined with increasing LAI. A small effect of changes in LAI on T(air) and svpd occurred during periods with low soil water content. The flux of O(3) in OTCs was largest (>100 microg m(-2) min(-1)) before and smallest (<20 microg m(-2) min(-1)) after each cut. Calculated deposition velocities for O(3) (nu(d)) in the range 0-3 cm s(-1) were generally higher than those measured under most field conditions. Overall, in OTCs the deficit in soil and atmospheric moisture was larger than in the open field, and the increase in daily mean T(air) was strongly influenced by the stage of canopy development. Changes in microclimate and incoming radiation affected pasture development. LAI was slightly reduced in OTCs as compared to AA plots. The total accumulated dry matter yield for all six growth periods was only about 7% lower in OTCs, but the contribution of clover to total forage mass declined during the experiment. OTCs had no significant effect on weeds. The results indicate that OTCs reduced the competitiveness of clover, and that the increase in growth of grasses compensates for the loss in clover yield. The shift in species composition caused by OTCs must be considered when studying the effect of pollutants on pasture.     

Assessing plant response to ambient ozone: growth of ozone-sensitive loblolly pine seedlings treated with ethylenediurea or sodium erythorbate

One-year-old seedlings from an ozone-sensitive half-sib family of loblolly pine (Pinus taeda L.) were transplanted into replicated plots in blocks in a large forest clearing near Nacogdoches, Texas. Seedlings were either non-treated (controls) or treated bi-weekly with foliar sprays of ethylenediurea (EDU), at 150, 300 or 450 ppm or sodium erythorbate (NaE), at 515, 1030, or 1545 ppm, for three growing seasons. Results from the final third year harvest indicated that both EDU and NaE increased all growth parameters, with significant differences only for EDU at 450 ppm. Both EDU and NaE would be useful for long-term studies on assessing the effects of ambient ozone on established native plants.     

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

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

Black cherry growth response to ambient ozone and EDU

Field-grown black cherry (Prunus serotina Ehrh.) seedlings were treated with the antioxidant ethylenediurea (EDU) to evaluate height, diameter, and above-ground dry-weight biomass growth response to ambient ozone over four years. Nine blocks with 44 trees/block were used in a randomized complete block design with three foliar spray treatments: (1) 1000 ppm EDU mixed with a surfactant and water; (2) surfactant mixed with water; and (3) water only. In each growing season treatments were applied seven times at approximately 10-day intervals. Repeated measures analysis of variance indicated significant (P< or =0.05) treatment and year effects for log-transformed height and diameter growth over the four-year period. After four years, EDU-treated trees were approximately 17% taller and stem diameters were 21% greater than non-EDU-treated trees. Total above-ground dry-weight biomass at the end of four years was 47% greater for EDU-treated trees compared to non-EDU-treated trees.     

Effects of air pollutants on the growth and antioxidative system of Norway spruce exposed in open-top chambers

Grafted Norway spruce trees were subjected to exposure beginning in April 1988, to one of four different air treatments in open-top chambers: Charcoal filtered air (CF), non-filtered air (NF), non-filtered air with the addition of O(3) during summer (NFO), and SO(2) plus NO(2) during winter (NFOSN). CF trees were considered as the reference group. No effects on growth parameters were observed. Samples of the two youngest needle year classes were taken late in November 1989 for enzyme determinations. The activity of ascorbic acid peroxidase (A-POD) increased the same level in all treatments, and activities of catalase and dehydroascorbic acid reductase (DHA-R) increased only in NF and NFO treatments. A higher level of activity in the NFOSN treatment was observed only for glucose-6-phosphate-dehydrogenase (Glc-6-P-DH) and non-specific peroxidase (POD). Isoelectric focusing of POD showed a changed pattern in the NFOSN treatment. Neither activity nor isoelectric focusing of superoxidase dismutase (SOD) was changed in any of the treatments.     

Effects of ozone on grain quality of wheat grown in open-top chambers: three years of experimentation

Field trials with winter wheat (Triticum aestivum L.) grown in open-top chambers and exposed to either ambient filtered (F) or unfiltered (NF) air were carried out for 3 years in northern Italy. The experiments were performed at two places, a rural and an urban site in the Po Plain. The data obtained show no significant difference in protein, starch and glucose contents between the treatments; nevertheless, a tendency towards an increase in glucose and a decrease in starch contents was observed in the F chambers compared with NF. In addition, potassium levels were lower in F than in NF treatment, while calcium were higher. Ozone levels observed in the Po Valley are not sufficiently high to cause significant differences in quality parameters of winter wheat.     

Effects of elevated carbon dioxide and ozone on the growth and yield of potatoes (Solanum tuberosum) grown in open-top chambers

Potato (Solanum tuberosum cv. Bintje) was grown in open-top chambers under three carbon dioxide (ambient and seasonal mean concentrations of 550 and 680 mumol mol-1 CO2) and two ozone concentrations (ambient and an 8 h day-1 seasonal mean of 50 nmol mol-1 O3) between emergence and final harvest. Periodic non-destructive measurements were made and destructive harvests were carried out at three key developmental stages (24, 49 and 101 days after emergence) to establish effects on growth and tuber yield. Season-long exposure to elevated O3 reduced above-ground dry weight at final harvest by 8.4% (P < 0.05), but did not affect tuber yields. There was no significant interaction between CO2 and O3 for any of the growth and yield variables examined. Non-destructive analyses revealed no significant effect of elevated CO2 on plant height, leaf number or green leaf area ratio. However, destructive harvests at tuber initiation and 500 degrees Cd after emergence showed that above-ground dry weight (8 and 7% respectively) and tuber yield (88 and 44%) were significantly increased (P < 0.05) in the 550 mumol mol-1 CO2 treatment. Responses to 550 and 680 mumol mol-1 CO2 were not significantly different for most parameters examined, suggesting the existence of an upper limit to the beneficial influence of CO2 enrichment. Significant effects on above-ground dry weight and tuber yield were no longer apparent at final harvest, although tuber numbers were increased (P < 0.05) under elevated CO2, particularly in the smaller size categories. The results show that the O3 treatment imposed was insufficient to reduce tuber yields and that, although elevated CO2 enhanced crop growth during the early stages of the season, this beneficial effect was not sustained to maturity.     

Effects of ozone on the grain composition of spring wheat grown in open-top field chambers

In a three-year study carried out at a rural site in Switzerland, spring wheat (Triticum aestivum L. cv. Albis) was exposed to different levels of ozone (O(3)) in open-top-field chambers from the two-leaf stage until harvest. Field plots in ambient air (AA) were used for comparison. Grain recovered from the different treatments was analyzed for minerals (Ca, Mg, K, P), starch, protein, amino acids and alpha-tocopherol, in order to investigate the effect of O(3) on grain composition. Chamber-enclosure had small effects on some parameters (K, protein), but not on others (starch), as shown by the comparison of data from the AA and non-filtered-air treatment (NF). Differences between NF and charcoal-filtered air (CF) were very small. At O(3) concentrations higher than in the NF treatment (O(3)-1 = 1.5xNF and O(3)-2 = 2.5xNF), mineral contents were higher than in the NF and CF treatments. Protein content was increased only in the O(3)-2 treatment. Starch contents decreased from about 63% in the CF treatment to 54% in the O(3)-2 tratment. No effect of O(3) on the content of alpha-tocopherol and on the essential amino acid index of the protein was observed. It is concluded that compositional changes in wheat grain in response to O(3) are minor, and that ambient O(3) is not likely to cause important changes.     

Effects of ozone on 14C translocation velocity and growth of spring wheat (Triticum aestivum L.) exposed in open-top chambers

Growth and yield were reduced but (14)C translocation velocity was not affected by increasing levels of ozone in spring wheat exposed in open top chambers to the following treatments: charcoal filtered air (CF), non-filtered ambient air (NF), or NF with addition of 30 microl litre(-1) ozone, 8 h daily (NFO). Destructive harvests were performed at anthesis and at maturity. Parts of the flag leaf or the second leaf were exposed to (14)CO(2) in small cuvettes for 5 min before, during and after anthesis. The translocation velocity was followed by autoradiography and scintillation counting of the plants frozen and lyophilized at different times after labelling. The label was transported at the same velocity in all the treatments. Ozone induced changes in carbon allocation or partitioning should probably be explained as amounts of carbon transported (mg s(-1)), rather than as transportation velocity (mm s(-1)). The amount translocated may be governed by source conditions under O(3) stress: reduced healthy green biomass and photosynthesis, but perhaps also by impairment of phloem loading because of membrane damage.     

Ozone flux to vegetation and its relationship to plant response and ambient air quality standards

The National Ambient Air Quality Standard (NAAQS) for ozone is based on occurrences of the maximum 8 h average ambient ozone concentration. However, biologists have recommended a cumulative ozone exposure parameter to protect vegetation. In this paper we propose a third alternative which uses quantifiable flux-based numerical parameters as a replacement for cumulative ambient parameters. Herein we discuss the concept of ozone flux as it relates to plant response and the NAAQS, and document information needed before a flux-based ozone NAAQS for vegetation can be implemented. Additional research is needed in techniques for determining plant uptake and in the quantification of plant defensive mechanisms to ozone. Models which include feedback mechanisms should be developed to relate ozone flux, loading, and detoxification with photosynthesis and plant productivity.     

Influence of ozone and ethylenediurea (EDU) on growth and yield of bean (Phaseolus vulgaris L.) in open-top field chambers

Greenhouse and ambient air experiments have shown ethylene diurea (EDU) to be a strong and specific protective suppressant of ozone injury in plants. To examine how EDU affects plant responses to various ozone (O(3)) levels under controlled field conditions, Phaseolus vulgaris L. cv. Lit was treated with 150 ppm EDU every 14 days and exposed in open-top chambers to charcoal-filtered air (CF), nonfiltered air (NF) or two cf treatments with ozone added. The ozone treatments were proportional additions of one (CF1) and two (CF2) times ambient ozone levels. The mean ozone concentrations in the CF, NF, CF1 and CF2 treatments were 0.98, 14.1, 14.98 and 31.56 nl litre(-1). A two-way split plot ANOVA revealed that shoot dry weight was significantly reduced by ozone. EDU treatment was highly significant for leaf dry weight, root dry weight and shoot dry weight, but not for pod dry weight; leading to a higher biomass of EDU-treated plants. Ozone/EDU interactions were significant for root weight only, indicating that EDU reduced growth suppression by ozone. These results show that EDU action on plant biomass could be interpreted as a delay in senescence since EDU-treated plants showed a significant decreased biomass loss even in the CF treatment.     

No general ozone-specific indicator among the hexane- and dichloromethane-soluble components of Picea abies needles exposed to ozone in open-top chambers

The effect of ozone (< 10, 200 or 400 microg m(-3) on hexane- and dichloromethane-soluble components of Picea abies needles was determined by fumigating potted grafts from mature trees. The trees (>55 and 125 years, 2.5 m high), representing six clones of Norway spruce, were fumigated in open-top chambers at two locations in Norway for one growth season. The needles were extracted with hexane and dichloromethane; 142 compounds from the hexane extract and 164 silylated compounds from the dichloromethane extract were analysed by gas chromatography although no identifications were made. The concentration of four of the compounds from the hexane extract changed with ozone dose in a way that made them promising as indicators, but the present analytical method could not verify this possibility. None of the other 302 compounds qualified as a general indicator of ozone stress in Norway spruce, as none changed its concentration with ozone dose consistently in all romets of all clones. Most of the variation in the experiment is mainly attributable to genetic variation and to climate.     

Size-mediated foliar response to ozone in black cherry trees

Local ozone concentration and visible foliar injury were measured over the 1994 growing season on open-grown black cherry (Prunus serotina Ehrh.) trees of varying size (age) within forest stands and adjacent openings at a site in north-central Pennsylvania. Relationships were determined between visible ozone injury and ozone exposure, as well as calculated between injury and ozone uptake expressed as the product of stomatal conductance and ozone concentration. In addition, simultaneous measurements of visible symptoms and leaf gas exchange were also conducted to determine the correlation between visible and physiological injury and ozone exposure. By September, the amount of leaf area affected by visible foliar ozone injury was greatest for seedlings (46%), followed by canopy trees (20%) and saplings (15%). A large amount of variability in foliar ozone symptom expression was observed among trees within a size class. Sum40 and Sum60 (ozone concentration > 40 and > 60 nl liter(-1)) cumulative exposure statistics were the most meaningful indices for interpretation of foliar injury response. Seedlings were apparently more sensitive to ozone injury than larger trees because their higher rates of stomatal conductance resulted in higher rates of ozone uptake. Seedlings also had higher rates of early leaf abscission than larger trees with an average of nearly 30% of the leaves on a shoot abscised by 1 September compared to approximately 5% for larger trees. However, per unit ozone uptake into the leaf, larger trees exhibited larger amounts of foliar injury. The amount of visible foliar injury was negatively correlated (r(2) = 0.82) with net photosynthetic rates, but was not related to stomatal conductance. Net photosynthesis and stomatal conductance thus became uncoupled at high levels of visible foliar injury.     

Exposure of oats, Avena sativa L., to filtered and unfiltered air in open-top chambers: effects on grain yield and quality

Field grown oats, Avena sativa L. cv Vital, were exposed to filtered and unfiltered air from anthesis until harvest in open-top chambers at a site in south-west Sweden. Ambient plots were used to study the influence of the chamber itself. With the exception of the number of grains per panicle, which was significantly higher in the charcoal-filtered treatment, no significant filtration effects were obtained for any of the plant growth parameters studied), i.e. grain yield, number of panicles per square metre, 1000-grain weight, straw yield and harvest index.) The chamber had a significant negative effect on grain yield, 1000-grain weight and straw yield. None of the yield quality parameters that were measured, such as crude protein content, crude fibre content, fat content, volume weight of the grain and water content of the grain at harvest, were significantly influenced by either air filtration, or the presence of the chamber. The chlorophyll content of the flag leaves was higher in the charcoal-filtered treatment than in the non-filtered and ambient air treatments towards the end of the experiment, indicating that filtering of the air delayed senescence. The decline of the shoot area after the onset of plant senescence proceeded faster in both chamber treatments. The faster development in the chamber was explained by the faster accumulation of thermal time in the chamber.     

Variation of ozone concentrations in the ambient air of Baghdad

Measurements of ozone concentration in the ambient air of the city of Baghdad were carried out for the period October 1983 to October 1984. The O₃, probably of local origin, showed a typical diurnal and seasonal variation. Maximum daily 1-h O₃ concentrations higher than the international ambient air quality standards were observed regularly during the summer months. High O₃ concentrations during the night were also observed. Scatter diagrams were used to relate the O₃ concentrations with temperature, solar radiation and humidity.     

Prediction of ozone concentration in ambient air using multivariate methods

Multivariate statistical methods including pattern recognition (Principal Component Analysis--PCA) and modeling (Multiple Linear Regression--MLR, Partial Least Squares--PLS, as well as Principal Component Regression--PCR) methods were carried out to evaluate the state of ambient air in Miskolc (second largest city in Hungary). Samples were taken from near the ground at a place with an extremely heavy traffic. Although PCA is not able to determine the significance of variables, it can uncover their similarities and classify the cases. PCA revealed that it is worth to separate day and night data because different factors influence the ozone concentrations during all day. Ozone concentration was modeled by MLR and PCR with the same efficiency if the conditions of meteorological parameters were not changed (i.e. morning and afternoon). Without night data, PCR and PLS suggest that the main process is not a photochemical but a chemical one.     

Effects of ozone exposure in open-top chambers on poplar (Populus nigra) and beech (Fagus sylvatica): a comparison

Rooted cuttings of poplar (Populus nigra) and seedlings of beech (Fagus sylvatica) were exposed to ozone in open-top chambers for one growing season. Three treatments were applied: charcoal-filtered (CF), non-filtered (NF) and non-filtered air plus 30 ppb (nl l(-1)) ozone (NF+). Extra ozone was only added on clear days, from 09:00 until 17:00-20:00. The AOT40s (accumulated exposure over a threshold of 40 ppb), calculated from April to September were 4055 ppb.h for the NF and 8880 ppb.h for the NF+ treatments. For poplar ozone exposure caused highly significant reductions in growth rate, light-saturated net CO(2) assimilation rate, stomatal conductance, F(v)/F(m) and chlorophyll content. The largest effects were observed in August at which time ozone concentrations were elevated. A reduction was noticed in new leaf production, while accelerated ageing and visible damage to leaves caused high leaf losses. For beech the responses were similar but less pronounced: ozone exposure resulted in non-significant growth reductions, slight changes in light-saturated photosynthesis and accelerated leaf abscission. The chlorophyll content of beech leaves was not affected by the ozone treatments. The results confirmed previous observations that fast-growing tree species, such as most poplar species and hybrids, are more sensitive and responsive to tropospheric ozone than slower-growing species, such as beech. The growth reductions observed and reported here for beech were within the range of those reported in relationship to the AOT40 (accumulated exposure over a threshold of 40 ppb) critical level for ozone.     

Detecting plant effects is necessary to give biological significance to ambient ozone monitoring data and predictive ozone standards

The use of mechanical monitors and passive samplers has made it possible to assess concentrations of ozone over wide areas and to develop air quality standards, like AOT40 and SUM60. Monitored ozone data and AOT40 and SUM60 are also used to predict ozone injury on local and regional scales. The data and the standards do not include or account for environmental and biological variables that affect ozone uptake and plant injury. Ground proofing via vegetation surveys must be done to verify and validate plant injury predictions. If this is not done, then the standards have no biological significance and are only exercises in air quality assessment.     

Modeling plant response to tropospheric ozone: a critical review

The advantages and disadvantages, benefits and limitations, of a number of published mathematical models representing the effects of ozone on crops and native vegetation are described. Several levels of modeling are addressed: word models, graphic models, mathematical models, and computer simulation implementation. Special attention is given to evaluating: (1) how the interaction between ozone exposure and vegetation effects is quantified, (2) the status of field testing of the model, and (3) the adequacy of information for enabling other investigators to replicate the model for further testing. Original contributions, not previously published, are made in this evaluation in the form of: (1) graphic model flow charts for published models, (2) clarification of mathematical equations for existing models, (3) graphic forms of functional relations comprising portions of models, and (4) graphic displays of model output performance versus observed data. The models that are evaluated cover acute exposure-response models, statistical and mechanistic-process models, including a partial model of ambient exposure versus ozone flux, and uptake. They also cover chronic exposure statistical approaches, including time-series modeling, mechanistic-process models, 'disintegrated' models of forest system simulations, chronic flux density-uptake-response, and models for regional effects assessment in forests and agricultural lands.