Effects of long-term, free-air ozone fumigation on the cytokinin content of mature beech trees

We present the results of a study of the effects of chronic exposure to elevated ozone on the cytokinins of mature beech trees. Methods for analysing the cytokinin (CK) content of beech (FAGUS SYLVATICA) were developed using seven enzyme-linked immunosorbent assays (ELISAs). Samples taken during 2003 and 2004 from 10 mature beech trees in Kranzberg forest, 5 trees exposed to twice ambient ozone (2 x O(3)) by free-air fumigation and 5 control trees (1 x O(3)), were analysed. In 2003 and 2004 the cytokinin content of leaf samples followed a similar seasonal pattern. In leaf samples, the content of aromatic types was equal to that of the isoprenoid types. In root samples, the level of aromatic types was no different from leaves, but that of the isoprenoid types was much higher. Leaf and phloem cytokinin contents for 2 x O(3) trees were lower than for 1 x O(3) at almost all sampling times. The effect of ozone was greater for leaves in the sun crown than for leaves in the shade crown. By contrast, the root and xylem contents of cytokinin for 2 x O(3) trees were greatly elevated over the values for 1 x O(3) trees early in the growing season. We propose that O(3)-associated CK destruction in leaves reduces CK-mediated root growth suppression. The resulting increases in root growth and ectomycorrhiza, reported by other groups in the Kranzberg forest project, are likely to be responsible for the increased CK export in xylem, although O(3)-associated CK destruction in the leaves appears to nullify this increase.     

CASIROZ: Root parameters and types of ectomycorrhiza of young beech plants exposed to different ozone and light regimes

Tropospheric ozone (O(3)) triggers physiological changes in leaves that affect carbon source strength leading to decreased carbon allocation below-ground, thus affecting roots and root symbionts. The effects of O(3) depend on the maturity-related physiological state of the plant, therefore adult and young forest trees might react differently. To test the applicability of young beech plants for studying the effects of O(3) on forest trees and forest stands, beech seedlings were planted in containers and exposed for two years in the Kranzberg forest FACOS experiment (Free-Air Canopy O(3) Exposure System, http://www.casiroz.de ) to enhanced ozone concentration regime (ambient [control] and double ambient concentration, not exceeding 150 ppb) under different light conditions (sun and shade). After two growing seasons the biomass of the above- and below-ground parts, beech roots (using WinRhizo programme), anatomical and molecular (ITS-RFLP and sequencing) identification of ectomycorrhizal types and nutrient concentrations were assessed. The mycorrhization of beech seedlings was very low ( CA. 5 % in shade, 10 % in sun-grown plants), no trends were observed in mycorrhization (%) due to ozone treatment. The number of Cenococcum geophilum type of ectomycorrhiza, as an indicator of stress in the forest stands, was not significantly different under different ozone treatments. It was predominantly occurring in sun-exposed plants, while its majority share was replaced by Genea hispidula in shade-grown plants. Different light regimes significantly influenced all parameters except shoot/root ratio and number of ectomycorrhizal types. In the ozone fumigated plants the number of types, number of root tips per length of 1 to 2 mm root diameter, root length density per volume of soil and concentration of Mg were significantly lower than in control plants. Trends to a decrease were found in root, shoot, leaf, and total dry weights, total number of root tips, number of vital mycorrhizal root tips, fine root (mass) density, root tip density per surface, root area index, concentration of Zn, and Ca/Al ratio. Due to the general reduction in root growth indices and nutrient cycling in ozone-fumigated plants, alterations in soil carbon pools could be predicted.     

Gas exchange and antioxidative compounds in young beech trees under free-air ozone exposure and comparisons to adult trees

Three-year-old beech (Fagus sylvatica) seedlings growing in containers were placed into the sun and shade crown of a mature beech stand exposed to ambient (1 x O(3)) and double ambient (2 x O(3)) ozone concentrations at a free-air exposure system ("Kranzberg Forst", Germany). Pigments, alpha-tocopherol, glutathione, ascorbate, and gas exchange were measured in leaves during 2003 (a drought year) and 2004 (an average year). Sun-exposed seedlings showed higher contents of antioxidants, xanthophylls, and beta-carotene and lower contents of chlorophyll, alpha-carotene, and neoxanthin than shade-exposed seedlings. In 2003 sun-exposed seedlings showed higher contents of carotenoids and total glutathione and lower net photosynthesis rates (A(max)) compared to 2004. O(3) exposure generally affected the content of chlorophyll, the xanthophyll cycle, and the intercellular CO(2) concentration (c(i)). Seedlings differed from the adjacent adult trees in most biochemical and physiological parameters investigated: Sun exposed seedlings showed higher contents of alpha-tocopherol and xanthophylls and lower contents of ascorbate, chlorophyll, neoxanthin, and alpha-carotene compared to adult trees. Shade exposed seedlings had lower contents of xanthophylls, alpha-carotene, and alpha-tocopherol than shade leaves of old-growth trees. In 2003, seedlings had higher A(max), stomatal conductance (g(s)), and c(i) under 2 x O(3) than adult trees. The results showed that shade acclimated beech seedlings are more sensitive to O(3), possibly due to a lower antioxidative capacity per O(3) uptake. We conclude that beech seedlings are uncertain surrogates for adult beech trees.     

Simulating stand climate, phenology, and photosynthesis of a forest stand with a process-based growth model

In the face of climate change and accompanying risks, forest management in Europe is becoming increasingly important. Model simulations can help to understand the reactions and feedbacks of a changing environment on tree growth. In order to simulate forest growth based on future climate change scenarios, we tested the basic processes underlying the growth model BALANCE, simulating stand climate (air temperature, photosynthetically active radiation (PAR) and precipitation), tree phenology, and photosynthesis. A mixed stand of 53- to 60-year-old Norway spruce (Picea abies) and European beech (Fagus sylvatica) in Southern Germany was used as a reference. The results show that BALANCE is able to realistically simulate air temperature gradients in a forest stand using air temperature measurements above the canopy and PAR regimes at different heights for single trees inside the canopy. Interception as a central variable for water balance of a forest stand was also estimated. Tree phenology, i.e. bud burst and leaf coloring, could be reproduced convincingly. Simulated photosynthesis rates were in accordance with measured values for beech both in the sun and the shade crown. For spruce, however, some discrepancies in the rates were obvious, probably due to changed environmental conditions after bud break. Overall, BALANCE has shown to respond to scenario simulations of a changing environment (e.g., climate change, change of forest stand structure).     

Suitability of a combined stomatal conductance and photosynthesis model for calculation of leaf-level ozone fluxes

Currently, the most important source of uncertainty in stomatal ozone flux ( FO3) modelling is the stomatal conductance ( gst) factor. Hence FO3 model accuracy will strongly depend on the gst model being implemented. In this study the recently developed semi-empirical Gst model of Dewar was coupled to the widely known biochemical photosynthesis ( An) model of Farquhar. The Gst performance of this model combination was evaluated with a 4-month time series of beech ( Fagus sylvatica L.) measurements. The Gst model was hereto optimized in two steps to a 4-day and a 8-day period. A comparison between the modelled and measured gst to O(3) (gstO3) revealed a rather good overall performance (R(2)=0.77). Errors between the model combination and the measurements are thought to be largely caused by a moderate performance of the AN model, due to poor parameterization. Two 2-day periods with distinctly differing soil and meteorological conditions were chosen to give a picture of the daily gst performance. Although instant relative differences between modelled and measured gstO3 are sometimes high, the model combination is able to simulate the rough daily courses of gstO3 and hence FO3 reasonably well. Further improvement on full parameterization of the gst model and a well-parameterized An model to be linked to are needed to draw founded conclusions about its performance. Future efforts hereto are certainly justified since the model's mechanistic nature makes it a tool able to model gst variation in space and time, O(3) effects on gst, and effective FO3.     

A mechanistic model of photoinhibition

A mechanistic model was developed, to simulate the main facets of photoinhibition in phytoplankton. Photoinhibition is modelled as a time dependent decrease in the initial slope of a photosynthesis versus irradiance curve, related to D1 (photosystem II reaction centre protein) damage and non-photochemical quenching. The photoinhibition model was incorporated into an existing ammonium-nitrate nutrition interaction model capable of simulating photoacclimation and aspects of nitrogen uptake and utilization. Hence the current model can simulate the effects of irradiance on photosynthesis from sub-saturating to inhibitory photon flux densities, during growth on different nitrogen sources and under nutrient stress. Model output conforms well to experimental data, allowing the extent of photoinhibition to be predicted under a range of nutrient and light regimes. The ability of the model to recreate the afternoon depression of photosynthesis and the enhancement of photosynthesis during fluctuating light suggests that these two processes are related to photoinhibition. The model may be used to predict changes in biomass and/or carbon fixation under a wide range of oceanographic situations, and it may also help to explain the progression to dominance of certain algal species, and bloom formation under defined irradiance and nutrient conditions.     

Physiological responses of Fagus sylvatica L. exposed to low levels of ozone in open-top chambers

Four-year-old beech seedlings were fumigated with three levels of ozone for 2 consecutive years in open-top chambers. During the second growth season different physiological measurements were conducted before and during daily fumigation. A 25–40% decrease in net photosynthesis was seen during fumigation, whereas no differences were detected before fumigation in July. In August lasting effects in net photosynthesis were seen. The apparent quantum yield was decreased after fumigation. Stomatal conductance was generally decreased during fumigation, but transpiration was reduced relatively less than net photosynthesis indicating a lower water use efficiency of the trees exposed to ozone. Chlorophyll fluorescence (F_v/F_m) showed additive reductions in relation to ozone and light.     

Assessing the future global impacts of ozone on vegetation

Ozone is a major secondary air pollutant, the current concentrations of which have been shown to have significant adverse effects on crop yields, forest growth and species composition. In North America and Europe, emissions of ozone precursors are decreasing but in other regions of the world, especially Asia, where much less is known about its impacts, they are increasing rapidly. There is also evidence of an increase in global background ozone concentrations, which will lead to significant changes in global ozone exposure over this century, during which direct and indirect effects of other changes in the global atmosphere will also modify plant responses to ozone. This paper considers how far our current understanding of the mechanisms of ozone impacts, and the tools currently used for ozone risk assessment, are capable of evaluating the consequences of these changing global patterns of exposure to ozone. Risk assessment based on relationships between external concentration and plant response is inadequate for these new challenges. New models linking stomatal flux, and detoxification and repair processes, to carbon assimilation and allocation provide a more mechanistic basis for future risk assessments. However, there are a range of more complex secondary effects of ozone that are not considered in current risk assessment, and there is an urgent need to develop more holistic approaches linking the effects of ozone, climate, and nutrient and water availability, on individual plants, species interactions and ecosystem function.     

Beech leaf colonization by the endophyte Apiognomonia errabunda dramatically depends on light exposure and climatic conditions

Ozone and light effects on endophytic colonization by Apiognomonia errabunda of adult beech trees (Fagus sylvatica) and their putative mediation by internal defence compounds were studied at the Kranzberg Forest free-air ozone fumigation site. A. errabunda colonization was quantified by "real-time PCR" (QPCR). A. errabunda-specific primers allowed detection without interference by DNA from European beech and several species of common genera of plant pathogenic fungi, such as Mycosphaerella, Alternaria, Botrytis, and Fusarium. Colonization levels of sun and shade leaves of European beech trees exposed either to ambient or twice ambient ozone regimes were determined. Colonization was significantly higher in shade compared to sun leaves. Ozone exhibited a marginally inhibitory effect on fungal colonization only in young leaves in 2002. The hot and dry summer of 2003 reduced fungal colonization dramatically, being more pronounced than ozone treatment or sun exposure. Levels of soluble and cell wall-bound phenolic compounds were approximately twice as high in sun than in shade leaves. Acylated flavonol 3- O-glycosides with putatively high UV-B shielding effect were very low in shade canopy leaves. Ozone had only a minor influence on secondary metabolites in sun leaves. It slightly increased kaempferol 3- O-glucoside levels exclusively in shade leaves. The frequently prominent hydroxycinnamic acid derivative, chlorogenic acid, was tested for its growth inhibiting activity against Apiognomonia and showed an IC50 of approximately 8 mM. Appearance of Apiognomonia-related necroses strongly correlated with the occurrence of the stress metabolite, 3,3',4,4'-tetramethoxybiphenyl. Infection success of Apiognomonia was highly dependent on light exposure, presumably affected by the endogenous levels of constitutive phenolic compounds. Ozone exerted only minor modulating effects, whereas climatic factors, such as pronounced heat periods and drought, were dramatically overriding.     

Analysing the role of soil properties, initial biomass and ozone on observed plant growth variability in a lysimeter study

This simulation study is based on a lysimeter experiment with juvenile beech trees (Fagus sylvatica L.) which were grown under ambient or doubled ambient atmospheric ozone concentrations. The aim of the study was to analyze the role of differences in soil properties, differences in initial biomass and ozone impacts on observed plant growth variability at the eight lysimeters of this experiment. For this purpose, we established a new simulation model based on the model system Expert-N by coupling soil water and nitrogen transport models with the plant growth model PLATHO, which was already tested and applied for juvenile beech. In order to parameterize the soil model, for all lysimeters soil hydraulic parameters as well as carbon and nitrogen stocks were measured. Simulation results reveal that the observed decreased growth rates under elevated ozone are due to ozone impacts on plant growth, whereas the high plant growth variability between lysimeters is to a major part the consequence of differences in soil hydraulic properties. Differences in initial biomass are of minor importance to explain plant growth variability in this experiment.     

Ozone effects on wheat in relation to CO2: modelling short‐term and long‐term responses of leaf photosynthesis and leaf duration

A combined stomatal–photosynthesis model was extended to simulate the effects of ozone exposure on leaf photosynthesis and leaf duration in relation to CO₂. We assume that ozone has a short‐term and a long‐term effect on the Rubisco‐limited rate of photosynthesis, A_c. Elevated CO₂ counteracts ozone damage via stomatal closure. Ozone is detoxified at uptake rates below a threshold value above which A_c decreases linearly with the rate of ozone uptake. Reduction in A_c is transient and depends on leaf age. Leaf duration decreases depending on accumulated ozone uptake. This approach is introduced into the mechanistic crop simulation model AFRCWHEAT2. The derived model, AFRCWHEAT2‐O3, is used to test the capability of these assumptions to explain responses at the plant and crop level. Simulations of short‐term and long‐term responses of leaf photosynthesis, leaf duration and plant and crop growth to ozone exposure in response to CO₂ are analysed and compared with experimental data derived from the literature. The model successfully reproduced published responses of leaf photosynthesis, leaf duration, radiation use efficiency and final biomass of wheat to elevated ozone and CO₂. However, simulations were unsatisfactory for cumulative radiation interception which had some impact on the accuracy of predictions of final biomass. There were responses of leaf‐area index to CO₂ and ozone as a result of effects on tillering which were not accounted for in the present model. We suggest that some model assumptions need to be tested, or analysed further to improve the mechanistic understanding of the combined effects of changes in ozone and CO₂ concentrations on leaf photosynthesis and senescence. We conclude that research is particularly needed to improve the understanding of leaf‐area dynamics in response to ozone exposure and elevated CO₂.     

Growth of adult Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica L.) under free-air ozone fumigation

This study attempted to detect the impact of ozone on adult trees of Norway spruce ( Picea abies [L.] Karst.) and European beech ( Fagus sylvatica L.) in an experimental mixed stand in Southern Bavaria, Germany. The aim was to examine whether there is a decrease in growth when trees are exposed to higher than atmospheric concentrations of ozone. This exposure was put into effect using a free-air fumigation system at tree crown level. Growth analysis was carried out on a group of 47 spruce and 36 beech trees, where radial stem increment at breast height - a sensitive index for stress - was measured. The ozone monitoring system allowed values to be obtained for the accumulated ozone exposure (SUM00) of each individual tree, so that their radial increment over three years could be correlated with the corresponding ozone exposure for the same time period. Correlation and regression analysis were then carried out to test the influence of ozone on diameter increment. In both spruce and beech, the initial stem diameter was the most influential factor on radial increment in the following year. A linear model was applied, including the diameter of the preceding year and the ozone exposure of the current year as predicting factors. For spruce trees, a significant negative influence of ozone exposure was found. In contrast, no significant ozone effect on diameter increment of beech was detected. The effect of ozone stress on a large spruce tree can lead to a decrease in potential radial increment of 22 %. The results are discussed in relation to other stress factors such as drought and lack of light.     

用光合-蒸散耦合模型模拟冬小麦CO2通量的日变化

根据SPAC理论建立了一个冬小麦光合和蒸散的耦合模型.冬小麦CO2通量包括冠层光合、呼吸和土壤呼吸.冠层光合采用了Farquhar光合作用生化模型,并通过冠层阻力的参数化将光合作用与蒸腾作用耦合起来.用涡度相关方法观测了CO2通量,对模型进行了验证,结果显示模型可以较好地模拟CO2通量日变化过程.对模型的敏感性分析发现日间CO2通量最敏感的参数是初始量子效率.其次,CO2通量对光响应曲线凸度、CO2补偿点、凋萎点和叶面积指数的变化也有着较强的敏感性;夜间CO2通量敏感的参数是最适温度下Rubisco催化能力和暗呼吸参数.     

Changes in the community of ectomycorrhizal fungi and increased fine root number under adult beech trees chronically fumigated with double ambient ozone concentration

Forest soils are an important but under-studied part of forest ecosystems. The effects of O(3) on below-ground processes in a mature forest have only received limited attention so far. In our study, we have analysed the community of ectomycorrhizal fungi and beech fine root dynamics over two growing seasons (2003 - 2004) in a 70-year old mixed spruce-beech forest stand, in which two groups of five adult beech trees were either fumigated by 2 x ambient ozone concentration or used as control. The main difference between previous studies and our approach was that previous studies were performed on seedlings in pot experiments or in closed or open top chambers, and not IN SITU, in a mature forest stand. Although beech is a relatively unresponsive species to tropospheric O(3), we found a pronounced effect of 2 x O(3) on the number of vital ectomycorrhizal root tips and non-turgescent fine roots. Both categories of roots were significantly increased when compared to controls in two consecutive years at each sampling event. The number of types of ectomycorrhizae and species richness increased in 2004, but not in the extremely dry year 2003. We hypothesised that the observed changes might be an expression of a transitional state in below-ground succession of niches caused by an O(3) induced effect on carbon allocation to roots and the rhizosphere. We have detected changes in ectomycorrhizal species level, however Shannon-Weavers species diversity index and percentage of types of ectomycorrhizae did not change significantly in any sampling year thus indicating our results cannot be unequivocally explained by summer drought in year 2003 or by O(3) exposure alone.     

Effects of elevated ozone and low light on diurnal and seasonal carbon gain in sugar maple

The long‐term interactive effects of ozone and light on whole‐tree carbon balance of sugar maple (Acer saccharum Marsh.) seedlings were examined, with an emphasis on carbon acquisition, foliar partitioning into starch and soluble sugars, and allocation to growth. Sugar maple seedlings were fumigated with ambient, 1·7 × ambient and 3·0 × ambient ozone in open‐top chambers for 3 years under low and high light (15 and 35% full sunlight, respectively). Three years of ozone fumigation reduced the total biomass of seedlings in the low‐ and high‐light treatments by 64 and 41%, respectively, but had no effect on whole‐plant biomass allocation. Ozone had no effect on net photosynthesis until late in the growing season, with low‐light seedlings generally exhibiting more pronounced reductions in photosynthesis. The late‐season reduction in photosynthesis was not due to impaired stomatal function, but was associated more with accelerated senescence or senescence‐like injury. In contrast, the 3·0 × ambient ozone treatment immediately reduced diurnal starch accumulation in leaves by over 50% and increased partitioning of total non‐structural carbohydrates into soluble sugars, suggesting that injury repair processes may be maintaining photosynthesis in late spring and early summer at the expense of storage carbon. The results in the present study indicate that changes in leaf‐level photosynthesis may not accurately predict the growth response of sugar maple to ozone in different light environments. The larger reduction in seedling growth under low‐light conditions suggests that seedlings in gap or closed‐canopy environments are more susceptible to ozone than those in a clearing. Similarly, understanding the effects of tropospheric ozone on net carbon gain of a mature tree will require scaling of leaf‐level responses to heterogeneous light environments, where some leaves may be more susceptible than others.     

Ultraviolet radiation,ozone depletion,and marine photosynthesis

Concerns about stratospheric ozone depletion have stimulated interest in the effects of UVB radiation (280–320 nm) on marine phytoplankton. Research has shown that phytoplankton photosynthesis can be severely inhibited by surface irradiance and that much of the effect is due to UV radiation. Quantitative generalization of these results requires a biological weighting function (BWF) to quantify UV exposure appropriately. Different methods have been employed to infer the general shape of the BWF for photoinhibition in natural phytoplankton, and recently, detailed BWFs have been determined for phytoplankton cultures and natural samples. Results show that although UVB photons are more damaging than UVA (320–400 nm), the greater fluxes of UVA in the ocean cause more UV inhibition. Models can be used to analyze the sensitivity of water column productivity to UVB and ozone depletion. Assumptions about linearity and time-dependence strongly influence the extrapolation of results. Laboratory measurements suggest that UV inhibition can reach a steady-state consistent with a balance between damage and recovery processes, leading to a non-linear relationship between weighted fluence rate and inhibition. More testing for natural phytoplankton is required, however. The relationship between photoinhibition of photosynthesis and decreases in growth rate is poorly understood, so long-term effects of ozone depletion are hard to predict. However, the wide variety of sensitivities between species suggests that some changes in species composition are likely. Predicted effects of ozone depletion on marine photosynthesis cannot be equated to changes in carbon flux between the atmosphere and ocean. Nonetheless, properly designed studies on the effects of UVB can help identify which physiological and ecological processes are most likely to dominate the responses of marine ecosystems to ozone depletion.Abbreviations BWF biological weighting function - BWF/P-I photosynthesis versus photosynthetically available irradiance as influenced by biologically-weighted UV - Chl chlorophyll a - DOM dissolved organic matter - E _PAR irradiance in energy units (PAR) - E _s saturation parameter for PAR in the BWF/P-I model - E _inh ^* biologically-weighted dimensionless fluence rate for photoinhibition of photosynthesis by UV and PAR - biological weighting coefficient - % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGqiVu0Je9sqqrpepC0xbbL8F4rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaGafqyTduMbae% baaaa!37AC!\[\bar \varepsilon \]_PAR biological weighting coefficient for damage to photosynthesis by E _PAR - k() diffuse attenuation coefficient for wavelength - MAAs mycosporine-like amino acids - PAR photosynthetically available radiation - P ^B rate of photosynthesis normalized to Chl - P _s ^B maximum attainable rate of photosynthesis in the absence of photoinhibition - UVA ultraviolet A (320–400 nm) - UVB ultraviolet B (280–320 nm)     

Beech trees exposed to high CO2 and to simulated summer ozone levels: Effects on photosynthesis, chloroplast components and leaf enzyme activity

Young trees of European beech ( Fagus sylvatica L.) were exposed in a phytotron to different levels of ozone and CO₂ under the climatic simulation of one vegetation period. High ozone levels were simulated similar to high ozone concentration in the field (up to 110 ppb), while CO₂ was added as 300 ppm to the present level of ca 380 ppm. Our study describes different aspects of photosynthesis from the leaf level to the reactions of selected thylakoid components at different harvest times during growth of the beech trees under the different fumigation regimes. Ozone effects appeared in the first weeks of the treatment as a stimulation of chlorophyll fluorescence (F_v/F_m), in oxygen production and in ribulose-1,5-bisphosphate carboxylase/oxygenase activity, while the summer and early autumn harvests showed strong reductions in these parameters. Only phosphoenolpyruvate carboxylase (PEPcase) activity remained higher under high ozone. The effects of high CO₂ appeared in general as a small stimulation in enzyme activity like PEPcase in spring. However, with increasing time of fumigation, reductions of all parameters were observed. Especially chlorophylls showed strong reductions under high CO₂. The combined treatment with high ozone plus high CO₂ resulted mostly in an amelioration of the negative ozone effects, although control levels were not reached.     

The contribution of ozone to forest decline

More than 10 years of intensive research into forest decline in Germany has ascertained that the full extent of the visible damage cannot be explained by just one of the current hypotheses on forest decline. One of the prominent hypotheses is that chronic exposure of forest trees to ozone is probably one of the primary causes of forest decline. The aim of this paper is the critical review of a number of recent results dealing with the ozone hypothesis from a plant physiological point of view. The synopsis focusses on the effects of ozone on conifers because the most extensive data are available for coniferous trees, especially for spruce ( Picea sp.) and pine ( Pinus sp.) trees.     

Extraordinary drought of 2003 overrules ozone impact on adult beech trees (Fagus sylvatica)

The extraordinary drought during the summer of 2003 in Central Europe allowed to examine responses of adult beech trees (Fagus sylvatica) to co-occurring stress by soil moisture deficit and elevated O₃ levels under forest conditions in southern Germany. The study comprised tree exposure to the ambient O₃ regime at the site and to a twice-ambient O₃ regime as released into the canopy through a free-air O₃ fumigation system. Annual courses of photosynthesis (A _max), stomatal conductance (g _s), electron transport rate (ETR) and chlorophyll levels were compared between 2003 and 2004, the latter year representing the humid long-term climate at the site. ETR, A _max and g _s were lowered during 2003 by drought rather than ozone, whereas chlorophyll levels did not differ between the years. Radial stem increment was reduced in 2003 by drought but fully recovered during the subsequent, humid year. Comparison of AOT40, an O₃ exposure-based risk index of O₃ stress, and cumulative ozone uptake (COU) yielded a linear relationship throughout humid growth conditions, but a changing slope during 2003. Our findings support the hypothesis that drought protects plants from O₃ injury by stomatal closure, which restricts O₃ influx into leaves and decouples COU from high external ozone levels. High AOT40 erroneously suggested high O₃ risk under drought. Enhanced ozone levels did not aggravate drought effects in leaves and stem.