Exemplifying whole-plant ozone uptake in adult forest trees of contrasting species and site conditions

Whole-tree O3 uptake was exemplified for Picea abies, Fagus sylvatica and Larix decidua in stands at high and low altitude and contrasting water availability through sap flow measurement in tree trunks, intrinsically accounting for drought and boundary layer effects on O3 flux. O3 uptake of evergreen spruce per unit foliage area was enhanced by 100% at high relative to low elevation, whereas deciduous beech and larch showed similar uptake regardless of altitude. The responsiveness of the canopy conductance to water vapor and, as a consequence, O3 uptake to soil moisture and air humidity did not differ between species. Unifying findings at the whole-tree level will promote cause-effect based O3 risk assessment and modeling.     

Response of Brazilian native trees to acute ozone dose

Ozone (O₃) is a toxic secondary pollutant able to cause an intense oxidative stress that induces visual symptoms on sensitive plant species. Controlled fumigation experiment was conducted with the aim to verify the O₃ sensibility of three tropical species: Piptadenia gonoachanta (Mart.) Macbr. (Fabaceae), Astronium graveolens Jacq. (Anacardiaceae), and Croton floribundus Spreng. (Euphorbiaceae). The microscopical features involved in the oxidative stress were recognized based on specific histochemical analysis. The three species showed visual symptoms, characterized as necrosis and stippling between the veins, mostly visible on the adaxial leaf surface. All the studied species presented hypersensitive-like response (HR-like), and peroxide hydrogen accumulation (H₂O₂) followed by cell death and proanthocyanidin oxidation in P. gonoachanta and A. graveolens. In P. gonoachanta, a decrease in chlorophyll autofluorescence occurred on symptomatic tissues, and in A. graveolens and C. floribundus, a polyphenol compound accumulation occurred. The responses of Brazilian native species were similar to those described for sensitive species from temperate climate, and microscopical markers may be useful for the detection of ozone symptoms in future studies in the field.     

Deriving ozone dose-response of photosynthesis in adult forest trees from branch-level cuvette gas exchange assessment

Branch-level gas exchange provided the basis for assessing ozone flux in order to derive the dose-response relationship between cumulative O3 uptake (COU) and carbon gain in the upper sun crown of adult Fagus sylvatica. Fluxes of ozone, CO2 and water vapour were monitored simultaneously by climatized branch cuvettes. The cuvettes allowed branch exposure to an ambient or twice-ambient O3 regime, while tree crowns were exposed to the same O3 regimes (twice-ambient generated by a free-air canopy O3 exposure system). COU levels higher than 20mmolm(-2) led to a pronounced decline in carbon gain under elevated O3. The limiting COU range is consistent with findings on neighbouring branches exposed to twice-ambient O3 through free-air fumigation. The cuvette approach allows to estimate O3 flux at peripheral crown positions, where boundary layers are low, yielding a meso-scale within-crown resolution of photosynthetic foliage sensitivity under whole-tree free-air O3 fumigation.     

The challenge of making ozone risk assessment for forest trees more mechanistic

Upcoming decades will experience increasing atmospheric CO₂ and likely enhanced O₃ exposure which represents a risk for the carbon sink strength of forests, so that the need for cause-effect related O₃ risk assessment increases. Although assessment will gain in reliability on an O₃ uptake basis, risk is co-determined by the effective dose, i.e. the plant's sensitivity per O₃ uptake. Recent progress in research on the molecular and metabolic control of the effective O₃ dose is reported along with advances in empirically assessing O₃ uptake at the whole-tree and stand level. Knowledge on both O₃ uptake and effective dose (measures of stress avoidance and tolerance, respectively) needs to be understood mechanistically and linked as a pre-requisite before practical use of process-based O₃ risk assessment can be implemented. To this end, perspectives are derived for validating and promoting new O₃ flux-based modelling tools.     

Advances in understanding ozone impact on forest trees: Messages from novel phytotron and free-air fumigation studies

Recent evidence from novel phytotron and free-air ozone (O₃) fumigation experiments in Europe and America on forest tree species is highlighted in relation to previous chamber studies. Differences in O₃ sensitivity between pioneer and climax species are examined and viewed for trees growing at the harsh alpine timberline ecotone. As O₃ apparently counteracts positive effects of elevated CO₂ and mitigates productivity increases, response is governed by genotype, competitors, and ontogeny rather than species per se. Complexity in O₃ responsiveness increased under the influence of pathogens and herbivores. The new evidence does not conflict in principle with previous findings that, however, pointed to a low ecological significance. This new knowledge on trees' O₃ responsiveness beyond the juvenile stage in plantations and forests nevertheless implies limited predictability due to complexity in biotic and abiotic interactions. Unravelling underlying mechanisms is mandatory for assessing O₃ risks as an important component of climate change scenarios.     

Mapping of exceedances of ozone critical levels for crops and forest trees in The Netherlands: Preliminary results

Within the framework of the UN-ECE Convention on Long-Range Transboundary Air Pollution (CLRTAP) critical levels of ozone for forest trees and crops have been determined. In this paper procedures are presented for constructing maps of The Netherlands showing measured ozone concentrations for comparison with the critical levels. The critical ozone levels for forest trees and crops are exceeded widely and frequently in The Netherlands. The frequency and size of exceedances depend on meteorological conditions: during a hot, sunny summer like 1989 or 1990 exceedances of the critical levels by a factor of two or three were observed in the whole of The Netherlands; in a cold, rainy summer like 1991 or 1993 only minor exceedances were observed in some parts (mainly in the south) of the country.     

Risk assessments for forest trees: the performance of the ozone flux versus the AOT concepts

Published ozone exposure-response relationships from experimental studies with young trees performed at different sites across Europe were re-analysed in order to test the performance of ozone exposure indices based on AOTX (Accumulated exposure Over a Threshold of X nmol mol(-1)) and AF(st)Y (Accumulated Stomatal Flux above a threshold of Y nmol m(-2) s(-1)). AF(st)1.6 was superior, as compared to AOT40, for explaining biomass reductions, when ozone sensitive species with differing leaf morphology were included in the analysis, while this was not the case for less sensitive species. A re-analysis of data with young black cherry trees, subject to different irrigation regimes, indicated that leaf visible injuries were more strongly related to the estimated stomatal ozone uptake, as compared to the ozone concentration in the air. Experimental data with different clones of silver birch indicated that leaf thickness was also an important factor influencing the development of ozone induced leaf visible injury.     

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.     

Effects of chronic elevated ozone exposure on gas exchange responses of adult beech trees (Fagus sylvatica) as related to the within-canopy light gradient

The effects of elevated O₃ on photosynthetic properties in adult beech trees (Fagus sylvatica) were investigated in relation to leaf mass per area as a measure of the gradually changing, within-canopy light availability. Leaves under elevated O₃ showed decreased stomatal conductance at unchanged carboxylation capacity of Rubisco, which was consistent with enhanced δ¹³C of leaf organic matter, regardless of the light environment during growth. In parallel, increased energy demand for O₃ detoxification and repair was suggested under elevated O₃ owing to enhanced dark respiration. Only in shade-grown leaves, light-limited photosynthesis was reduced under elevated O₃, this effect being accompanied by lowered F_v/F_m. These results suggest that chronic O₃ exposure primarily caused stomatal closure to adult beech trees in the field regardless of the within-canopy light gradient. However, light limitation apparently raised the O₃ sensitivity of photosynthesis and accelerated senescence in shade leaves.     

Understanding consistencies and gaps between desired forest futures: An analysis of visions from stakeholder groups in Sweden

Conflicting perspectives on forests has for a long time challenged forest policy development in Sweden. Disagreements about forest futures create intractable deadlocks when stakeholders talk past each other. The purpose of this study is to move beyond this situation through the application of participatory backcasting. By comparing visions of the future forest among stakeholder groups, we highlight contemporary trajectories and identify changes that were conceived as desirable. We worked with four groups: the Biomass and Bioenergy group, the Conservation group, the Sami Livelihood group and the Recreation and Rural Development group; in total representatives from 40 organizations participated in workshops articulating the groups’ visions. Our results show well-known tensions such as intrinsic versus instrumental values but also new ones concerning forests’ social values. Identified synergies include prioritization of rural development, new valued-added forest products and diversified forest management. The results may feed directly into forest policy processes facilitating the process and break current deadlocks.     

Assessing the risk caused by ground level ozone to European forest trees: a case study in pine, beech and oak across different climate regions

Two different indices have been proposed for estimation of the risk caused to forest trees across Europe by ground-level ozone, (i) the concentration based AOT40 index (Accumulated Over a Threshold of 40 ppb) and (ii) the recently developed flux based AFstY index (Accumulated stomatal Flux above a flux threshold Y). This paper compares the AOT40 and AFstY indices for three forest trees species at different locations in Europe. The AFstY index is estimated using the DO(3)SE (Deposition of Ozone and Stomatal Exchange) model parameterized for Scots pine (Pinus sylvestris), beech (Fagus sylvatica) and holm oak (Quercus ilex). The results show a large difference in the perceived O(3) risk when using AOT40 and AFstY indices both between species and regions. The AOT40 index shows a strong north-south gradient across Europe, whereas there is little difference between regions in the modelled values of AFstY. There are significant differences in modelled AFstY between species, which are predominantly determined by differences in the timing and length of the growing season, the periods during which soil moisture deficit limits stomatal conductance, and adaptation to soil moisture stress. This emphasizes the importance of defining species-specific flux response variables to obtain a more accurate quantification of O(3) risk.     

Physiological and foliar symptom response in the crowns of Prunus serotina, Fraxinus americana and Acer rubrum canopy trees to ambient ozone under forest conditions

The crowns of five canopy dominant black cherry (Prunus serotina Ehrh.), five white ash (Fraxinus americana L.), and six red maple (Acer rubrum L.) trees on naturally differing environmental conditions were accessed with scaffold towers within a mixed hardwood forest stand in central Pennsylvania. Ambient ozone concentrations, meteorological parameters, leaf gas exchange and leaf water potential were measured at the sites during the growing seasons of 1998 and 1999. Visible ozone-induced foliar injury was assessed on leaves within the upper and lower crown branches of each tree. Ambient ozone exposures were sufficient to induce typical symptoms on cherry (0-5% total affected leaf area, LAA), whereas foliar injury was not observed on ash or maple. There was a positive correlation between increasing cumulative ozone uptake (U) and increasing percent of LAA for cherry grown under drier site conditions. The lower crown leaves of cherry showed more severe foliar injury than the upper crown leaves. No significant differences in predawn leaf water potential (psi(L)) were detected for all three species indicating no differing soil moisture conditions across the sites. Significant variation in stomatal conductance for water vapor (g(wv)) was found among species, soil moisture, time of day and sample date. When comparing cumulative ozone uptake and decreased photosynthetic activity (P(n)), red maple was the only species to show higher gas exchange under mesic vs. drier soil conditions (P < 0.05). The inconsistent differences in gas exchange response within the same crowns of ash and the uncoupling relationship between g(wv) and P(n) demonstrate the strong influence of heterogeneous environmental conditions within forest canopies.     

Determining temporal and spatial variations in ozone air quality

Unless the change in emissions is substantial, the resulting improvement in ozone air quality can be easily masked by the meteorological variability. Therefore, the meteorological and chemical signals must be separated in examining ozone trends. In this paper, we discuss the use of the Kolmogorov-Zurbenko filter in evaluating the temporal and spatial variations in ozone air quality utilizing ozone concentration data from several monitoring locations in the northeastern United States. The results indicate a downward trend in the ozone concentrations during the period 1983-1992 at most locations in the northeastern United States. The results also reveal that ozone is a regional-scale problem in the Northeast.     

Limitations and perspectives about scaling ozone impacts in trees.

We review the need for scaling effects of ozone (O3) from juvenile to mature forest trees, identify the knowledge presently available, and discuss limitations in scaling efforts. Recent findings on O3/soil nutrient and O3/CO2 interactions from controlled experiments suggest consistent scaling patterns for physiological responses of individual leaves to whole-plant growth, carbon allocation, and water use efficiency of juvenile trees. These findings on juvenile trees are used to develop hypotheses that are relevant to scaling O3 effects to mature trees, and these hypotheses are examined with respect to existing research on differences in response to O3 between juvenile and mature trees. Scaling patterns of leaf-level physiological response to O3 have not been consistent in previous comparisons between juvenile and mature trees. We review and synthesize current understanding of factors that may cause such inconsistent scaling patterns, including tree-size related changes in environment, stomatal conductance, O3 uptake and exposure. carbon allocation to defense, repair, and compensation mechanisms, and leaf production phenology. These factors should be considered in efforts to scale O3 responses during tree ontogeny. Free-air O3 fumigation experiments of forest canopies allow direct assessments of O3 impacts on physiological processes of mature trees, and provide the opportunity to test current hypotheses about ontogenetic variation in O3 sensitivity by comparing O3 responses across tree-internal scales and ontogeny.     

Seasonal differences and within-canopy variations of antioxidants in mature spruce (Picea abies) trees under elevated ozone in a free-air exposure system

The effect of free-air ozone fumigation and crown position on antioxidants were determined in old-growth spruce (Picea abies) trees in the seasonal course of two consecutive years (2003 and 2004). Levels of total ascorbate and its redox state in the apoplastic washing fluid (AWF) were increased under double ambient ozone concentrations (2xO3), whilst ascorbate concentrations in needle extracts were unchanged. Concentrations of apoplastic and symplastic ascorbate were significantly higher in 2003 compared to 2004 indicating a combined effect of the drought conditions in 2003 with enhanced ozone exposure. Elevated ozone had only weak effects on total glutathione levels in needle extracts, phloem exudates and xylem saps. Total and oxidised glutathione concentrations were higher in 2004 compared to 2003 and seemed to be more affected by enhanced ozone influx in the more humid year 2004 compared to the combined effect of elevated ozone and drought in 2003 as observed for ascorbate.     

Ozone uptake by citrus trees exposed to a range of ozone concentrations

The Citrus genus includes a large number of species and varieties widely cultivated in the Central Valley of California and in many other countries having similar Mediterranean climates. In the summer, orchards in California experience high levels of tropospheric ozone, formed by reactions of volatile organic compounds (VOC) with oxides of nitrogen (NO_x). Citrus trees may improve air quality in the orchard environment by taking up ozone through stomatal and non-stomatal mechanisms, but they may ultimately be detrimental to regional air quality by emitting biogenic VOC (BVOC) that oxidize to form ozone and secondary organic aerosol downwind of the site of emission. BVOC also play a key role in removing ozone through gas-phase chemical reactions in the intercellular spaces of the leaves and in ambient air outside the plants. Ozone is known to oxidize leaf tissues after entering stomata, resulting in decreased carbon assimilation and crop yield. To characterize ozone deposition and BVOC emissions for lemon (Citrus limon), mandarin (Citrus reticulata), and orange (Citrus sinensis), we designed branch enclosures that allowed direct measurement of fluxes under different physiological conditions in a controlled greenhouse environment. Average ozone uptake was up to 11 nmol s^?1 m^?2 of leaf. At low concentrations of ozone (40 ppb), measured ozone deposition was higher than expected ozone deposition modeled on the basis of stomatal aperture and ozone concentration. Our results were in better agreement with modeled values when we included non-stomatal ozone loss by reaction with gas-phase BVOC emitted from the citrus plants. At high ozone concentrations (160 ppb), the measured ozone deposition was lower than modeled, and we speculate that this indicates ozone accumulation in the leaf mesophyll.     

Spatial patterns concentrations in upland Wales in relation to catchment forest cover and forest age

Data on nitrate nitrogen were collected weekly during 1984 from 136 sites on streams in upland Wales. Mean nitrate concentrations in summer (0.02 to 1.5 mg litre(-1)) were significantly lower (P<0.001) than in winter (0.02 to 1.26 mg litre(-1)), particularly at sites with mature conifers (>30 years old). Mean concentrations increased significantly with the average age of conifers on each catchment (P<0.001), and with increasing areal cover by trees over 30 years old (P<0.001). Nitrate concentrations increased significantly with stream total hardness (P<0.001), possibly reflecting nitrogen mineralisation in soils of higher base status. Concentrations also increased with stream chloride (P<0.001), which is predominantly atmospherically derived, implying that increased nitrate occurred where general atmospheric inputs of solutes were increased. After accounting for variation in hardness, residual nitrate concentrations still increased with the average age of the conifers (P<0.001), and with catchment cover by mature trees (P<0.001). We infer that some additional nitrate under older conifers is thus independent of catchment sources associated with increasing hardness. Two possibilities are increased inputs and decreased retention of nitrogen within the ecosystem of maturing conifer forest. Residual nitrate after accounting for variations in chloride also increased significantly with conifer age (P<0.01) and cover (P<0.01), a pattern implying that some sources of nitrate may also be independent of increased sea-salt deposition. We allude to the possibility that additional nitrogen deposition adds to nitrogen throughputs from maturing forests, and we discuss the potential ecological role of additional nitrogen in runoff.     

Summer and autumn ozone fluxes to a forest in the Czech Republic Brdy Mountains

In an effort to examine ozone (O3) deposition over a forest site in the Czech Republic, a low cost eddy flux experiment using slow response ozone and temperature sensors was implemented in July 1993 within the Brdy Mountains. Half-hour 2-Hz ozone and sensible heat measurements made at the Brdy Mountains for 98 days during the period 7 July 1994-20 October 1994 are analyzed and reported. While the Czech Brdy Mountains AOT40 level for the overall 104 day period was 7.6 ppm h (15.1 ppm h for the full 24-h summation), indicating a slight potential for 03 injury, the 1994 summer to autumn'measured forest O3 uptake was 2.4 (+/- 0.9) g m(-2), not unusually high compared to other studies. Average summer midday 03 fluxes and depositidn velocities were -1.0 (+/- 0.6) microg m(-2) s(-1) and 1.1 (+/- 0.7) cm s(-1). and autumn values were -0.36 (+/- 0.4) microg m(-2) s(-1) and 0.7 (+/- 0.5) cm s(-1) respectively. A unique contribution of this study is the first time demonstrated use of slow responding sensors for eddy covariance flux measurements at heights of 20 m above a forest.     

A theoretical investigation of the pressure and temperature dependence of atmospheric ozone deposition to trees

Methods for describing the exposure patterns of forests to atmospheric ozone concentrations are compared with special emphasis on the situation at high altitudes, such as the Appalachian Mountains of the eastern USA. Limitations to the use of ozone concentration as mass per unit volume are discussed and a correction for temperature and pressure changes is derived. If identical ozone mass concentrations were measured at two sites separated by 2000 m elevation, the ozone flux at the lower site would exceed the flux at the higher site by 4-8% due to temperature and pressure effects on both air volume and ozone deposition velocity. It is recommended that ozone exposures be described in terms of 'flux-corrected' mass concentrations or volumetric mixing ratios when ambient ozone data from sites at different altitudes are to be compared.