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.     

Seasonal ozone response of mature beech trees (Fagus sylvatica) at high altitude in the Bavarian forest (Germany) in comparison with young beech grown in the field and in phytotrons

Mature beech trees (Fagus sylvatica) grown at two different altitudes in the Bavarian forest were compared with young beech trees grown at nearby field sites or in phytotrons for their macroscopic and physiological responses to different ozone (O(3)) exposures. Cumulative O(3) exposure expressed as the sum of hourly mean concentrations above the canopy ranged between 100 and 150 microl l(-1) h, with the vertical O(3) profiles at the higher altitude site being enhanced by 30%. O(3) profiles at all sites were reduced by up to 20% with increasing depth within and beneath the canopy. The leaf discoloration that developed in the absence of premature leaf loss was similar in the sun foliage of mature and young trees (including plant grown in the phytotron). Injury became apparent at low O(3) exposures, expressed as accumulated hourly means over a threshold of 40 nl l(-1) (AOT40 <3.5 microl l(-1) h) at the lower site in both the mature trees and the young beech at the field site, but only occurred when AOT40 values reached 7 microl l(-1) h at the upper site, and 6 microl l(-1) h in the phytotrons. However, the association between injury and O(3) exposure was improved when cumulative ozone uptake to sun leaves was the ozone index, used with values of about 3 mmol m(-2) resulting in visible injury in both mature and young beech growing in phytotrons. Under high ozone exposure levels of inositol were lowered, whilst concentrations of lignin-like materials were enhanced in mature beech. Similar responses were observed in young beech grown in phytotrons. As the sun foliage was affected by only a small and variable extent each year, the seasonal O(3) impact at high altitude did not appear to pose an acute risk to mature beech trees.     

Vegetation of the selected forest stands and land use in the Carpathian Mountains

Within the framework of the project "Effects of forest health on biodiversity with emphasis on air pollution in the Carpathian Mountains" 26 permanent study sites were established in the vicinity of the ozone monitoring sites. The study sites were located on the NW-SE transect through the Western (12 sites), Eastern (11 sites) and Southern (3 sites) Carpathians in forest ecosystems typical of each area. Some of the forest monitoring sites were located in national parks, biosphere reserves and areas of protected landscape. Each permanent site of 0.7 ha area consisted of 5 small 500m(2) circular plots, arranged in the form of a cross, i.e. four placed on the cardinal points (N, E, S, W) and one in the center. Phytosociological records were done twice during the 1998 growing season using the Braun-Blanquet's method. The study sites represented various types of forest: Picea abies stands (8), beech (Fagus sylvatica) stands (10), fir (Abies alba) stands (2) and mixed beech-fir, spruce-fir and beech-spruce stands (6). Age of most stands was 80-100 years. Degree of crown damage varied greatly between sites, a percentage of damaged trees decrease in Carpathians from West to East. It corresponds well with the O(3) level in these areas. Typical damage by O(3) in herb layer species in several Carpathian sites were found. Land-use map for the entire Carpathian Mountains and two detailed land use maps for Tatras (Western Carpathians) and Retezat (Southern Carpathians) are presented. A little more than half of the Carpathian territory is forested. The most densely forested are Eastern Carpathians, while the most sparsely Western Carpathians. Arable lands occupy 22.6% of the Carpathians, pastures and meadows 6.2%, water bodies 1.9%, and build up areas several percent. In the highest elevation of the Carpathians alpine meadows (11.3%) and rocks (3.5%) are distributed.     

Performance of some growth variables

European beech (Fagus sylvatica L.), Norway spruce (Picea abies L. Karst.) and Silver fir (Abies alba Mill.) were exposed to low concentrations of ozone (O(3)) and sulfur dioxide (SO(2)), alone and combined, and simulated acid rain (pH 4.0) in sheltered open-top chambers in Hohenheim (Southwest Germany) for almost five years. The concentrations of O(3) and SO(2) used were related to annual ambient average found in southern West Germany. Two control chambers were ventilated with charcoal filtered air and rainfall was simulated at pH 4.0 and 5.0. Because of large dense plant growth in the chambers it was only possible to measure uncompleted growth of shoots in the upper canopy. Therefore, growth analysis was restricted to this area. The treatment with acidic precipitation decreased the annual shoot growth of beech and reduced leaf surface area of those trees. Exposure to SO(2), O(3) alone and in combination resulted in further reduction of shoot length and leaf surface area. Fumigation with SO(2) and O(3) + SO(2) caused insignificant decreases of shoot length, total dry weight and needle surface area of spruce. The lateral leader shoot growth of spruce exposed to O(3) was significantly reduced only in the last year of the experiment. Growth rates of the spruce exposed to charcoal filtered air and non-acidic precipitation were reduced more than those of beech and fir. Growth variables determined for fir reflected different rates of incremental change. Exposure to O(3) resulted in the largest dry matter production of all fir groups but those exposed to charcoal filtered air and non-acidic precipitation responded with the best lateral leader shoot growth, lowest specific leaf area (SLA) and leaf area ratio (LAR) respectively indicating best metabolic efficiency. At the conclusion of this study a classification of sensitivity was developed for the tree species.     

Tree and stand growth of mature Norway spruce and European beech under long-term ozone fumigation

In a 50- to 70-year-old mixed stand of Norway spruce (Picea abies (L.) Karst.) and European beech (Fagus sylvatica L.) in Germany, tree cohorts have been exposed to double ambient ozone (2×O₃) from 2000 through 2007 and can be compared with trees in the same stand under the ambient ozone regime (1×O₃). Annual diameter growth, allocation pattern, stem form, and stem volume were quantified at the individual tree and stand level. Ozone fumigation induced a shift in the resource allocation into height growth at the expense of diameter growth. This change in allometry leads to rather cone-shaped stem forms and reduced stem stability in the case of spruce, and even neiloidal stem shapes in the case of beech. Neglect of such ozone-induced changes in stem shape may lead to a flawed estimation of volume growth. On the stand level, 2×O₃ caused, on average, a decrease of 10.2 m³ ha⁻¹ yr⁻¹ in European beech.     

Evidence from nitrogen fertilisation in the forests of Germany

In the first part, this contribution presents German results of nitrogen fertilisation experiments on stand growth, published 1958-1987. Over this period Norway spruce and Scots pine have generally responded positively to nitrogen fertilisation, without showing signs of damage, at levels up to at least 1000 kg N ha(-1). In the second part, growth patterns of forest trees and stands in Southern Germany are presented. Norway spruce in particular is now growing better than in earlier years of this century, starting around 1960. It is plausible that increasing nitrogen deposition, which coincides with the growth increase, is the cause, though this cannot be shown unequivocally. At a few sites with poor soil and management and high acid deposition a deterioration is taking place.     

Ozone exposure-response relationships for biomass and root/shoot ratio of beech (Fagus sylvatica), ash (Fraxinus excelsior), Norway spruce (Picea abies) and Scots pine (Pinus sylvestris)

Current-year seedlings of beech, ash, Norway spruce and Scots pine were exposed during one growing season to different, but moderate, ozone (O(3)) scenarios representative for Switzerland (50, 85, 100% ambient, 50% ambient+30 nl l(-1)) in open-top chambers (OTCs) and to ambient O(3) concentrations in the field. Biomass significantly decreased with increasing O(3) dose in all species except for spruce. Losses of 25.5% (ash), 17.4% (beech), 9.9% (Scots pine) were found per 10 microl l(-1) h accumulated O(3) exposure over a threshold concentration of 40 nl l(-1) during daylight hours (AOT40). Ratios of root/shoot biomass (RSR) also significantly decreased with increasing AOT40 levels in beech and ash, but not in Norway spruce and Scots pine. The data show that the deciduous species beech and ash were more susceptible to O(3) with respect to RSR and biomass than the coniferous species Norway spruce and Scots pine.     

Use of sap flow measurements to validate stomatal functions for mature beech (Fagus sylvatica) in view of ozone uptake calculations

For a quantitative estimate of the ozone effect on vegetation reliable models for ozone uptake through the stomata are needed. Because of the analogy of ozone uptake and transpiration it is possible to utilize measurements of water loss such as sap flow for quantification of ozone uptake. This technique was applied in three beech (Fagus sylvatica) stands in Switzerland. A canopy conductance was calculated from sap flow velocity and normalized to values between 0 and 1. It represents mainly stomatal conductance as the boundary layer resistance in forests is usually small. Based on this relative conductance, stomatal functions to describe the dependence on light, temperature, vapour pressure deficit and soil moisture were derived using multivariate nonlinear regression. These functions were validated by comparison with conductance values directly estimated from sap flow. The results corroborate the current flux parameterization for beech used in the DO₃SE model.     

Comparison of ozone uptake and sensitivity between a phytotron study with young beech and a field experiment with adult beech (Fagus sylvatica)

Chamber experiments on juvenile trees have resulted in severe injury and accelerated loss of leaves along with reduced biomass production under chronically enhanced O3 levels. In contrast, the few studies conducted on adult forest trees in the field have reported low O3 sensitivity. In the present study, young beech in phytotrons was more sensitive to O3 than adult beech in the field, although employed O3 regimes were similar. The hypotheses tested were that: (1) differences in O3 uptake were caused by the ontogenetically higher stomatal conductance of young compared to adult trees, (2) the experimental settings in the phytotrons enhanced O3 uptake compared to field conditions, and (3) a low detoxification capacity contributes to the higher O3 sensitivity of the young trees. The higher O3 sensitivity of juvenile beech in the phytotrons is demonstrated to relate to both the experimental conditions and the physiological responsiveness inherent to tree age.     

Impacts of ozone on the growth and yield of field-grown winter wheat

Seed of winter wheat (Triticum aestivum L. cv Riband) was sown on 29 August 1992 in eight field plots. Four plots were exposed to elevated ozone (O(3)) concentrations on 16 days between 29 August and 2 October 1992, for 6 h day(-1), and on 27 days between 29 March and 24 August 1993, for 7 h day(-1). Mean daily O(3) concentrations were approximately 30 and 80 nmol mol(-1) in ambient and fumigated plots, respectively. Plants were sampled on 5 November (1992), 14 January, 16 February, 1 April, 25 May, 23 June and 24 August (1993). No visible symptoms of O(3) damage or premature senescence were observed at any time over the course of the experiment. Exposure to elevated O(3) decreased the above ground biomass by reducing plant density and individual plant relative growth rate. However, there was no significant influence of the pollutant on the growth of the root relative to the shoot. Assessment of yield characteristics at the final harvest revealed an O(3)-induced decrease in the number of grains per ear, as a result of fewer grains per spikelet and an increase in the number of infertile florets per spikelet. No significant effects of the pollutant on the number of ears per plant, spikelets per ear, or 1000 grain weight were found. As a result of the combined effects on the number of grains per ear and the decrease in plant density and growth rate, O(3) exposure reduced grain and straw yields (tonnes ha(-1)) by 13 and 8%, respectively. However, no significant change in the partitioning of dry matter between the grain and the straw was observed in fumigated plots. The findings are discussed within the context of United Nation Economic Commission for Europe critical level guidelines for the protection of crop yields, in relation to their application to winter-sown crops.     

Air pollution, precipitation chemistry and forest health in the Retezat Mountains, Southern Carpathians, Romania

In the Retezat Mountains concentrations of O3, NO2 and SO2 in summer season 2000-2002 were low and below toxicity levels for forest trees. While NH3 concentrations were low in 2000, the 2001 and 2002 concentrations were elevated indicating possibility for increased N deposition to forest stands. More than 90% of the rain events were acidic with pH values <5.5, contributing to increased acidity of soils. Crown condition of Norway spruce (Picea abies) and European beech (Fagus sylvatica) was good, however, defoliation described as >25% of foliage injured increased from 9.1% in 2000 to 16.1% in 2002. Drought that occurred in the southern Carpathians between fall 2000 and summer 2002 and frequent acidic rainfalls could cause the observed decline of forest condition. Both Norway spruce and European beech with higher defoliation had lower annual radial increments compared to the trees with low defoliation. Ambient O3 levels found in the Retezat did not affect crown condition of Norway spruce or European beech.     

A statistical approach to estimate O3 uptake of ponderosa pine in a mediterranean climate

In highly polluted sites, stomatal behavior is sluggish with respect to light, vapor pressure deficit, and internal CO2 concentration (Ci) and poorly described by existing models. Statistical models were developed to estimate stomatal conductance (gs) of 40-year-old ponderosa pine at three sites differing in pollutant exposure for the purpose of calculating O3 uptake. Gs was estimated using julian day, hour of day, pre-dawn xylem potential and photosynthetic photon flux density (PPFD). The median difference between estimated and observed field gs did not exceed 10 mmol H2O m(-2) s(-1), and estimated gs within 95% confidence intervals. 03 uptake was calculated from hourly estimated gs, hourly O3 concentration, and a constant to correct for the difference in diffusivity between water vapor and 03. The simulation model TREGRO was also used to calculate the cumulative 03 uptake at all three sites. 03 uptake estimated by the statistical model was higher than that simulated by TREGRO because gas exchange rates were proportionally higher. O3 exposure and uptake were significantly correlated (r2>0.92), because O3 exposure and gs were highly correlated in both statistical and simulation models.     

Growth responses of Populus tremuloides clones to interacting elevated carbon dioxide and tropospheric ozone.

The Intergovernmental Panel of Climate Change (IPCC) has concluded that the greenhouse gases carbon dioxide (CO2) and tropospheric ozone (O3) are increasing concomitantly globally. Little is known about the effect of these interacting gases on growth, survival, and productivity of forest ecosystems. In this study we assess the effects of three successive years of exposure to combinations of elevated CO2 and O3 on growth responses in a five trembling aspen (Populus tremuloides) clonal mixture in a regenerating stand. The experiment is located in Rhinelander, Wisconsin, USA (45 degrees N 89 degrees W) and employs free air carbon dioxide and ozone enrichment (FACE) technology. The aspen stand was exposed to a factorial combination of four treatments consisting of elevated CO2 (560 ppm), elevated O3 (episodic exposure-90 microl l(-1) hour(-1)), a combination of elevated CO2 and O3, and ambient control in 30 m treatment rings with three replications. Our overall results showed that our three growth parameters including height, diameter and volume were increased by elevated CO2, decreased by elevated O3, and were not significantly different from the ambient control under elevated CO2 + O3. However, there were significant clonal differences in the responses; all five clones exhibited increased growth with elevated CO2, one clone showed an increase with elevated O3, and two clones showed an increase over the control with elevated CO2 + O3, two clones showed a decrease, and one was not significantly different from the control. Notably. there was a significant increase in current terminal shoot dieback with elevated CO2 during the 1999-2000 dormant season. Dieback was especially prominent in two of the five clones, and was attributed to those clones growing longer into the autumnal season where they were subject to frost. Our results show that elevated O3 negates expected positive growth effects of elevated CO2 in Populus tremuloides in the field, and suggest that future climate model predictions should take into account the offsetting effects of elevated O3 on CO2 enrichment when estimating future growth of trembling aspen stands.     

Response patterns in adult forest trees to chronic ozone stress: identification of variations and consistencies

The responsiveness of adult beech and spruce trees to chronic O(3) stress was studied at a free-air O(3) exposure experiment in Freising/Germany. Over three growing seasons, gas exchange characteristics, biochemical parameters, macroscopic O(3) injury and the phenology of leaf organs were investigated, along with assessments of branch and stem growth as indications of tree performance. To assess response pattern to chronic O(3) stress in adult forest trees, we introduce a new evaluation approach, which provides a comprehensive, readily accomplishable overview across several tree-internal scaling levels, different canopy regions and growing seasons. This new approach, based on a three-grade colour coding, combines statistical analysis and the proficient ability of the "human eye" in pattern recognition.     

Ozone effects on growth and assimilate partitioning in alfalfa, Medicago sativa L

Alfalfa (Medicago sativa L.) were exposed to O(3) concentrations varying between 118 x 10(-6) microg cm(-3) (0.06 ppm) and 157 x 10(-6) microg cm(-3) (0.08 ppm) for 6 h per day 5 days per week for several weeks. Typical plants were sacrificed weekly, and growth parameters were measured. O(3) reduced overall growth, relative growth rates and unit leaf rates in alfalfa before it was cut, indicating that O(3) had reduced photosynthesis. However, after the alfalfa was cut, these same parameters indicated that in some cases, O(3)-stressed plants had greater photosynthetic rates than controls during regrowth. O(3) also altered dry matter partitioning. Roots were most affected, followed by leaves and stems, respectively. In general, O(3) reduced photosynthate production and reduced the proportion of photosynthate partitioned to roots relative to leaves and stems. This could reduce starch reserves in alfalfa, and be detrimental to stand longevity. However, the post-cutting study indicated that at least some alfalfa cultivars may be able to acclimate to O(3)-stress, though plants did not fully recover from pre-cutting differences.     

From critical levels to critical loads for ozone: a discussion of a new experimental and modelling approach for establishing flux-response relationships for agricultural crops and native plant species

Present critical levels for ozone (O3) for protecting vegetation against adverse effects are based on exposure-response relationships mainly derived from open-top chamber experiments and are expressed as an Accumulated exposure Over a Threshold of 40 ppb (AOT40). In that context with a revision of the UN (United Nations)-ECE (Economic Commission for Europe) Gothenburg protocol, AOT40 values should be replaced by flux-oriented quantities, i.e. in the end by critical loads. At present, the database for the derivation of critical loads for O3 is extremely inadequate. Furthermore, the currently available flux-response relationships are also derived from open-top chamber experiments. The use of a relationship for spring wheat in a risk assessment for an agricultural site in Hesse, Germany, demonstrates in principle, the applicability of the critical load concept for O3. Comparisons of diurnal variation of stomatal uptake and AOT40 showed that a major part of toxicologically effective stomatal uptake occurred before noon whereas the AOT40 values were dominated by the O3 concentrations during afternoon. In other words, the AOT40 exposure index do not adequately address the O3 burden during hours when plants are sensitive to O3 uptake. However, due to the differences in radiation, air temperature and humidity between the chamber and the ambient microclimates, a derivation of flux-response relationships from chamber experiments is likely to be questionable, especially for species rich ecosystems: Here, without any changes in the pollution climate, significant modifications of species composition as well as an earlier beginning of the growing season has been previously observed. To overcome the problems associated with chamber-derived flux-response relationships, a new experimental and modelling concept, was developed. The approach, briefly described in this paper, combines methods in air pollution toxicology and micrometeorology. As an analogy to the free-air fumigation concept, O3 is released into the air by an injection system above the plant canopy. The assessment of dispersion and surface deposition of O3 released is based on Lagrangian trajectory modelling. Depending on wind direction and velocity, atmospheric stratification and surface roughness, without any disturbance of the microclimate and micrometeorology, several sub-areas can be identified around the source position with differing deposition rates above the ambient level. Taking into account the actual O3 background deposition, deposition rates and vegetation responses observed in these sub-areas can easily be used to derive flux-effect relationships under ambient conditions and more realistic limiting values to protect our environment.     

New exposure-based metric approach for evaluating O(3) risk to North American aspen forests

The United States and Canada currently use exposure-based metrics to protect vegetation from O(3). Using 5 years (1999-2003) of co-measured O(3), meteorology and growth response, we have developed exposure-based regression models that predict Populus tremuloides growth change within the North American ambient air quality context. The models comprised growing season fourth-highest daily maximum 8-h average O(3) concentration, growing degree days, and wind speed. They had high statistical significance, high goodness of fit, include 95% confidence intervals for tree growth change, and are simple to use. Averaged across a wide range of clonal sensitivity, historical 2001-2003 growth change over most of the 26 Mha P. tremuloides distribution was estimated to have ranged from no impact (0%) to strong negative impacts (-31%). With four aspen clones responding negatively (one responded positively) to O(3), the growing season fourth-highest daily maximum 8-h average O(3) concentration performed much better than growing season SUM06, AOT40 or maximum 1h average O(3) concentration metrics as a single indicator of aspen stem cross-sectional area growth.     

Role of climate, crown position, tree age and altitude in calculated ozone flux into needles of Picea abies and Pinus cembra: a synthesis

Ozone (O(3)) flux into Norway spruce (Picea abies) and cembran pine (Pinus cembra) needles was estimated under ambient conditions at six rural sites between 580 and 1950 m a.s.l. We also assessed age-related differences in O(3) flux by examining changes in leaf conductance across the life span of Norway spruce. At the leaf level O(3) flux into the needles was effectively controlled by stomatal conductance and, hence by factors such as temperature, irradiance and humidity, which control stomatal conductance. Seasonal variations in O(3) flux were mainly attributed to the course of the prevailing temperature. During the growing season, however, data have emphasised leaf-air vapour pressure difference as the environmental factor most likely to control stomatal conductance and O(3) flux into the needles. In the sun crown stomatal conductance averaged over the growing season decreased with increasing tree age from 42.0+/-3.5 mmol O(3) m(-2) s(-1) in 17-year-old trees to 7.1+/-1.0 mmol O(3) m(-2) s(-1) in 216-year-old trees, indicating that O(3) concentration in the substomatal cavities is higher in young than in old trees. Independent from tree age stomatal conductance and O(3) flux were approximately 50% lower in shade needles as compared to sun-exposed needles. Stomatal conductance was also greater in the current flush (24+/-5.6 mmol O(3) m(-2) s(-1)) and in 1-year old needles (16+/-4 mmol O(3) m(-2) s(-1)) than in older needle age classes (12+/-1 mmol O(3) m(-2) s(-1), averaged across the four older needle age classes). In trees similar in age (60-65 years old) average O(3) flux into sun needles increased from 0.55+/-0.36 nmol m(-2) s(-1) at the valley floor to 0.9 nmol m(-2) s(-1) in 1950 m a.s.l. Cumulative O(3) uptake during the vegetation period increased from 11.4+/-1.7 mol m(-2) in the valley to 14 mol m(-2) at the alpine timberline. Although stomatal conductance provides the principal limiting factor for O(3) flux, additional field research is necessary in order to improve our understanding concerning the quantitative 'physiological threshold dose' which internally can be active and can have adverse effects of O(3) on forest trees.     

The variations of aluminium species in mountainous forest soils and its implications to soil acidification

Aluminium (Al) speciation is a characteristic that can be used as a tool for describing the soil acidification process. The question that was answered is how tree species (beech vs spruce) and type of soil horizon affect Al speciation. Our hypotesis is that spruce and beech forest vegetation are able to modify the chemical characteristics of organic horizon, hence the content of Al species. Moreover, these characteristics are seasonally dependent. To answer these questions, a detailed chromatographic speciation of Al in forest soils under contrasting tree species was performed. The Jizera Mountains area (Czech Republic) was chosen as a representative mountainous soil ecosystem. A basic forestry survey was performed on the investigated area. Soil and precipitation samples (throughfall, stemflow) were collected under both beech and spruce stands at monthly intervals from April to November during the years 2008–2011. Total aluminium content and Al speciation, pH, and dissolved organic carbon were determined in aqueous soil extracts and in precipitation samples. We found that the most important factors affecting the chemistry of soils, hence content of the Al species, are soil horizons and vegetation cover. pH strongly affects the amount of Al species under both forests. Fermentation (F) and humified (H) organic horizons contain a higher content of water extractable Al and Al³⁺ compared to organo-mineral (A) and mineral horizons (B). With increasing soil profile depth, the amount of water extractable Al, Al³⁺ and moisture decreases. The prevailing water-extractable species of Al in all studied soils and profiles under both spruce and beech forests were organically bound monovalent Al species. Distinct seasonal variations in organic and mineral soil horizons were found under both spruce and beech forests. Maximum concentrations of water-extractable Al and Al³⁺ were determined in the summer, and the lowest in spring.     

Quantifying ozone uptake at the canopy level of spruce,pine and larch trees at the alpine timberline: an approach based on sap flow measurement

Micro-climatic and ambient ozone data were combined with measurements of sap flow through tree trunks in order to estimate whole-tree ozone uptake of adult Norway spruce (Picea abies), cembran pine (Pinus cembra), and European larch (Larix decidua) trees. Sap flow was monitored by means of the heat balance approach in two trees of each species during the growing season of 1998. In trees making up the stand canopy, the ozone uptake by evergreen foliages was significantly higher than by deciduous ones, when scaled to the ground area. However, if expressed per unit of whole-tree foliage area, ozone flux through the stomata into the needle mesophyll was 1.09, 1.18 and 1.40 nmol m(-2) s(-1) in Picea abies, Pinus cembra and Larix decidua, respectively. These fluxes are consistent with findings from measurements of needle gas exchange, published from the same species at the study site. It is concluded that the sap flow-based approach offers an inexpensive, spatially and temporally integrating way for estimating ozone uptake at the whole-tree and stand level, intrinsicly covering the effect of boundary layers on ozone flux.