Effect of simulated acid fog on membrane-bound calcium (mCa) in fir (Abies firma) and cedar (Cryptomeria japonica) mesophyll cells

The decline of virgin fir (Abies firma) forest at Mt. Oyama has been reported. Related field observations suggest that high acidity fog is linked with its decline. However, cedar (Cryptomeria japonica) in the same area shows no symptoms of decline. For assessing effects of acid fog on membrane-bound calcium (mCa) of the leaf mesophyll cells, 9-year-old seedlings of fir (Abies firma) and 8-year-old seedlings of cedar (Cryptomeria japonica) were exposed twice a week to simulated acid fog (SAF at pH 3 with pH 5 as control) for 2 h per day in a chamber during May–December 2007 (except August). Current and 1-year-old needles were collected from seedlings and analyzed at 1-month intervals. For current-year needles of fir, mCa levels in cells exposed to SAF at pH 3 were significantly lower than in cells exposed to pH 5, especially during September 2007–March 2008. In contrast, it is noteworthy that mCa levels of cedar were maintained as virtually constant irrespective of SAF acidity, indicating that fir is more sensitive to acid fog than is cedar. Based on these results, mCa loss by acid fog might also be caused in the declining virgin fir forest at Mt. Oyama.     

Effects of complex stress of chronic acid fog exposure with rhizosphere aluminum on nutrient status in Abies firma seedlings

We exposed momi fir (Abies firma Sieb. et Zucc.) seedlings to simulated acid fog of pH 3.0 for about 3 years (from July 1999). During the last year of acid fog exposure, half of the seedlings were subjected to rhizosphere Al stress (complex stress) and the nutrient status of seedlings was determined. Chronic acid fog exposure decreased Fe and Zn concentrations in current-year and 1-year needles, and Al in 1-year needles, but had little effect on major element concentrations. Aluminum treatment had a broad impact on nutrient status in fine roots and needles. In fine roots, increases in Al and Cu concentrations and decreases in B, Mn, and Zn concentrations were observed. In 1-year needles, Al treatment increased Al, B, and Mn concentrations and decreased Cu concentration. The complex effect of acid fog with aluminum on nutrient status was relatively slight. These results show that changes in nutrient status under chronic acid fog exposure and/or Al stress are induced before the decline of photosynthesis in momi fir seedlings, mainly due to Al stress.     

Growth recovery of mature Norway spruce and European beech from chronic O<Subscript>3</Subscript> stress

Elevated O₃ levels can strongly impair the health and vitality of forest ecosystems. Free-air exposure systems reveal that forest tree and stand growth can be reduced strongly under chronic O₃ stress. Detailed knowledge of the effect of O₃ exposure on photosynthesis, carbon sequestration, allometry and growth during chronic stress is available. However, knowledge of growth response after O₃ reduction is scarce. Here, we analyse the growth of mature Norway spruce (Picea abies (L.) Karst.) and European beech (Fagus sylvatica L.) in the free-air O₃ fumigation experiment at Kranzberg Forest. We compare tree growth over a 9-year period (2008–2016) after exposure to O₃ (2000–2007). During 2?×?O₃ exposure, the annual basal area growth of Norway spruce and European beech decreased by 24 and 32%, respectively. After cessation of 2?×?O₃ exposure, the annual basal area growth of Norway spruce and European beech not only recovered but exceeded the growth of the trees in the control condition by 14 and 24%, respectively. The growth resilience and resistance of trees previously exposed to 2?×?O₃ towards drought stress and late frost was hardly lower than that of the trees in the control condition. The capacity for growth recovery even after long-term chronic O₃ stress emphasizes the strong beneficial effect of air pollution control on the health of forest ecosystems and on the global land carbon sink.     

Detection and quantification of changes in membrane-associated calcium in red spruce saplings exposed to acid fog

Five-year-old red spruce saplings (Picea rubens Sarg.) were exposed to either (1) acid fog consisting of a mixture of H(2)SO(4) and HNO(3) adjusted to pH 2.5, (2) distilled-water fog at pH 5.6, or (3) no fog (dry control) for 3.5 hours per day, five times a week during the 1996 and 1997 growing seasons. The effect of fog on cell membrane-associated calcium (mCa) of leaf mesophyll cells was investigated with the fluorescence probe chlortetracycline (CTC). In both years, mean mCa concentrations were significantly less in needles exposed to acid fog than in needles exposed to distilled-water fog or in untreated needles. In 1997, acid-fog treatment resulted in 25 and 12% reductions in mCa in current-year needles, and 18 and 15% reductions in 1-year-old needles, compared with untreated needles and needles exposed to distilled-water fog, respectively, indicating that acid deposition induced calcium leaching from the membranes of photosynthetic mesophyll cells. Exposure to distilled-water fog also led to reductions in mCa in young needles, suggesting that water films on needle surfaces can induce losses by diffusion between the needle interior and surface. Consistent with the chamber studies, field data obtained from red spruce trees at two sites in Maine showed that low mCa concentrations in needles were associated with exposure to acid fog.     

Effects of chronic acid fog exposure with soil acidification or nitrogen loading on nutrient status in Abies firma seedlings

To clarify the nutrient status in momi fir (Abies firma Sieb. et Zucc.) seedlings under complex stress conditions of acid fog with soil acidification or nitrogen loading, we exposed seedling shoots to simulated acid fog (pH 3) and simultaneously loaded the soil with acid or excess nitrogen for 17 months. Proton and nitrogen loading reduced K concentration in soil, but these treatments had little effect on the nutrient status of fine roots in momi fir seedlings. Acid fog exposure resulted in reduced concentrations of Mg, Al, and Fe in current-year needles and Mg, Al, Cu, Fe, and Zn in 1-year-old needles. The complex effects of acid fog exposure with proton or nitrogen loading on nutrient status were relatively slight. However, elements such as Al and Cu notably reflect the effects of proton loading. These results show that the nutrient status of trace elements, rather than major elements, in needles of momi fir seedlings is sensitive to exposure to acid fog or proton loading.     

Effects of partial defoliation on carbon and nitrogen partitioning and photosynthetic carbon uptake by two-year-old cork oak (Quercus suber) saplings

At the end of the growing season in late July, 20-month-old cork oak (Quercus suber L.) saplings were partially defoliated (63% of leaf area) to evaluate their ability to recover leaf area after defoliation. At 18 and 127 days after defoliation, changes in starch and nitrogen pools were determined in leaves and perennial organs, and variations in photosynthetic carbon uptake were investigated. To determine the role of stored nitrogen in regrowth after defoliation, plant nitrogen was labeled in the previous winter by enriching the nutrient solution with 15N. Plants recovered the lost leaf area in 127 days. Although there was remobilization of starch and nitrogen from leaves and perennial organs, the availability of resources for growth in the following spring was not decreased by defoliation. On the contrary, starch concentration in coarse roots was higher in defoliated saplings than in control saplings, presumably as a result of the higher net CO2 exchange rate in newly developed leaves compared with pre-existing leaves.     

Response of gas exchange rates in <Emphasis Type="Italic">Abies firma</Emphasis> seedlings to various additional stresses under chronic acid fog stress

In order to clarify the chronic influence of acid fog on the gas exchange rates of momi fir (Abies firma Sieb. et Zucc.) trees, we exposed them to simulated acid fog (pH 3) for 3 years. The composition of the acid fog was similar to that observed in a region where momi fir trees have been declining, and it contained organic acids. We then treated the firs with various additional stresses, such as drought, low temperature, fine root cutting, ozone exposure, soil acidification, nitrogen load, and rhizosphere aluminum stress. Under chronic exposure to acid fog, the momi fir seedlings exhibited a pattern of stomatal behavior whereby they excessively opened in summer and closed in winter. Furthermore, the stomata of these seedlings tended to open during drought stress, and their needles were visibly injured after ozone exposure. The net photosynthesis rates of the seedlings exposed to acid fog were regulated by their stomatal aperture, rather than directly by acid fog. These results suggest that acid fog exposure disturbs the control of stomatal function in the momi fir seedlings. In addition, we found that chronic acid fog exposure suppressed the decrease in net photosynthesis rate, due to its nitrogen load.     

Physiological defence reactions of young beech trees (Fagus sylvatica) to attack by Phyllaphis fagi

The physiological reaction to attack by the lachnid Phyllaphis fagi compared with non-infested control plants was examined in whole leaves and leaf phloem of beech (Fagus sylvatica) saplings as well as in beach seedlings cultivated in different nutrient substrates. The colonisation density of the lachnids on beech seedlings increased with improved nutrient supply. However, growth was not depressed to a comparable extent. Defence reactions were found at three hierarchic levels. Infested plants exhibited a strongly decreased amino acid/monosaccharide ratio in the entire leaf and leaf phloem. In addition, secondary plant defence substances were detected in beech saplings and lamma shoot leaves of beech seedlings. A systematic reaction, dependent on the infestation density of the lachnid was observed in the leaf phloem of the beech saplings.     

Influence of foliar phenology and shoot inclination on annual photosynthetic gain in individual beech saplings: A functional–structural modeling approach

We developed a functional–structural plant model for Fagus crenata saplings and calculated annual photosynthetic gains to determine the influences of foliar phenology and shoot inclination on the carbon economy of saplings. The model regenerated the three-dimensional shoot structure and spatial and temporal display of leaves; we calculated the hourly light interception of each leaf with a detailed light model that allowed us to estimate hourly leaf photosynthetic gain taking leaf age into account. To evaluate the importance of simultaneous foliar phenology and slanting shoots in beech saplings, we calculated the photosynthetic budgets for saplings with contrasting foliar phenologies and shoot inclinations. In our simulations, we distinguished between simultaneous and successive foliar phenologies, upright and slanting shoot inclinations, and environments with and without a vertical gradient in light intensity. Other model parameters (including photosynthesis vs. light curve, leaf size, and leaf shape) were obtained directly from live beech saplings. With no vertical gradient in light intensity, modeled saplings with simultaneous foliar phenology and slanting shoots (as in live beech) had larger annual photosynthetic gains than saplings with other combinations of traits. Hence, simultaneous foliar phenology and slanting shoots are efficient ways to display leaves in the shaded forest understory light regime where beech saplings thrive. In the presence of vertical light gradients, which can occur in canopy gaps, saplings with upright shoots had larger annual photosynthetic gains than counterparts with slanting shoots. Although mean daily photosynthetic gains of saplings with successive foliar phenology were elevated by exposing leaves to strong light when young and productive, the annual photosynthetic budget of these saplings was reduced (compared to saplings with simultaneous foliar phenology) by their relatively short leaf lifespan. Overall, our results suggest that slanting shoots with simultaneous foliar phenology are particularly successful in shaded environments, where beech often dominates, because they appear to maximize the annual carbon budget by avoiding self-shading and extending leaf lifespans.     

Soluble N compound profiles and concentrations in European beech (Fagus sylvatica L.) are influenced by local climate and thinning

We assessed seasonal changes of total soluble nonprotein nitrogen compounds (TSNN) in adult European beech trees (Fagus sylvatica, L.) growing under different local climate during the growing season immediately following a thinning treatment and 3 years later. In both years, samples of leaves, xylem sap and phloem exudates from beech trees growing in thinned and unthinned (control) stands on a dry, warm SW exposed and a cooler, moist NE exposed site were collected in May, July and September. In May of both years, asparagine (Asn) and glutamine (Gln) were most abundant in leaves and xylem, respectively, whereas arginine (Arg) dominated in the phloem. In July, TSNN concentrations decreased in all tissues and sites, but differences in water availability between aspects were reflected in TSNN concentrations. In September, differences in the increase of Arg concentration in the phloem were related to differences in the onset of senescence between treatments. Thinning treatment increased amino compound concentrations of beech tissues in July on both aspects, particularly at the NE thinned site. It is supposed that, the N balance of adult beech is favoured by both, the thinning treatments as well as the cool-moist climate prevailing at the NE aspect.     

Biomass,nutrient and pigment content of beech (Fagus sylvatica) saplings infected with Phytophthora citricola,P. cambivora,P. pseudosyringae and P. undulata

Fagus sylvatica saplings were infected with Phytophthora citricola, Phytophthora cambivora, Phytophthora pseudosyringae and Phytophthora undulata to study the influence of these root pathogens on total belowground and aboveground biomass, on the nutrient distribution within plants, on the concentration of plastid pigments, including tocopherol and on components of the xanthophyll cycle. Phytophthora citricola and P. cambivora infection significantly reduced total biomass of beech when compared with control plants and finally most of these plants died at the end of the experiment. However, beech invaded by the other two Phytophthora spp. did not differ from control plants and none of them was killed. Fine root length as well as the number of root tips of all infected beeches were reduced between 30 and 50%. The excellent growth of beech infected with P. pseudosyringae and P. undulata when compared with control plants was correlated with a strong increase of important root efficiency parameters. Phytophthora citricola and P. cambivora caused a significant reduction in nitrogen concentration of leaves in comparison with control and other infected plants, whereas this nutrient was slightly increased in fine and coarse roots. Furthermore, the phosphorus and potassium concentrations in leaves were impaired after infection with P. citricola. However, foliar concentrations of Ca and Mg were not affected by the different Phytophthora spp., whereas fine and coarse roots were significantly enriched with Ca in beech infected with P. citricola or P. cambivora. The concentrations of α‐tocopherol and xanthophyll cycle pigments were increased in plants infected by P. citricola and P. cambivora, indicating that several reactive oxygen species might be formed in leaves during infection.     

Biomass and nutrients allocation in pot cultured beech seedlings： influence of nitrogen fertilizer

Allocation of biomass and nutrient elements including Nitrogen to above and belowground compartments of beech seedlings (Fagus sylvatica L.) treated by labeled nitrogen fertilizer in the form of ¹⁵NH₄ and ¹⁵NO₃ were investigated at the end of two successive growing seasons. Pot cultured beech seedlings were grown at a green house on intact soil cores sampled from three adjacent stands including beech, Norway spruce and mixed beech-spruce cultures of Solling forest, Germany. Comparing biomass allocation and nutrients concentrations of the seedlings between the control and ¹⁵N-fertilized treatments revealed no significant effect of N fertilization on nutrients uptake by seedlings over the experiment. The form of N input influenced its movement into plant pools. It was demonstrated that beech seedlings take up nitrogen mainly in the form of nitrate, which is then reduced in the leaves, although the differences between the retention of NO₃ ^?-N and NH₄ ⁺-N in plants were not statistically significant. Percent recoveries of ¹⁵N in trees were typically greater after ¹⁵NO₃ than after ¹⁵NH₄ additions. It was indicated that immobilization of 15N tracer in fine roots was a slower process comparing other plant compartments such as stem and coarse roots, but a powerful sink for N during the course of study.     

UV‐B radiation constrains the photosynthesis of Quercus robur through impacts on the abundance of Microsphaera alphitoides

Quercus robur saplings were exposed at an outdoor facility in the UK to supplemental levels of UV‐B radiation (280–315 nm) under arrays of cellulose diacetate‐filtered fluorescent lamps which also produced UV‐A radiation (315–400 nm). Saplings were also exposed to supplemental UV‐A radiation under arrays of polyester‐filtered lamps and to ambient levels of solar radiation under arrays of unenergized lamps. The UV‐B treatment was modulated to maintain a 30% elevation above the ambient level of erythemally weighted UV‐B radiation. Naturally occurring infections by oak powdery mildew (Microsphaera alphitoides) were more abundant, and developed more rapidly, on lammas leaves of saplings which were exposed to treatment levels of UV‐B radiation than on leaves of saplings exposed to supplemental UV‐A or to ambient levels of solar radiation over 12 weeks in summer and autumn 1996. An analysis of leaf photosynthetic capacities revealed that M. alphitoides infection reduced the quantum efficiency of photosystem (PS) II by 14% at moderate irradiance. Although there was no direct effect of UV‐B radiation on PSII photochemistry, exposure of saplings to supplemental UV‐A radiation under polyester‐filtered lamps resulted in a 17.5% decrease in PSII quantum efficiency, compared with saplings exposed to ambient solar radiation. The results from our study suggest that photosynthesis of Q. robur may be constrained by exposure to UV‐B radiation in the natural environment through impacts on the abundance of M. alphitoides.     

Effects of elevated CO(2) concentration on leaf characteristics and photosynthetic capacity of beech (Fagus sylvatica) during the growing season

Two-year-old beech (Fagus sylvatica L.) saplings were planted directly in the ground at high density (100 per m(2)), in an experimental design that realistically mimicked field conditions, and grown for two years in air containing CO(2) at either ambient or an elevated (ambient + 350 ppm) concentration. Plant dry mass and leaf area were increased by a two-year exposure to elevated CO(2). The saplings produced physiologically distinct types of sun leaves associated with the first and second growth flushes. Leaves of the second flush had a higher leaf mass per unit area and less chlorophyll per unit area, per unit dry mass and per unit nitrogen than leaves of the first flush. Chlorophyll content expressed per unit nitrogen decreased over time in plants grown in elevated CO(2), which suggests that, in elevated CO(2), less nitrogen was invested in machinery of the photosynthetic light reactions. In early summer, the photosynthetic capacity measured at saturating irradiance and CO(2) was slightly but not significantly higher in saplings grown in elevated CO(2) than in saplings grown in ambient CO(2). However, a decrease in photosynthetic capacity was observed after July in leaves of saplings grown in CO(2)-enriched air. The results demonstrate that photosynthetic acclimation to elevated CO(2) can occur in field-grown saplings in late summer, at the time of growth cessation.     

Compared effects of competition by grasses (Graminoids) and broom (Cytisus scoparius) on growth and functional traits of beech saplings (Fagus sylvatica)

? The effect of two weeds (grasses and broom, Cytisus scoparius) competition on the growth and functional traits of European beech saplings (Fagus sylvatica) was investigated in an experimental plantation in the French Massif central.

? We hypothesized that grasses would have a much more harmful effect than broom on beech growth through strong competition for soil water and nitrogen.

? A randomized block design was used with three separate blocks, each possessing three types of vegetation; grasses (mainly Agrostis capillaris, Holcus mollis and H. lanatus), broom, and bare soil as a control.

? Two years after plantation beech sapling growth was significantly greater in bare soil than with competition from grasses, and was intermediate on broom. Growth was positively correlated with beech leaf nitrogen content (N_m) and xylem water potential (Ψ_x). N and Ψ_x were dependent, with nitrogen uptake probably decreasing with increasing drought. There was no clear correlation between growth and light transmitted 10 cm below the apex of the saplings.

? Beech leaf mass on an area basis (LMA) was correlated with the three resource variables (water, nitrogen and light) and with growth, confirming that LMA is a sensitive functional trait integrating the degree of stress experienced by the saplings.

? The results suggest that broom is less competitive than grasses and, if properly managed on a sufficiently well-watered site, can generate satisfactory conditions for beech establishment and growth.

     

Effects of the absorption of artificial acidic solutions on the root systems of Japanese red cedar (Cryptomeria japonica D. Don) cutting and saplings

The effects of the absorption of artificial acidic solutions only from below-ground parts on root systems were examined for 60 days in summer and 72 days in fall using cuttings and saplings of Japanese red cedar (Cryptomeria japonica D. Don). Cuttings and saplings absorbed the solutions through the potted soils from the reservoir of an autoirrigator and/or a simplified-autoirrigator. Nitric acid solutions of pH 2.0, 4.0 and 6.0 and distilled water (control) were used in summer, mixtures of H₂SO₄ and HNO₃ solutions of pH 2.0, 3.0 and 4.0, HNO₃ solution of pH 3.0, H₂SO₄ solution of pH 3.0 and distilled water (control) were used in fall. Although no detrimental effects of acidic solutions on saplings were observed, the rooting rate of cuttings treated at pH 2.0 in summer was significantly lower than that of the control. Root lengths and root dry weight per cutting at pH 2.0 in summer and fall were smaller than those of the control, and the difference in fall was significant. The present study revealed that the acidic treatment only from below-ground parts had detrimental effects on the root systems of cuttings, suggesting the importance of quantification of the effects on below-ground parts, as well as on above-ground parts, to evaluate the effects of acidic precipitation on trees.     

Physiological responses of birch (Betula pendula) to ozone: a comparison between open-soil-grown trees exposed for six growing seasons and potted seedlings exposed for one season

Physiological responses of 4-year-old potted saplings of an O3-tolerant clone of Betula pendula Roth to short-term ozone (O3) exposure (one growing season) were compared with those of 6-year-old open-soil-grown trees of the same clone fumigated with O3 for six growing seasons. In the 2001 growing season, both groups of plants were exposed to ambient (control) and 1.6x ambient (elevated) O3 concentration under similar microclimatic conditions in a free air O3 exposure facility. Growth, net photosynthesis, stomatal conductance, stomatal density, visible foliar injury, starch and nutrient concentrations, bud formation and differences in O3 responses between lower, middle and upper sections of the canopy were determined. The potted saplings were unaffected by elevated O3 concentration, whereas the open-soil-grown trees showed a 3-38% reduction in shoot growth, a 22% reduction in number of overwintering buds, a 26-65% decrease in autumnal net photosynthesis, 30% and 20-23% reductions in starch and nitrogen concentrations of senescing leaves, respectively, and disturbances in stomatal conductance. The greater O3 sensitivity of open-soil-grown trees compared with potted saplings was a result of senescence-related physiological factors. First, a lower net photosynthesis to stomatal conductance ratio in open-soil-grown trees at the end of the season promoted O3 uptake and decreased photosynthetic gain, leading to the onset of visible foliar injuries. Second, decreased carbohydrate reserves may have resulted in deleterious carry-over effects arising from the reduced formation of over-wintering buds. Finally, the leaf-level O3 load was higher for open-soil-grown trees than for potted saplings because of slower leaf senescence in the trees. Thus, O3 sensitivity in European white birch increases with increasing exposure time and tree size.     

Nutrient concentrations in roots, leaves and wood of seedling and mature sugar maple and American beech at two contrasting sites

Differences in sensitivity to soil conditions across tree species and developmental stage are important to predicting forest response to environmental change. This study was conducted to compare elemental concentrations in leaves, stems, and roots of (1) sugar maple (Acer saccharum Marsh.) seedlings vs. mature trees and (2) mature sugar maple vs. mature American beech (Fagus grandifolia Ehrh.) in two sites that differ in soil base saturation and pH. Both sites are located in Huntington Forest, NY, USA; one site (hereafter ‘H’) has higher soil pH and Ca, Mg, and Mn concentrations than the other site (hereafter ‘L’). Sugar maple growth at H (14.8 cm² year⁻¹ per tree) was much greater than at L (8.6 cm² year⁻¹ per tree), but the growth of beech was not different between the two sites. Leaves, roots, and stem wood of mature beech trees and sugar maple seedlings and mature trees were sampled for nutrient analysis. Foliar Ca, K, and Al concentrations were positively correlated with soil elements, but Mn concentrations were negatively correlated. Sugar maple differed more than beech between sites in foliar K and Mn concentrations. Root Mg and P concentrations reflected soil chemistry differences, in contrast to foliar concentrations of Mg and P, which were indistinguishable between the sites. In sugar maple, seedlings differed more than in mature trees in nutrient concentrations in roots, especially for Mg and Mn. Although beech was not as responsive to nutrient availability as sugar maple in foliar and root nutrient concentrations, Ca and Mg concentrations in beech wood were higher in H (52% higher for Ca and 68% higher for Mg), while sugar maple did not differ between sites. Sugar maple regeneration failure on acidic soils in the same region is consistent with our finding that sugar maple seedlings were very sensitive to nutrient availability. This sensitivity could ultimately contribute to the replacement of sugar maple by American beech in regions of low pH and base cations if base cation leaching by anthropogenic deposition and tree harvesting continues.     

Nutrient contents and efficiencies of beech and spruce saplings as influenced by competition and O<Subscript>3</Subscript>/CO<Subscript>2</Subscript> regime

Saplings of Fagus sylvatica and Picea abies were grown under conditions of intra and interspecific competition in a 2-year phytotron study under combinations of ambient and elevated ozone (+O₃ which is 2 × O₃, but <150 nl l⁻¹) as well as carbon dioxide concentrations (+CO₂ which is amb. CO₂ + 300 μl CO₂ l⁻¹) in a full factorial design. Saplings were analysed for various mineral nutrients in different plant organs as well as biomass production and crown development. The study was based on the assumption that nutritional parameters important for growth and competitiveness are affected by stress defence under limiting nutrient supply. The hypotheses tested were (1) that nutrient uptake-related parameters (a) as well as efficiencies in nutrient use for above-ground competition (b) of beech rather than spruce are impaired by the exposure to elevated O₃ concentrations, (2) that the efficiency in nutrient uptake of spruce is enhanced by elevated CO₂ concentrations in mixed culture, and (3) that the ability to occupy above-ground space at low nutrient cost is co-determinant for the competitive success in mixed culture. Clear nitrogen deficiencies were indicated for both species during the 2-year phytotron study, although foliar nitrogen-biomass relationships were not so close for spruce than for beech. O₃ stress did not impair nutrient uptake-related parameters of beech; thus hypothesis (1a). was not supported. A negative effect of elevated O₃ (under amb. CO₂) on the N and P based efficiencies in above-ground space occupation (i.e. lower crown volume per unit of N or P invested in stems, limbs and foliage) of beech supported hypothesis (1b). It appeared that ozone stress triggered a nutrient demand for stress defence and tolerance at the expense of above-ground competition (trade-off). Crown volume of beech under O₃ stress was stabilized in monoculture by increased nutrient uptake. In general, the +CO₂-treatment was able to counteract the impacts of 2 × O₃. Elevated CO₂ caused lower N and S concentrations in current-year foliage of both tree species, slightly higher macronutrient amounts in the root biomass of spruce, but did not increase the efficiencies in nutrient uptake of spruce in mixed culture. Therefore hypothesis (2) was not supported. At the end of the experiment spruce turned out to be the stronger competitor in mixed culture as displayed by its higher total shoot biomass and crown volume. The amounts of macronutrients in the above-ground biomass of spruce individuals in mixed culture distinctly exceeded those of beech, which had been strongly reduced by interspecific competition. The superior competitiveness of spruce was related to higher N and P-based efficiencies in above-ground space occupation as suggested in hypothesis (3). This article belongs to the special issue “Growth and defence of Norway spruce and European beech in pure and mixed stands”.