The three youngest age-classes of needles of Norway spruce (Picea abies) were collected from four sites in the Krusne Hory Mountains (Czech Republic) characterized by different levels of damage caused by environmental pollution. Histochemical methods did not reveal any differences in localization of phenolics among the needles. Mesophyll cells close to the epidermis of needles and cells around resin ducts and substomatal cavities often accumulated higher amounts of phenolics than the rest of the mesophyll cells, but this was independent of age and damage. Needles of different age- and damage-class did not show any marked changes in general lignification pattern. However, a lower intensity of histochemical detection of lignin was observed in needles from the most damaged site. This finding was confirmed by chemical analysis using thioglycolic acid. Generally, the amount of lignin in mesophyll cells was lower in damaged trees than in healthy ones. Using the Folin–Ciocalteau method, no significant differences in the total content of phenolics were observed in the needles, although HPLC revealed marked alterations in the forms of seven phenolic acids. Concentrations of conjugated forms of phenolic acids (esters and glycosides) were higher in damaged needles (255.9 μg g−1 f. wt) than in healthy needles (189.8 μg g−1 f. wt). By contrast, content of esterified phenolic acids incorporated into cell walls was higher in needles from healthy trees (101.1 μg g−1 f. wt) than in damaged needles (78.3 μg g−1 f. wt). Marked differences were also observed in the activity of soluble peroxidases, although the activity of ionically bound forms was approximately the same in healthy and damaged needles. The total amounts of chlorophylls and carotenoids decreased as environmental damage increased.

Growth responses of two provenances of European beech (Fagus sylvatica) were studied. The seedlings were grown in closed-top chambers at four temperature regimes (−2.9 °C below ambient, ambient, +2.3 °C and +4.8 °C above ambient) in combination with two CO2 partial pressures (40 Pa and 74 Pa). Growth was followed by making destructive harvests c. every 25 d from germination in early June to senescence in late September. Allocation patterns were significantly affected by the temperature regimes. However, changes in dry matter allocation and morphology associated with the different treatments at a given time were mostly a result of differences in tree size. Temperature regimes only had a significant effect on the relative growth rate, RGR, at the beginning of the experiment. In contrast to temperature, high [CO2] increased RGR throughout the experiment when compared with plants of equal size. As the trees increased in size net assimilation rate, NAR, decreased but the effect of [CO2] on both NAR and RGR had a tendency to increase. Increases in NAR caused by elevated [CO2] were partly counteracted by reductions in the leaf area ratio, LAR. Reductions in LAR were caused by concomitant reductions in specific leaf area, SLA, whereas the level of [CO2] did not significantly affect leaf weight ratio, LWR, nor other dry weight ratios. The interactions between temperature and [CO2] are highly dependent on whether they are expressed as instantaneous values for plants at a common age or instantaneous values at a common size (and thereby extracting the effects of ontogenetic drift). When comparing instantaneous values at common sizes, the positive effect of [CO2] on RGR increased with plant size in every temperature regime. This also occurred in every temperature regime when comparing plants of equal age but the response to [CO2] was less. The effect of [CO2] on RGR was dependent on growth temperature. The positive effects of elevated [CO2] on RGR were less than the positive effect on photosynthesis. The two provenances did not differ significantly in the response of RGR to [CO2] which is in agreement with measurements of photosynthesis.

We conducted two experiments to investigate the expression of shade avoidance in response to low ratios of red to far-red irradiation (R[ratio ]FR) in the C4 perennial grass Schizachyrium scoparium at two levels of above-ground (photosynthetic photon flux density (PFD) 400–700 nm) and below-ground (soil volume) resource availability and in plants of two ages. Young plants showed greater sheath and ramet height in response to low R[ratio ]FR and old plants showed reduced ramet initiation in experiments one and two, respectively, but both responses were not expressed simultaneously in either group. Growth of all ramet variables, including ramet initiation, was suppressed in small as opposed to large soil volumes. By contrast, architectural variables, but not ramet initiation, were greater for plants grown in low as opposed to ambient PFD. However, expression of shade avoidance to low R[ratio ]FR was not significantly affected by either level of PFD or soil volume. We must conclude that the levels of resource availability provided do not modulate shade avoidance in this perennial grass. These results demonstrate that S. scoparium is capable of expressing shade avoidance in response to low R[ratio ]FR, but inconsistent juvenile ramet initiation and architectural responses in plants of different ages and phenological stages of development indicate that the shade avoidance response might not be expressed consistently throughout the life of plants.

Plants use chemical defences to reduce damage from herbivores and the effectiveness of these defences can be altered by biotic and abiotic factors, such as herbivory and soil resource availability. Streptanthus polygaloides, a nickel (Ni) hyperaccumulator, possesses both Ni-based defences and organic defences (glucosinolates), but the extent to which these defences interact and respond to environmental conditions is unknown. S. polygaloides plants were grown on high-Ni and low-Ni soil and concentrations of Ni and glucosinolates were compared with those of the congeneric non-hyperaccumulator, S. insignus spp. insignus, grown under the same conditions. Ni contents were highest (4000 μg g−1 dry tissue) in S. polygaloides plants grown on high-Ni soil. Glucosinolate content was significantly higher in S. insignus than in S. polygaloides suggesting that plants defended by Ni produce a lower concentration of organic defences. In a separate experiment, high-Ni S. polygaloides plants were exposed to simulated herbivory or live folivores to determine the inducibility of Ni-based and organic defences. Contents of Ni were not affected by either herbivory treatment, whereas glucosinolate concentrations were >30% higher in damaged plants. We concluded that the Ni-based defence of S. polygaloides is not induced by herbivory.

Seedling mortality of Alyssum serpyllifolium ssp. lusitanicum and A. murale, both nickel hyperaccumulators, was reduced by increasing concentrations of metal within plant tissues when inoculated with the fungi Pythium mamillatum or P. ultimum, both of which cause damping-off disease of seedlings. Pythium mamillatum, isolated from nickel-rich serpentine soil, was more tolerant of nickel than P. ultimum, isolated from low-metal control soil, and was more pathogenic than P. ultimum towards seedlings containing high concentrations of metal. These results support the hypothesis that metal hyperaccumulation by plants is closely linked to increased protection against disease.

The effects of elevated concentrations of atmospheric carbon dioxide and ozone on diurnal patterns of photosynthesis have been investigated in field-grown spring wheat (Triticum aestivum). Plants cultivated under realistic agronomic conditions, in open-top chambers, were exposed from emergence to harvest to reciprocal combinations of two carbon dioxide and two ozone treatments: [CO2] at ambient (380 μmol mol−1, seasonal mean) or elevated (692 μmol mol−1) levels, [O3] at ambient (27 nmol mol−1, 7 hr seasonal mean) or elevated (61 nmol mol−1) levels. After anthesis, diurnal measurements were made of flag-leaf gas-exchange and in vitro Rubisco activity and content. Elevated [CO2] resulted in an increase in photoassimilation rate and a loss of excess Rubisco activity. Elevated [O3] caused a loss of Rubisco and a decline in photoassimilation rate late in flag-leaf development. Elevated [CO2] ameliorated O3 damage. The mechanisms of amelioration included a protective stomatal restriction of O3 flux to the mesophyll, and a compensatory effect of increased substrate on photoassimilation and photosynthetic control. However, the degree of protection and compensation appeared to be affected by the natural seasonal and diurnal variations in light, temperature and water status.

This paper reports the effects of nutrient magnesium (Mg) concentrations on the growth and photosynthetic physiology of clonal Pinus radiata from four female parents (families) known to differ in their tolerance to Mg deficiency and in their needle Mg concentrations. Plants were grown in flowing nutrient solutions with 2 mg l−1 (control) and 0.8 mg l−1 (low) Mg. Plant growth, needle Mg concentration, photosynthesis, chlorophyll fluorescence and carotenoid pigment content were measured. At low Mg, needle Mg concentration was about half that of control plants, height growth was reduced 15–25%, and the needles showed strong visual characteristics of Mg deficiency. Photosynthesis was also halved, and was associated with closure of the stomata under low Mg and with reductions in the residual conductance. In needles from plants grown at low Mg, photochemical yield was reduced both in the light and in the dark, and was strongly dependent on needle Mg concentrations below a threshold concentration of 0.02–0.025% (d. wt basis). The electron transport rate (ETR) at saturating photon flux density in low-Mg-grown needles was reduced to about half that of their Mg controls, but the photon efficiency of ETR was unaffected by the Mg concentration the plants were grown in. Photosynthetic quenching was markedly reduced and non-photosynthetic quenching was increased following growth in low Mg. Growth under low Mg also increased levels of zeaxanthin. Although family differences in growth and photosynthetic physiology were present, few family × Mg interactions were significant. We conclude that Mg deficiency probably affects growth through severe reductions in photosynthesis.

Zinc tolerance was investigated in five populations of Arabidopsis halleri (syn.: Cardaminopsis halleri) raised from seeds collected from contaminated and uncontaminated sites. Tolerance was measured by determining the concentration which inhibited root growth (EC100). A. halleri populations from contaminated and uncontaminated sites were found to be Zn-tolerant compared with the Zn-nontolerant species Arabidopsis thaliana and A. lyrata subsp. petraea. At very high Zn concentrations, populations of A. halleri from uncontaminated sites were slightly less Zn-tolerant than those from contaminated sites. These observations support the hypothesis that in A. halleri, Zn tolerance is largely a constitutive property. One population from an uncontaminated site and one population from a contaminated site were studied for Zn uptake. Zinc content was measured in shoots and roots using a colorimetric test under laboratory conditions. The results showed that whatever their origin, individuals from both populations are Zn accumulators compared with the nonaccumulator species A. thaliana. Moreover, the population from the uncontaminated area accumulated Zn in its shoots and roots more quickly than the population from the contaminated site. These results suggest that, in A. halleri, Zn accumulation to very high concentration is a constitutive property.

Seedlings of Quercus petraea (oak), Fraxinus excelsior (ash) and Pinus sylvestris (Scots pine) were grown at two CO2 concentrations with two O3 and two water supply treatments for 3 yr in a factorial experiment. Oak was the most responsive species to all three treatments; elevated CO2 and irrigation increased biomass by an average of 79% and 41%, respectively, whereas the ozone treatment resulted in a 30% reduction in growth. Significant treatment interactions in this species demonstrated that CO2 ameliorated and irrigation exacerbated the effects of ozone. For Scots pine and ash, irrigation and elevated CO2 increased growth by approx. 60% and 20%, respectively, whereas ozone had no detectable effect on ash and resulted in a 15% reduction in growth of Scots pine. Carbon partitioning to the shoot was enhanced by both the CO2 and H2O treatments in oak, while branching was also increased in this species in response to elevated O3, resulting in changes to the allometric relationships. CO2 enhanced leaf production in oak and Scots pine, and together with the promotion of shoot allocation, this indicates an increased susceptibility to windthrow. Biomass accumulation expressed as relative growth rate, suggested three different time-dependent growth responses to elevated CO2; the CO2 fertilization effect was maintained through the third year of growth in oak, had disappeared in Scots pine and a negative effect was evident in ash. Foliar nitrogen and chlorophyll concentrations indicated a CO2-induced nitrogen deficiency in oak and ash, but not in Scots pine. Chlorophyll degradation in response to ozone was observed in oak, an effect that was enhanced by irrigation and reduced by CO2, presumably through stomatal mediated changes in effective ozone dose. These results therefore suggest that elevated atmospheric CO2 concentrations will enhance growth of some UK forest tree species, although this might only be apparent during the juvenile phase. However, nitrogen deficiencies might limit this enhancement on some sites while changes in allocation and leaf area might promote susceptibility to windthrow. Elevated CO2 also provides some protection against ozone pollution, especially in combination with limited soil moisture availability. These interactions between CO2, ozone and water supply should be taken into account when predicting the effects of environmental change on tree growth and forest productivity.

Ring widths of five Mediterranean forest tree species (Arbutus unedo, Fraxinus ornus, Quercus cerris, Quercus ilex and Quercus pubescens) growing close to a natural source of CO2 in Tuscany, Italy and at a nearby control site were compared. At the CO2-enriched site, trees have been growing for decades under elevated CO2 concentrations. They originated from parent trees that also grew under elevated CO2 in natural conditions, and they have been continuously exposed to elevated CO2 throughout their growth. Tree-ring series from each of the species were prepared. Assigning calendar dates to rings was difficult but possible, and ring-width series were built for all species. The ring-width data were analysed using a two-sided t-test to assess if there was a difference between the radial growth at the CO2-enriched site and the control site. The cumulative basal area at the same cambial age at both sites was also compared using a Wilcoxon test. Radial growth of trees at the CO2-enriched site was not significantly different from growth at the control site. For each species, year by year, radial growth at the CO2-enriched site was tested against the control site and significant differences were found in only a few years; these differences were not synchronous with extreme climatic events. The expected increase in above-ground productivity, as one of the ecosystem responses to increasing CO2 during drought stress, was not observed in this Mediterranean woody plant community, despite being water-limited. Other resource limitations, such as low nutrient availability (common in the Mediterranean region), may have counteracted the positive effect of elevated CO2 under drought stress, or trees may have acclimated to the high CO2.

Plants of Molinia caerulea were grown in sand culture for two seasons. All nitrogen (N) supplied to the plants was enriched with 15N throughout the first season, and at natural isotopic abundance throughout the second season. A series of destructive harvests was taken during the second season. At each harvest, the N mobilized from roots and swollen basal internodes and the protease activity (ability to degrade azocasein) of these storage tissues were measured. The pH response curves of protease activity of both basal internodes and roots exhibited optima at pH 5 throughout the season. The protease activity of roots and basal internodes increased in spring, both on a unit fresh weight and unit protein basis, concomitant with mobilization of N from both these tissues to new shoot growth. In absolute terms more N was mobilized from roots than from basal internodes. However, basal internodes which, compared with roots, showed the greatest protease activity (on a f. wt or unit protein basis) were also the tissues that mobilized a greater proportion of N present in the tissue over winter to new shoot material. Individual clones of M. caerulea varied both in the amount of N mobilized from roots and in root protease activity (per plant). Individuals with a higher protease activity mobilized more N compared with individuals of lower protease activity. Therefore in M. caerulea, relationships between N mobilization and protease activity exist at several levels: (1) between different tissues; (2) temporally throughout the season; and (3) between individual clones.

We assessed the ability of several populations of the metal-hyperaccumulator species, Thlaspi caerulescens, to mobilize non-labile cadmium in soils historically contaminated by Pb/Zn mine spoil or sewage sludge. Radio- labile Cd was determined chemically as an ‘E-value’, [CdE], and biologically as an ‘L-value’, [CdL]. For comparison, chloride-extractable Cd, [Cdchlor], was also determined using 1 M CaCl2 as a single-step soil extractant. Values of [CdL] were measured for six populations of T. caerulescens that varied substantially in their ability to assimilate soil Cd, and a non-accumulator species with a similar growth habit, Lepidium heterophyllum. Seeds were sown in soil spiked with 109Cd and grown for 9–12 wk in a controlled environment room. Values of [CdL] were determined from the specific activity of 109Cd and concentration of Cd in the plant leaves. For the six soils studied, [CdE] ranged from 4.9 to 49% of total soil Cd [CdT]. Values of [CdL] were, in general, in close agreement with both [CdE] and [Cdchlor] and substantially less than [CdT]. However, [CdL] showed no correlation with the concentration of Cd in plant tissue, [Cdshoot]. This suggests that, in the soils studied, T. caerulescens did not mobilize non-labile soil Cd by producing root exudates or altering rhizosphere pH. The results imply that there may be significant restrictions to metal bioavailability, even to hyperaccumulator species, in heavily contaminated soils in which a large proportion of the metal may be present in ‘non-labile’ forms.

Xylem maturation in elongating leaf blades of tall fescue (Festuca arundinacea) was studied using staining and microcasting. Three distinctive regions were identified in the blade: (1) a basal region, in which elongation was occurring and protoxylem (PX) vessels were functioning throughout; (2) a maturation region, in which elongation had stopped and narrow (NMX) and large (LMX) metaxylem vessels were beginning to function; (3) a distal, mature region in which most of the longitudinal water movements occurred in the LMX. The axial hydraulic conductivity (Kh) was measured in leaf sections from all these regions and compared with the theoretical axial hydraulic conductivity (Kt) computed from the diameter of individual inner vessels. Kt was proportional to Kh throughout the leaf, but Kt was about three times Kh. The changes in Kh and Kt along the leaf reflected the different stages of xylem maturation. In the basal 60 mm region, Kh was about 0.30±0.07 mmol s−1 mm MPa−1. Beyond that region, Kh rapidly increased with metaxylem element maturation to a maximum value of 5.0±0.3 mmol s−1 mm MPa−1, 105 mm from the leaf base. It then decreased to 3.5±0.2 mmol s−1 mm MPa−1 near the leaf tip. The basal expanding region was observed to restrict longitudinal water movement. There was a close relationship between the water deposition rate in the elongation zone and the sum of the perimeters of PX vessels. The implications of this longitudinal vasculature on the partitioning of water between growth and transpiration is discussed.

Roots of angiosperms differ in the behaviour of their apical meristems, and this diversity has been studied through the patterns of differentiation in a range of families. Except for three families of the Nymphaeales, all dicotyledons derive the epidermis wholly or partially with the root cap. In closed meristems the inner cell layer of the cap complex forms the epidermis; in open meristems, where cortex and cap are intermittently of common origin, linkages between the epidermis and the cap predominate. Roots of most monocotyledons derive the epidermis as the outer layer of the cortex. Nymphaeaceae, Cabombaceae and Barclayaceae resemble monocotyledons in this respect, and this applies even in the species with open meristems, in which a cleft forms between the cap and the rest of the apex. This feature fits with the close relationship between the Nymphaeales and monocotyledons revealed by other data. But none of the other Magnoliidae with close links to Liliidae has this type of epidermis. The Nymphaeales are also almost unique among dicotyledons in having trichoblasts in a vertical pattern. This results from transverse divisions of epidermal cells giving two conspicuously different daughters. The smaller and most densely staining one later bears a root hair. This is the way trichoblasts develop in many monocotyledons. The only other dicotyledons found with trichoblasts in vertical patterns are in the Piperaceae, though their closed meristems derive the epidermis in the normal dicotyledon pattern. All other dicotyledons that bear trichoblasts develop them in radial patterns probably related to the radial disposition of cells across the cortex. This pattern is more widespread than was previously known, though it can occur only in roots with closed meristems. In plants of both subclasses with vertical patterns of trichoblasts, it is usually the proximal daughter of the initiating mitosis which becomes the trichoblast. But it is the distal daughter in Juncaceae, Restionaceae, Cyperaceae and Poaceae. Vertical patterns differ further as to whether the atrichoblast daughters divide after initiation, thus changing the alternation of trichoblasts and atrichoblasts along a file of epidermal cells. Monocotyledons and the Nymphaeaceae can produce trichoblasts in open or closed meristems. In some embryos and root primordia the epidermis becomes discrete from cortex and cap and remains so in the meristems. This has now been found in six aquatic genera of the Liliidae, but reports of it occurring elsewhere are probably due to misinterpretation of sections. Discrete epidermises differentiate into trichoblasts and atrichoblasts in Hydrocharitaceae, but not in Araceae or Lemnaceae.

Increased leaf phosphorus (P) concentration improved the water-use efficiency (WUE) and drought tolerance of regularly defoliated white clover plants by decreasing the rate of daily transpiration per unit leaf area in dry soil. Night transpiration was around 17% of the total daily transpiration. The improved control of transpiration in the high-P plants was associated with an increased individual leaf area and WUE that apparently resulted from net photosynthetic assimilation rate being reduced less than the reductions in the transpiration (27% vs 58%). On the other hand, greater transpiration from low-P plants was associated with poor stomatal control of transpirational loss of water, less ABA in the leaves when exposed to dry soil, and thicker and smaller leaf size compared with high-P leaves. The leaf P concentration was positively related with leaf ABA, and negatively with transpiration rates, under dry conditions (P < 0.001). However, leaf ABA was not closely related to the transpiration rate, suggesting that leaf P concentration has a greater influence than ABA on the transpiration rates.

Two-yr-old Scots pine (Pinus sylvestris) seedlings were grown in sand culture for 1 yr with a generous supply of a balanced nutrient solution. Trees were repotted into clean sand in February 1998 and given either a reduced or adequate nutrient supply containing enriched 15N, 41K and 26Mg to label nutrient uptake during spring 1998. Trees doubled their biomass during the experiment. Whole-tree net photosynthesis was reduced by 43% after 95 d in trees that received the lower nutrient supply (P < 0.001), although differences in biomass between the two treatments were less pronounced. Remobilization contributed 83, 82 and 52% of the N, K and Mg, respectively, used to support growth of new tissues in trees that received reduced nutrient supply. Those receiving the higher nutrient supply still obtained 44–59% of nutrients used for spring growth of new tissues from remobilization. Current nutrient supply had no significant effect on the amount of N or Mg remobilized to new tissues but K remobilization was less in trees that received the lower nutrient supply (P = 0.025). The importance of remobilization in young trees and problems associated with quantifying internal cycling of nutrients are discussed.

Hordeum spontaneum shows a large genetic variation and occupies a wide range of different habitats. The aim of this study was to quantify variation in growth characteristics of H. spontaneum from different sites in Israel and to relate this variation to different environmental conditions. To this end, 84 accessions of 21 populations were grown in a growth chamber in near-optimal conditions and a range of physiological, morphological, allocation- related and chemical characteristics were measured. These parameters included rates of photosynthesis, shoot and root respiration, specific leaf area, biomass allocation and seed mass. Averaged over all traits variation explained by differences between populations was 26%, between accessions 21%, whereas that within accessions was 53%. By contrast with most genetic studies, we found variation between populations larger than between accessions. The largest between-population variation (46%) was for morphological traits. In particular, seed mass, leaf thickness and leaf width differed strongly between populations. Variation in growth characteristics between populations was poorly related to mean annual rainfall, mean humidity or January temperature at the sites of origin. We expect that differences between populations to be larger and correlation with environmental parameters stronger in plants grown in stressful conditions. According to our study, seed mass is more important than relative growth rate in determining variation in early plant biomass in H. spontaneum.

Osmotic adjustment (OA) and increased cell-wall extensibility required for expansive leaf growth are well defined components of adaptation to water stress in dry soil, which might interact with soil phosphorus (P) concentration and defoliation frequency for intensively grazed white clover in legume-based pastures. Experiments were conducted with frequently and infrequently defoliated mini-swards of white clover growing in dry soil with low and high P concentrations. The higher yielding high-P plants were able to dry the soil to greater soil water suctions; their leaves had lower water potential values, yet they showed fewer water stress symptoms and underwent a more complete recovery from the water stress symptoms on rewatering, than the low-P plants. High- P plants had greater OA, proline concentration and leaf expansion rate. On the other hand, low-P plants showed an increased osmotic concentration when there was no change in the total solute content per unit of leaf d. wt, indicating more loss of water from the leaf tissue. The key measures that appeared to be directly associated with plant recovery over a short period following water stress were increased proline concentration and leaf expansion rate, probably resulting from increased cell-wall extensibility rather than increased production of cells for the high-P plants.

Parameters of convective ventilation and also the content and composition of free amino acids and free carbohydrates in roots, basal culm internodes and leaves were investigated in five stands of Phragmites australis that were subjected to different growth conditions and salinity levels. Shoot length, stem diameter and productivity decreased with increasing salinity. These morphometric variations in turn affected parameters of convective ventilation and amino acid patterns. Phragmites populations growing at the highest salinity levels possessed the lowest rhizome conductivities, which was probably caused by their more pronounced xeromorphic growth. High counterpressures of the rhizome system and low stem diameters were limiting factors for convective flows. The Phragmites stand with lowest flow rate and ventilation efficiency had the highest percentage of alanine, serine and γ-aminobutyric acid (55% of total free amino acids) in its roots, indicating hypoxic stress in basal plant parts. Increasing salinity was connected with significantly higher fractions of glutamine (in basal culm internodes), glutamate (in roots) and proline (in both) as well as slightly increasing osmolality of plant sap. However, total contents of free amino acids and carbohydrates were low, and not related to either salinity level or osmolality of plant sap. This suggests that they might not have an important, direct role as an osmolyte in P. australis. Stand-specific variations in amino acid and carbohydrate patterns of leaves were negligible. Ratios of amino acids to carbohydrate were positively related to potential N supply. It is concluded that (1) salinity affects convective rhizome ventilation by varying the reed morphometry, and (2) the pattern of free amino acids responds characteristically to conditions of salinity and hypoxic growth.

The effects of ambient and elevated ozone levels on growth and photosynthesis of beech (Fagus sylvatica) were studied by exposing seedlings in open-top chambers for one growing season to three treatments: charcoal-filtered (CF), non-filtered (NF) and non-filtered air with addition of ozone (30 ppb ozone) on clear days for 8–10 h d−1 (NF +). Ambient levels were relatively low and accumulated to an AOT40 (accumulated exposure over a threshold of 40 ppb) of 4055 ppb h (for the period 23 Apr–30 Sept). The NF + chambers received an AOT40 of 8880 ppb h. Throughout the growing season we measured growth and photosynthetic properties. The treatments did not cause strong effects: measurements of gas exchange (light-saturated assimilation rate, CO2 and light-response curves) and chlorophyll fluorescence showed slight and mostly non-significant reductions of several parameters. No significant differences were found for growth, though in the NF + treatment (AOT40 8880 ppb h) the relative growth rate for diameter increment was at times reduced by 12% compared with the control treatment.