Growth responses of Scots pine to climatic factors on reclaimed oil shale mined land

Afforestation on reclaimed mining areas has high ecological and economic importance. However, ecosystems established on post-mining substrate can become vulnerable due to climate variability. We used tree-ring data and dendrochronological techniques to study the relationship between climate variables and annual growth of Scots pine (Pinus sylvestris L.) growing on reclaimed open cast oil shale mining areas in Northeast Estonia. Chronologies for trees of different age classes (50, 40, 30) were developed. Pearson’s correlation analysis between radial growth indices and monthly climate variables revealed that precipitation in June–July and higher mean temperatures in spring season enhanced radial growth of pine plantations, while higher than average temperatures in summer months inhibited wood production. Sensitivity of radial increment to climatic factors on post-mining soils was not homogenous among the studied populations. Older trees growing on more developed soils were more sensitive to precipitation deficit in summer, while growth indices of two other stand groups (young and middle-aged) were highly correlated to temperature. High mean temperatures in August were negatively related to annual wood production in all trees, while trees in the youngest stands benefited from warmer temperatures in January. As a response to thinning, mean annual basal area increment increased up to 50 %. By managing tree competition in the closed-canopy stands, through the thinning activities, tree sensitivity and response to climate could be manipulated.     

Interactive effects of CO2 and O3 on a ponderosa pine plant/litter/soil mesocosm.

To study individual and combined impacts of two important atmospheric trace gases, CO2 and O3, on C and N cycling in forest ecosystems; a multi-year experiment using a small-scale ponderosa pine (Pinus ponderosa Laws.) seedling/soil/litter system was initiated in April 1998. The experiment was conducted in outdoor, sun-lit chambers where aboveground and belowground ecological processes could be studied in detail. This paper describes the approach and methodology used, and presents preliminary data for the first two growing seasons. CO2 treatments were ambient and elevated (ambient + 280 ppm). O3 treatments were elevated (hourly averages to 159 ppb, cumulative exposure > 60 ppb O3, SUM 06 approximately 10.37 ppm h), and a low control level (nearly all hourly averages <40 ppb. SUM 06 approximately 0.07 ppm h). Significant (P < 0.05) individual and interactive effects occurred with elevated CO2 and elevated O3. Elevated CO2 increased needle-level net photosynthetic rates over both seasons. Following the first season, the highest photosynthetic rates were for trees which had previously received elevated O3 in addition to elevated CO2. Elevated CO2 increased seedling stem diameters, with the greatest increase at low O3. Elevated CO2 decreased current year needle % N in the summer. For 1-year-old needles measured in the fall there was a decrease in % N with elevated CO2 at low O3, but an increase in % N with elevated CO2 at elevated O3. Nitrogen fixation (measured by acetylene reduction) was low in ponderosa pine litter and there were no significant CO2 or O3 effects. Neither elevated CO2 nor elevated O3 affected standing root biomass or root length density. Elevated O3 decreased the % N in coarse-fine (1-2 mm diameter) but not in fine (< 1 mm diameter) roots. Both elevated CO2 and elevated O3 tended to increase the number of fungal colony forming units (CFUs) in the AC soil horizon, and elevated O3 tended to decrease bacterial CFUs in the C soil horizon. Thus, after two growing seasons we showed interactive effects of O3 and CO2 in combination, in addition to responses to CO2 or O3 alone for a ponderosa pine plant/litter/soil system.     

Variable emissions of microbial volatile organic compounds (MVOCs) from root-associated fungi isolated from Scots pine

Soils emit a large variety of volatile organic compounds. In natural ecosystems, measurements of microbial volatile organic compound (MVOC) exchange rates between soil and atmosphere are difficult due to e.g. the spatial heterogeneity of the belowground organisms, and due to the many potential sources for the same compounds. We measured in laboratory conditions the MVOC emission rates and spectra of eight typical fungi occurring in boreal forest soils. The studied species are decomposers (Gymnopilus penetrans, Ophiostoma abietinum), ectomycorrhizal (Cenococcum geophilum, Piloderma olivaceum, Suillus variegatus, Tomentellopsis submollis) and endophytic fungi (Meliniomyces variabilis, Phialocephala fortinii). The MVOC emissions contained altogether 21 known and 6 unidentified compounds whose emission rates were >0.1 μg g(DW)^?1 h^?1. The most abundant compounds were the short-chain carbonyl compounds (acetone and acetaldehyde). The greatest carbonyl emissions were measured from P. olivaceum (1.9 mg acetone g(DW)^?1 h^?1) and P. fortinii (0.114 mg acetaldehyde g(DW)^?1 h^?1). Terpenoid emissions (isoprene, mono- and sesquiterpenes) were detected from some fungal cultures, but in relatively small amounts. We conclude that soil micro-organisms can potentially be responsible for significant emissions of volatiles, especially short-chain oxygenated compounds, to the below-canopy atmosphere.     

Response of needle sulphur and nitrogen concentrations of Scots pine versus Norway spruce to SO2 and NO2

The results of two field studies and an open-top chamber fumigation experiment showed that the response of mature Scots pine to SO(2) and NO(2) differed from that of mature Norway spruce. Moreover, the response of pine seedlings to SO(2) and NO(2) differed from that of mature trees. The greater increase in the needle total S concentrations of pine suggested more abundant stomatal uptake of SO(2) compared to spruce. Both pine seedlings and mature trees also seemed to absorb more N from atmospheric deposition. Mature pine was able to assimilate SO(4)(2-) derived from SO(2) into organic S more effectively than mature spruce at the high S and N deposition sites, whereas both pine and spruce seedlings accumulated SO(4)-S under NO(2)+SO(2) exposure. Spruce, in turn, accumulated SO(4)-S even when well supplied with N. Net assimilation of SO(4)(2-) in conifer seedlings was enhanced markedly by elevated temperature. To protect the northern coniferous forests against the harmful effects of S and N deposition, it is recommended that the critical level for SO(2) as a growing season mean be set at 5-10 microg m(-3) and NO(2) at 10-15 microg m(-3), depending on the 'effective temperature sum' and/or whether SO(2) and NO(2) occur alone or in combination.     

Ectomycorrhizal colonization of loblolly pine seedlings during three growing seasons in response to ozone, acidic precipitation, and soil Mg status

A three-year study was initiated in 1987 to evaluate the impact of O3, acidic precipitation, and soil Mg on ectomycorrhizal colonization of loblolly pine (Pinus taeda L.) seedlings. Thirty-six open-top chambers equipped with a rainfall exclusion/addition system were utilized to administer three levels of O3 (subambient, ambient, or twice ambient) and two precipitation acidity levels (pH 3.8 or 5.2) to seedlings growing in 24-liter plastic pots containing soil having either 35 or 15 mg kg(-1) of exchangeable Mg. Seedlings exposed to the twice ambient O3 treatment exhibited smaller percentages of total ectomycorrhizal short roots at the end of each year of the study, but trends were statistically significant in 1989 only. Changes in number of specific ectomycorrhizal morphotypes in response to O3 were not consistent from year to year. Acidic precipitation treatments had no effect on number or percent of mycorrhizal short roots, and responses of two morphotypes to soil Mg treatments were probably due to differences in the soil environment rather than a result of changes in aboveground processes. Temporal shifts in morphotype frequencies were observed for seedlings in all treatments and indicate that mycorrhizal succession occurred during the study period.     

Assessing the critical level of SO2 for Scots pine in situ

A field survey was performed in eastern Finland, where measured ambient SO2 concentrations were 1.4-3.8 microg m(-3) a(-1) and bulk S deposition 0.17-0.32 g m(-2) a(-1) in 1991-1993. The accumulation of sulphur (S) in needles of Scots pine (Pinus sylvestris L.) was studied with XRF, IC and FESEM analyses and the needle damage examined under a light microscope and by SEM. Foliar N concentrations were also measured. Foliar total S concentrations were observed to be above the normal S level (500-700 microg g(-1)) over almost the whole area. Slight chlorosis and/or necrosis of the needle tips and stomatal areas, changes in the needle surface waxes and localization of S into needle tips and mesophyll cells around the stomata suggested the impact of S deposition, as did the calculations of St/Nt, and 'predicted' and 'excess' S. A concentration of about 900 microg g(-1) may be considered a critical level for foliar St in areas with low N supply.     

Contribution of canopy leaching to sulphate deposition in a Scots pine forest

Radioactive sulphate (35SO4) was applied to the soil below a Scots pine forest on 23 June 1989, and its movement into the canopy and into throughfall and stemflow was measured over 4 months. The specific activity, Bq (mg S)(-1), of the canopy increased monotonically; uptake by current-year (1989) expanding needles was initially twice as fast as by older needles or live twigs. By 10 October the canopy average specific activity was 62 Bq (mg S)(-1). The specific activity of net throughfall (throughfall + stemflow - rain), deduced from measurements from six throughfall collectors, six stemflow collectors and two rain collectors, fell rapidly from 12.6 Bq (mg S)(-1) in late July to <1 Bq (mg S)(-1) in mid-August. The results suggest (assuming rapid equilibration of 35S with sulphate in soil) that root-derived sulphate contributed c. 3% of sulphate in net throughfall and that dry deposition of SO2 and sulphate particles contributed c. 97% of the 0.56 g S m(-2) measured in net throughfall over the period. Simultaneous measurements of SO2 at canopy height and of NH3 above and within the canopy gave mean concentrations of 5.9 and 0.86 microg m(-3), respectively, sufficient to account for the sulphate measured in net throughfall only if codeposition of NH3 and SO2 occurred to canopy surfaces. The large values of specific activity observed in July, however, indicate that throughfall composition may be closely related to recent soil input of sulphate, and that equilibrium cannot be safely assumed. The possibility of a significant contribution of soil-derived sulphate to sulphate deposition in net throughfall cannot be ruled out on the basis of this experiment.     

Seasonal soil and leaf CO2 exchange rates in a Mediterranean holm oak forest and their responses to drought conditions

We measured the soil and leaf CO₂ exchange in Quercus ilex and Phillyrea latifolia seasonally throughout the year in a representative site of the Mediterranean region, a natural holm oak forest growing in the Prades Mountains in southeastern Catalonia. In the wet seasons (spring and autumn), we experimentally decreased soil moisture by 30%, by excluding rainfall and water runoff in 12 plots, 1×10 m, and left 12 further plots as controls. Our aim was to predict the response of these gas exchanges to the drought forecasted for the next decades for this region by GCM and ecophysiological models.Annual average soil CO₂ exchange rate was 2.27±0.27 μmol CO₂ m⁻² s⁻¹. Annual average leaf CO₂ exchange rates were 8±1 and 5±1 μmol m⁻² s⁻¹ in Q. ilex and P. latifolia, respectively. Soil respiration rates in control treatments followed a seasonal pattern similar to photosynthetic activity. They reached maximum values in spring and autumn (2.5–3.8 μmol m⁻² s⁻¹ soil CO₂ emission rates and 7–15 μmol m⁻² s⁻¹ net photosynthetic rates) and minimum values (almost 0 for both variables) in summer, showing that soil moisture was the most important factor driving the soil microbial activity and the photosynthetic activity of plants. In autumn, drought treatment strongly decreased net photosynthesis rates and stomatal conductance of Q. ilex by 44% and 53%, respectively. Soil respiration was also reduced by 43% under drought treatment in the wet seasons. In summer there were larger soil CO₂ emissions in drought plots than in control plots, probably driven by autotrophic (roots) metabolism. The results indicate that leaf and soil CO₂ exchange may be strongly reduced (by ca. 44%) by the predicted decreases of soil water availability in the next decades. Long-term studies are needed to confirm these predictions or to find out possible acclimation of those processes.     

Forest floor CO2 flux estimated from soil CO2 and radon concentrations

Having a quantitative understanding of the carbon cycle in forests is of great importance for predicting global warming issues. Carbon dioxide production in soil is the largest CO₂ source in forests, and exhibits large temporal and spatial variations. Continuous observation of soil CO₂ flux at many sites over a forest is therefore necessary to obtain representative soil CO₂ fluxes for the forest. In this study, a gradient method to measure soil CO₂ flux indirectly from soil radon and CO₂ measurements was theoretically modified to conveniently measure the soil CO₂ flux from soil radon and CO₂ concentrations measured at one soil depth. To experimentally test the modified method, a field observation was conducted continuously in a forest over a 31-day period.Since changes in the soil water content near the soil surface were small throughout the observation, a constant effective diffusivity for CO₂ was assumed for the soil CO₂ flux estimation. The soil CO₂ flux was then calculated as the product of the effective diffusivity and the gradient of the soil CO₂ concentration, each calculated from soil radon and CO₂ concentrations. The estimated flux ranged from 1.9 to 5.8 μmol m^?2 s^?1, and, correlating well with the reference value, measured with a conventional ventilated-chamber method. We therefore conclude that the modified gradient method based on the measurement of soil CO₂ and radon concentration at one depth is reliable, at least under conditions where the change in the soil water content is small.     

Adaptation strategies and referencing trial of Scots and black pine populations subjected to heavy metal pollution

The impact of industrial heavy metal pollution on Scots pine (Pinus sylvestris L.) and black pine (Pinus nigra Arn.) populations was investigated. Sampled pine stands, which were located in Upper Silesia (southern Poland) in an area strongly polluted by heavy metals, consisted of resistant and sensitive trees. To evaluate the adaptation process, genetic structure and diversity was tested using isozyme analysis. Higher levels of Zn, Pb, Cd and Cu were detected in needles of sensitive trees compared with resistant ones. With respect to morphology, Scots pines were more distinctly impaired than black pines. Although black pines had lower heavy metal concentrations, levels in 1-year-old needles, other than Cu, significantly exceeded “reference plant” values (Markert 1994). In both species, resistant trees demonstrated a lower degree of genetic variation than metal-sensitive trees with respect to some enzyme loci (SHDH A, PGI, PGM, MDH C and DIA). This observation was corroborated in sensitive trees by the smaller number of identified alleles and alleles per locus, absence of private alleles and significant excess of homozygotes in relation to expected Hardy–Weinberg equilibrium values. Assuming that only resistant trees of both species survive under conditions of prolonged soil contamination, the observed genetic structure implies that remaining populations will be depleted of some alleles of unknown adaptive value to future selection pressures. Genetic changes induced by heavy metals suggest an important role for specific enzymes—FEST, SHDH A and B, GOT B and PGI—in the adaptation process. Our results may serve as a basis for selection and propagation of individuals appropriate for re-cultivation of areas chemically degraded by industrial activity.     

Interactions between precipitation and Scots pine canopies along a heavy-metal pollution gradient

Bulk precipitation and stand throughfall were collected during 1992-96 at distances of 0.5, 4 and 8 km from the Harjavalta Cu-Ni smelter, southwestern Finland. The amounts of heavy metals (Cu, Ni, Zn, Fe) and mineral nutrients in bulk precipitation and throughfall were highest at 0.5 km. Although the canopy coverage was low at 0.5 km, the amounts of heavy metals intercepted by the canopy were extremely high. The proportion of foliar leaching relative to the wash-off of dry deposition from the needle surfaces decreased on moving towards the smelter for all elements, except for K. The high rate of K leaching from the needle tissues close to the smelter demonstrated that the K throughfall flux has been greatly altered by the heavy pollution load.     

Combined use of open-air and indoor fumigation systems to study effects of SO2 on leaching processes in Scots pine litter

Both an open-air fumigation system and a laboratory-based system were used to expose decomposing Scots pine (Pinus sylvestris L.) needles to controlled concentrations of SO(2) (arithmetic mean 相似文献   

Total vs. internal element concentrations in Scots pine needles along a sulphur and metal pollution gradient

Analysis of foliar elements is a commonly used method for studying tree nutrition and for monitoring the impacts of air pollutants on forest ecosystems. Interpretations based on the results of foliar element analysis may, however, be different in nutrition vs. monitoring studies. We studied the impacts of severe sulphur and metal (mainly Cu and Ni) pollution on the element concentrations (Al, Ca, Cu, Fe, K, Mg, Mn, Ni, P, Pb, S and Zn) in Scots pine (Pinus sylvestris L.) foliage along an airborne sulphur and metal pollution gradient. Emphasis was put on determining the contribution of air-borne particles that have accumulated on needle surfaces to the total foliage concentrations. A comparison of two soil extraction methods was carried out in order to obtain a reliable estimate of plant-available element concentrations in the soil. Element concentrations in the soil showed only a weak relationship with internal foliar concentrations. There were no clear differences between the total and internal needle S concentrations along the gradient, whereas at the plot closest to the metal smelter complex the total Cu concentrations in the youngest needles were 1.3-fold and Ni concentrations over 1.6-fold higher than the internal needle concentrations. Chloroform-extracted surface wax was found to have Ni and Cu concentrations of as high as 3000 and 600 microg/g of wax, respectively. Our results suggest that bioindicator studies (e.g. monitoring studies) may require different foliar analysis techniques from those used in studies on the nutritional status of trees.     

Long term changes in atmospheric N and S throughfall deposition and effects on soil solution chemistry in a Scots pine forest in the Netherlands

In a Scots pine forest the throughfall deposition and the chemical composition of the soil solution was monitored since 1984. (Inter)national legislation measures led to a reduction of the deposition of nitrogen and sulphur. The deposition of sulphur has decreased by approximately 65%. The total mineral-nitrogen deposition has decreased by ca. 25%, which is mainly due to a reduction in ammonium-N deposition (−40%), since nitrate-N deposition has increased (+50%). The nitrogen concentration in the upper mineral soil solution at 10 cm depth has decreased, leading to an improved nutritional balance, which may result in improved tree vitality. In the drainage water at 90 cm depth the fluxes of NO₃⁻ and SO₄²⁻ have decreased, resulting in a reduced leeching of accompanying base cations, thus preserving nutrients in the ecosystem. It may take still several years, however, before this will meet the prerequisite of a sustainable ecosystem.     

Influence of seedling roots, environmental factors and soil characteristics on soil CO2 efflux rates in a 2-year-old loblolly pine (Pinus taeda L.) plantation in the Virginia Piedmont

To understand the role of managed forests in carbon sequestration an understanding of factors controlling soil CO2 efflux will be necessary. This study examined the influence of seedling roots, environmental factors, nutrient availability, and soil characteristics on soil CO2 efflux patterns in a 2-year-old pine plantation in the Virginia Piedmont. Efflux rates were measured both near the base of seedlings and midway between rows in plots that had received fertilization and mulch treatments in a factorial combination. Soil CO2 efflux rates were consistently higher near the base of seedlings, fertilization increased seedling growth with no significant effect on rates. and mulching increased winter efflux rates. In a regression analysis of seasonal soil CO2 efflux, soil temperature explained 42.2% of the variance followed by the interaction of soil temperature and moisture and of soil temperature and plot position, which together explained an additional 9.8% of the observed variance in seasonal rates. During March 2000 measurements, the spatial pattern of soil CO2 efflux between plots was most influenced by differences in soil nitrogen and pine root biomass. Furthermore, spatial differences observed in mean annual efflux rates were found to be highly influenced by the amount of soil coarse fragments in the upper soil profile.     

Organochlorine pollutants in Scots pine needles--biological and site related variation within a forest stand

To study the biological variation in the content of some persistent organic pollutants, viz hexachlorobenzene (HCB), alpha- and gamma-hexachlorocyclohexane (alpha- and gamma-HCH), and 1,1,1-trichloro-2,2-bis-(4-chlorophenyl)-ethane (DDT) were analysed in needles from Scots pine trees growing at an isolated peninsula south of Stockholm, Sweden. The concentration variations of each compound was evaluated by a nested analysis of variance (ANOVA) with sampling site specific compass direction, individual tree, and sampling height as factors. Two pair-wise post hoc tests were used to test significant results from the ANOVA. The hexachlorocyclohexanes showed no significant differences between sampling site, trees or sampling height. DDT concentration was significantly lower at the sampling site with the densest vegetation compared to at least three of the other sampling sites. HCB was significantly lower at the lowest sampling height (0.5 m).     

Plant responses to atmospheric CO2 enrichment with emphasis on roots and the rhizosphere

Empirical records provide incontestable evidence of global changes: foremost among these changes is the rising concentration of CO(2) in the earth's atmosphere. Plant growth is nearly always stimulated by elevation of CO(2). Photosynthesis increases, more plant biomass accumulates per unit of water consumed, and economic yield is enhanced. The profitable use of supplemental CO(2) over years of greenhouse practice points to the value of CO(2) for plant production. Plant responses to CO(2) are known to interact with other environmental factors, e.g. light, temperature, soil water, and humidity. Important stresses including drought, temperature, salinity, and air pollution have been shown to be ameliorated when CO(2) levels are elevated. In the agricultural context, the growing season has been shortened for some crops with the application of more CO(2); less water use has generally, but not always, been observed and is under further study; experimental studies have shown that economic yield for most crops increases by about 33% for a doubling of ambient CO(2) concentration. However, there are some reports of negligible or negative effects. Plant species respond differently to CO(2) enrichment, therefore, clearly competitive shifts within natural communities could occur. Though of less importance in managed agro-ecosystems, competition between crops and weeds could also be altered. Tissue composition can vary as CO(2) increases (e.g. higher C: N ratios) leading to changes in herbivory, but tests of crop products (consumed by man) from elevated CO(2) experiments have generally not revealed significant differences in their quality. However, any CO(2)-induced change in plant chemical or structural make-up could lead to alterations in the plant's interaction with any number of environmental factors-physicochemical or biological. Host-pathogen relationships, defense against physical stressors, and the capacity to overcome resource shortages could be impacted by rises in CO(2). Root biomass is known to increase but, with few exceptions, detailed studies of root growth and function are lacking. Potential enhancement of root growth could translate into greater rhizodeposition, which, in turn, could lead to shifts in the rhizosphere itself. Some of the direct effects of CO(2) on vegetation have been reasonably well-studied, but for others work has been inadequate. Among these neglected areas are plant roots and the rhizosphere. Therefore, experiments on root and rhizosphere response in plants grown in CO(2)-enriched atmospheres will be reviewed and, where possible, collectively integrated. To this will be added data which have recently been collected by us. Having looked at the available data base, we will offer a series of hypotheses which we consider as priority targets for future research.     

Relations between Scots pine needle element concentrations and decreased needle longevity along pollution gradients

Scots pine (Pinus sylvestris L.) shoots were sampled along transects near one urban pollution source and two smelters. Needle Mg, P and K concentrations decreased from the second to the fourth age class linearly with needle survival along the urban pollution gradient. Still, over 80% of the average concentration of these nutrients remained in the fourth needle age class. Decreased needle longevity was closely related to the increased heavy metal concentrations near the smelters. Near the urban pollution source, it was related to the increased annual needle mass and the increased needle nutrient concentrations. Decreased Mn accumulation along with needle age was detected near all pollution sources. Leaching of Mn from needles and especially from soil as a cause of decreased needle concentrations is discussed.     

Productivity responses of Acer rubrum and Taxodium distichum seedlings to elevated CO2 and flooding

Elevated levels of atmospheric CO2 are expected to increase photosynthetic rates of C3 tree species, but it is uncertain whether this will result in an increase in wetland seedling productivity. Separate short-term experiments (12 and 17 weeks) were performed on two wetland tree species, Taxodium distichum and Acer rubrum, to determine if elevated CO2 would influence the biomass responses of seedlings to flooding. T. distichum were grown in replicate glasshouses (n = 2) at CO2 concentrations of 350 or 700 ppm. and A. rubrum were grown in growth chambers at CO2 concentrations of 422 or 722 ppm. Both species were grown from seed. The elevated CO2 treatment was crossed with two water table treatments, flooded and non-flooded. Elevated CO2 increased leaf-level photosynthesis, whole-plant photosynthesis, and trunk diameter of T. distichum in both flooding treatments, but did not increase biomass of T. distichum or A. rubrum. Flooding severely reduced biomass, height, and leaf area of both T. distichum and A. rubrum. Our results suggest that the absence of a CO2-induced increase in growth may have been due to an O2 limitation on root production even though there was a relatively deep (approximately 10 cm) aerobic soil surface in the non-flooded treatment.     

Photosynthetic responses to temperature, light flux-density, CO2 concentration and vapour pressure deficit in Eucalyptus tetrodonta grown under CO2 enrichment

Seeds of Eucalyptus tetrodonta were sown under ambient or CO(2) enriched (700 microl litre(-1)) conditions in tropical Australia. Four sets of measurements were made, the first two after 12 months, on trees growing either in pots or planted in the ground. The third and fourth set were made after 18 and 30 months exposure to CO(2) enrichment, on trees growing in the ground. After 12 months exposure to CO(2) enrichment, the rate of light-saturated assimilation (Amax) of plants growing in the ground was determined. Responses of CO(2) assimilation to variations in leaf temperature, leaf-to-air vapour pressure deficit (LAVPD), light flux density and CO(2) concentration were also measured in the laboratory using plants growing in large pots. There was no significant difference in Amax between pot and ground located plants. Assimilation of E. tetrodonta was relatively insensitive to changes in LAVPD for both ambient and CO(2) enriched plants but the temperature optimum of assimilation was increased in plants grown and measured under CO(2) enrichment. Plants grown with CO(2) enrichment had an increased rate of light-saturated assimilation and apparent quantum yield was significantly increased by CO(2) enrichment. In contrast, carboxylation efficiency was decreased significantly by CO(2) enrichment. After 18 months growth with CO(2) enrichment, there was no sign of a decline in assimilation rate compared to measurements undertaken after 12 months. At low LAVPD values, assimilation rate was not influenced by CO(2) treatment but at moderate to high LAVPD, plants grown under CO(2) enrichment exhibited a larger assimilation rate than control plants. Specific leaf area and chlorophyll contents decreased in response to CO(2) enrichment, whilst foliar soluble protein contents and chlorophyll a/b ratios were unaffected by CO(2) treatment. Changes in soluble protein and chlorophyll contents in response to CO(2) enrichment did not account for changes in assimilation between treatments. After 30 months exposure to CO(2) enrichment, the rate of light-saturated assimilation was approximately 50% larger than controls and this enhancement was larger than that observed after 18 months exposure to CO(2) enrichment.