Vertical profiles reveal impact of ozone and temperature on carbon assimilation of Betula pendula and Populus tremula

Rising temperature and tropospheric ozone (O(3)) concentrations are likely to affect carbon assimilation processes and thus the carbon sink strength of trees. In this study, we investigated the joint action of elevated ozone and temperature on silver birch (Betula pendula) and European aspen (Populus tremula) saplings in field conditions by combining free-air ozone exposure (1.2?×?ambient) and infrared heaters (ambient +1.2 °C). At leaf level measurements, elevated ozone decreased leaf net photosynthesis (P(n)), while the response to elevated temperature was dependent on leaf position within the foliage. This indicates that leaf position has to be taken into account when leaf level data are collected and applied. The ozone effect on P(n) was partly compensated for at elevated temperature, showing an interactive effect of the treatments. In addition, the ratio of photosynthesis to stomatal conductance (P(n)/g(s) ratio) was decreased by ozone, which suggests decreasing water use efficiency. At the plant level, the increasing leaf area at elevated temperature resulted in a considerable increase in photosynthesis and growth in both species.     

Biochemical acclimation patterns of Betula pendula and Pinus sylvestris seedlings to elevated carbon dioxide concentrations

Acclimation of photosynthesis to increasing atmospheric carbon dioxide concentration ([CO2]; 350 to 2,000 micromol mol-1) was followed in silver birch (Betula pendula Roth.) and Scots pine (Pinus sylvestris L.) seedlings for two years. Chlorophyll fluorescence and concentrations of Rubisco, chlorophyll, total soluble protein and nitrogen were monitored together with steady-state gas exchange at three CO2 concentrations (ambient [CO2] (345 +/- 20 micromol mol-1), the growth [CO2] and 1950 +/- 55 micromol mol-1). Rubisco and chlorophyll concentrations decreased in birch and Scots pine with increasing growth [CO2]. A nonlinear response was recorded for Rubisco and chlorophyll concentrations in birch, which was correlated with a significant decrease in specific leaf area. Nitrogen concentration decreased in birch leaves, but was unchanged in Scots pine needles. The species differed substantially in their steady-state CO2 exchange response to increasing growth [CO2]. The principal effect in birch was a significant nonlinear decrease in the steady-state gas exchange rate at the ambient [CO2], whereas in Scots pine the main effect was a significant increase in the steady-state gas exchange rate at the growth [CO2].     

Ozone air pollution effects on tree-ring growth, delta(13)C, visible foliar injury and leaf gas exchange in three ozone-sensitive woody plant species

We assessed the effects of ambient tropospheric ozone on annual tree-ring growth, delta(13)C in the rings, leaf gas exchange and visible injury in three ozone-sensitive woody plant species in southern Switzerland. Seedlings of Populus nigra L., Viburnum lantana L. and Fraxinus excelsior L. were exposed to charcoal-filtered air (CF) and non-filtered air (NF) in open-top chambers, and to ambient air (AA) in open plots during the 2001 and 2002 growing seasons. Ambient ozone exposures in the region were sufficient to cause visible foliar injury, early leaf senescence and premature leaf loss in all species. Ozone had significant negative effects on net photosynthesis and stomatal conductance in all species in 2002 and in V. lantana and F. excelsior in 2001. Water-use efficiency decreased and intercellular CO(2) concentrations increased in all species in response to ozone in 2002 only. The width and delta(13)C of the 2001 and 2002 growth rings were measured for all species at the end of the 2002 growing season. Compared with CF seedlings, mean ring width in the AA and NF P. nigra seedlings was reduced by 52 and 46%, respectively, in 2002, whereas in V. lantana and F. excelsior, ring width showed no significant reductions in either year. Although delta(13)C was usually more negative in CF seedlings than in AA and NF seedlings, with the exception of F. excelsior in 2001, ozone effects on delta(13)C were significant only for V. lantana and P. nigra in 2001. Among species, P. nigra exhibited the greatest response to ozone for the measured parameters as well as the most severe foliar injury and was the only species to show a significant reduction in ring width in response to ozone exposure, despite significant negative ozone effects on leaf gas exchange and the development of visible foliar injury in V. lantana and F. excelsior. Thus, significant ozone-induced effects at the leaf level did not correspond to reduced tree-ring growth or increased delta(13)C in all species, indicating that the timing of ozone exposure and severity of leaf-level responses may be important in determining the sensitivity of tree productivity to ozone exposure.     

Mycorrhizal status of an ozone-sensitive poplar clone treated with the antiozonant ethylene diurea

The antiozonant ethylene diurea is proven to prevent growth reductions in forest trees induced by ozone. The community of mycorrhizal fungi could be useful indicator of environmental stress. In this study, response of mycorrhizal fungi and fine roots to a 4-year exposure to ambient ozone and treatment with antiozonant was investigated in ozone-sensitive poplar clone under field conditions. The community of ectomycorrhizal fungi and root length colonization with ectomycorrhizal, arbuscular mycorrhizal fungi, and root endophytic fungi was analyzed in antiozonant-treated poplar plants and in poplar plants irrigated with water. In general, plants protected by antiozonant showed higher total number of fine roots, number of ectomycorrhizal types, Shannon–Weaver diversity index, and Species richness index compared to the plants treated with water. The ectomycorrhizal community shifted from contact exploration type in the trees irrigated with water to short-distance exploration type in ethylene diurea-treated trees. Ozone protectant may beneficially affect the belowground community of mycorrhizal fungi colonizing roots of ozone-sensitive poplar clone.     

Midday depression of net photosynthesis in the tropical rainforest tree Eperua grandiflora: contributions of stomatal and internal conductances,respiration and Rubisco functioning

High midday temperatures can depress net photosynthesis. We investigated possible mechanisms underlying this phenomenon in leaves of Eperua grandiflora (Aubl.) Benth. saplings. This tropical tree establishes in small gaps in the rainforest canopy where direct sunlight can raise midday temperatures markedly. We simulated this microclimate in a growth chamber by varying air temperature between 28 and 38 degrees C at constant vapor pressure. A decrease in stomatal conductance in response to an increase in leaf-to-air vapor pressure difference (deltaW) caused by an increase in leaf temperature (Tleaf) was the principal reason for the decrease in net photosynthesis between 28 and 33 degrees C. Net photosynthesis decreased further between 33 and 38 degrees C. Direct effects on mesophyll functioning and indirect effects through deltaW were of similar magnitude in this temperature range. Mitochondrial respiration during photosynthesis was insensitive to Tleaf over the investigated temperature range; it thus did not contribute to midday depression of net photosynthesis. Internal conductance for CO2 diffusion in the leaf, estimated by combined gas exchange and chlorophyll fluorescence measurements, decreased slightly with increasing Tleaf. However, the decrease in photosynthetic rate with increasing Tleaf was larger and thus the difference in CO2 partial pressure between the substomatal cavity and chloroplast was smaller, leading to the conclusion that this factor was not causally involved in midday depression. Carboxylation capacity inferred from the CO2 response of photosynthesis increased between 28 and 33 degrees C, but remained unchanged between 33 and 38 degrees C. Increased oxygenation of ribulose-1,5-bisphosphate relative to its carboxylation and the concomitant increase in photorespiration with increasing Tleaf were thus not compensated by an increase in carboxylation capacity over the higher temperature range. This was the principal reason for the negative effect of high midday temperatures on mesophyll functioning.     

土壤干旱胁迫及复水对紫叶矮樱光合特性的影响

以盆栽3年生紫叶矮樱为试材,采用称重控水法,研究土壤干旱胁迫及复水对紫叶矮樱光合特性的影响。结果表明：干旱条件下,随着胁迫程度的加重,紫叶矮樱的叶水势、净光合速率、蒸腾速率、气孔导度和胞间CO2浓度下降,叶绿素、花青苷含量和水分利用效率先升高再降低;复水后,叶水势、净光合速率、蒸腾速率、叶绿素含量、花青苷含量和水分利用效率升高,表明紫叶矮樱具有胁迫后补偿生长特征。     

Effects of soil moisture regimes on growth and photosynthesis of the riparian plant Bolboschoenus planiculmis

Plants distributed in riparian regions experience frequent episodes of flooding and drought between years, and hence, riparian plants need to be floodand drought-tolerant. Riparian plants possess various traits to survive flooding, while their sensitivity to drought has received less attention. To investigate the growth and photosynthetic responses of a riparian species (Bolboschoenus planiculmis) to flooding and drought, plants of this species were subjected to 60-d flooding or drought stress under greenhouse conditions. Growth and photosynthetic traits were measured at the end of the treatments. As well, we determined the efficiency of photosynthetic apparatus in mature leaves. Plants of B. planiculmis adequately adjusted their growth and photosynthetic traits under both flooding and drought conditions. Flooding did not affect the above-ground growth of B. planiculmis. Increased growth of roots and rhizomes and the generation of new tubers suggested a high ability of below-ground lateral growth by capturing resources under flooding conditions. Enhanced photosynthetic capacity, retained leaf pigment concentrations and chlorophyll a fluorescence capacity indicated photosynthetic adaptation to flooding. In contrast, drought significantly decreased the above-ground growth of B. planiculmis, especially the leaves, thereby minimizing water loss due to transpiration. Its increased root to shoot ratio and "phalanx" asexual propagation pattern might enhance soil water uptake ability. Although the functional leaves of B. planiculmis could retain their leaf pigment concentrations, as well as photosynthesis and chlorophyll a fluorescence, the total biomass of plants decreased, which may be a consequence of the reduced leaf area, suggesting adverse effects by drought. Therefore, both growth and photosynthetic responses of B. planiculmis are likely to contribute to the ability of this species to thrive in riparian regions, but remain susceptive to drought.     

Carbon partitioning and allocation in northern red oak seedlings and mature trees in response to ozone

Northern red oak (Quercus rubra L.) seedlings and trees differ in their response to ozone. Previous work reported reductions in net photosynthesis, carboxylation efficiency and quantum yield of mature tree leaves, whereas seedling processes were unaffected by the same ozone exposure. To further characterize differences in ozone response between seedlings and mature trees, we examined carbon partitioning and allocation in 32-year-old trees and 4-year-old seedlings of northern red oak after exposure to subambient (seasonal SUM00 dose (sum of all hourly ozone exposures) = 31 ppm-h), ambient (SUM00 dose = 85 ppm-h) and twice ambient (SUM00 dose = 151 ppm-h) ozone concentrations for three growing seasons. For mature trees, ozone exposure decreased foliar starch partitioning, increased starch partitioning in branches and increased (14)C retention in leaves. In contrast, starch partitioning in leaves and branches, and foliar (14)C retention in seedlings were unaffected by ozone exposure, but soluble carbohydrate concentrations in coarse and fine roots of seedlings were reduced. Differences in carbohydrate demand between seedlings and mature trees may underlie the higher leaf ozone uptake rates and greater physiological response to ozone in mature northern red oak trees compared with seedlings.     

氮素对楸树无性系幼苗的生长影响研究

用沙培法培养楸树无性系I-4幼苗,进行4种供氮量（1 mmol/L、4 mmol/L、8 mmol/L、16 mmol/L）处理,对幼苗生长和部分生理性状进行试验。结果表明,对不同供氮量下苗木生长相比较,随着供氮量的增大,苗高呈先升高后降低的趋势,地径、叶片数、叶面积、茎、叶的生物量呈增大的趋势,而根冠比不断降低;叶绿素a、叶绿素b及类胡萝卜素含量升高,但蛋白质含量却不断降低。通过综合分析苗高、地径和生物量3个性状,确定楸树无性系I-4幼苗生长的最适氮浓度为16 mmol/L。     

Genotypic variation in growth and physiological responses of Finnish hybrid aspen (Populus tremuloides x P. tremula) to elevated tropospheric ozone concentration

Saplings of six Finnish hybrid aspen (Populus tremuloides Michx. x P. tremula L.) clones were exposed to 0, 50, 100 and 150 ppb ozone (O3) for 32 days in a chamber experiment to determine differences in O3 sensitivity among genotypes. Based on the chamber experiment, three clones with intermediate sensitivity to O3 were selected for a free-air O3 enrichment experiment in which plants were exposed for 2 months to either ambient air (control) or air containing 1.3 x the ambient O3 concentration. We measured stem height and radial growth, number of leaves, dry mass and relative growth rate of leaves, stem and roots, visible leaf injuries, net photosynthesis and stomatal conductance of the clones. There was high clonal variation in susceptibility to O3 in the chamber experiment, indicated by foliar injuries and differential reductions in growth and net photosynthesis. In the free-air O3 enrichment experiment, ozone caused a shift in resource allocation toward stem height growth, thereby altering the shoot to root balance. In both experiments, low O3 concentrations tended to stimulate growth of most clones, whereas 100 and 150 ppb O3 in the chamber experiment impaired growth of most clones. However, growth of the most O3-tolerant clone was not significantly affected by any O3 treatment.     

Water stress, photosynthesis and early growth patterns of cuttings of three Populus clones

Photosynthetic attributes, leaf area and early root growth patterns were studied in three Populus clones to identify traits associated with superior growth potential on sites where water could be a limiting factor. It was found that early root growth and superior leaf area production were more closely related to growth potential than were photosynthetic capacity or carboxylation efficiency. A hybrid clone of Populus nigra var. charkowiensis (syn. P. nigra var. plantierensis) x P. nigra cv. 'Incrassata' (NE308) had more leaf area production and greater root system development in both wet and dry soil than did a P. trichocarpa clone (T6) and a P. balsamifera clone (B3). Despite greater above- and below-ground productivity, plants of clone NE308 had significantly lower photosynthetic capacity and carboxylation efficiency and a slightly higher CO(2) compensation point than plants of clones T6 and B3. Rapid early leaf and root growth appear to be key attributes associated with productivity in these clones regardless of soil water availability.     

Genotypic variation in drought response of silver birch (Betula pendula): leaf water status and carbon gain

To assess genotypic variation in drought response of silver birch (Betula pendula Roth), we studied the plasticity of 16 physiological traits in response to a 12-14-week summer drought imposed on four clones in two consecutive years. In a common garden experiment, 1-year-old clonal trees from regions with low (550 mm year(-1)) to high rainfall (1270 mm year(-1)) were grown in 45-l pots, and leaf gas exchange parameters, leaf water potentials, leaf osmotic potentials and leaf carbon isotope signatures were repeatedly measured. There were no clonal differences in leaf water potential, but stomatal conductance (gs), net photosynthesis at ambient carbon dioxide concentration, photosynthetic water-use efficiency, leaf carbon isotope composition (delta13C) and leaf osmotic potentials at saturation (Pi0) and at incipient plasmolysis (Pip) were markedly influenced by genotype, especially gs and osmotic adjustment. Genotypes of low-rainfall origin displayed larger osmotic adjustment than genotypes of high-rainfall origin, although their Pi0 and Pip values were similar or higher with ample water supply. Genotypes of low-rainfall origin had higher gs than genotypes of high-rainfall origin under both ample and limited water supply, indicating a higher water consumption that might increase competitiveness in drought-prone habitats. Although most parameters tested were significantly influenced by genotype and treatment, the genotype x treatment interactions were not significant. The genotypes differed in plasticity of the tested parameters and in their apparent adaptation to drought; however, among genotypes, physiological plasticity and drought adaptation were not related to each other. Reduction of gs was the first and most plastic response to drought in all genotypes, and allowed the maintenance of high predawn leaf water potentials during the drought. None of the clones exhibited non-stomatal limitation of photosynthesis. Leaf gs, photosynthetic capacity, magnitude of osmotic adjustment and delta13C were all markedly lower in 2000 than in 1999, indicating root limitation in the containers in the second year.     

生长抑制剂对大叶黄杨形态及光合作用的影响

目的]探究抑制剂对大叶黄杨生长的抑制作用以及对其叶片形态和光合作用的影响,为灌木绿篱的化学修剪提供技术指导。[方法]在北京林业大学林场苗圃采用3种生长抑制剂(多效唑(PP_(333))、三碘苯甲酸(TIBA)以及脱落酸(ABA))对密植成绿篱状的1年生大叶黄杨扦插苗进行叶面喷施,对其生长、叶片形态及光合作用等指标进行测定。[结果]3种抑制剂均有矮化植株、抑制新梢生长的作用,矮化效果最佳、抑制作用最强的为PP_(333),且高浓度PP_(333)对于高生长的抑制作用持效性较长。叶宽、叶厚以及叶面积在短期PP_(333)处理下高于对照。PP_(333)能提高大叶黄杨净光合速率,主要通过增加叶厚、气孔导度、叶肉导度及叶绿素含量来实现,并且PP_(333)使蒸腾速率提高的同时降低了水分利用效率。TIBA有显著减小叶长、叶宽的作用,但能使叶厚增加,且随浓度的增加作用效果增强,主要通过减小叶面积,降低叶肉导度、气孔导度、胞间CO_2浓度及叶绿素含量来降低净光合速率。ABA能增加叶面积(10 mg·L~(-1)处理除外)、叶长以及减小叶片厚度,通过抑制气孔导度、叶肉导度、叶绿素含量降低净光合速率。TI-BA及ABA均通过降低蒸腾速率,使水分利用效率提高。[结论]3种抑制剂均有矮化植株、抑制新梢生长的作用,并且影响叶片发育,进而影响叶片光合作用,且800 mg·L~(-1)的多效唑对大叶黄杨具有较好的正向作用。     

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.     

光叶子花不同叶位叶片叶绿素含量和光合作用研究

本文以光叶子花炼苗60 d组培苗为材料,对光叶子花不同叶位叶绿素含量、呼吸作用(Re)、气孔导度(Gs)、胞间CO2浓度(Ci)、净光合速(Pn)、蒸腾速率(Tr)、叶面饱和水汽压亏缺(Vpdl)等进行了测定。试验结果表明,光叶子花叶片单位重量和单位面积叶绿素含量、净光合速度和水分利用效率(WUE)均随叶位上升呈先增大后减小趋势变化,其最大值都出现在第6叶位叶片;叶片呼吸速率、气孔导度和蒸腾速率均随叶位上升而减小;叶片胞间CO2浓度随叶位上升呈先减小后增大变化;叶面饱和水汽压亏缺随叶位上升而增大。由于光叶子花中部叶片叶绿素含量、Pn和WUE相对较高,因此在今后的栽培管理中应对中部叶片加强保护。     

油茶光合作用及其影响因素研究进展

综述了光合作用及其影响因素的研究进展,影响因素包括内部因素(品种特性、叶龄、叶绿素、叶面积指数)和外界因素(光照、水分、温度、CO_2体积分数)．概述了油茶不同品种、不同叶龄叶片的光合作用规律及提高油茶光合作用的途径的研究进展;探讨了油茶光合作用的研究方向．     

How does drought tolerance compare between two improved hybrids of balsam poplar and an unimproved native species?

Poplars are one of the woody plants that are very sensitive to water stress, which may reduce the productivity of fast-growing plantations. Poplars can exhibit several drought tolerance strategies that may impact productivity differently. Trees from two improved hybrids, Populus balsamifera?×?Populus trichocarpa Torr. & Gray (clone B?×?T) and P. balsamifera?×?Populus maximowiczii A. Henry (clone B?×?M), having P. balsamifera L. as a parent and trees from native and unimproved P. balsamifera were subjected to a 1-month drying cycle in a growth chamber and then rewatered. The unimproved and native B clone maintained higher stomatal conductance (g(s)) than the hybrids, and high photosynthetic activity and transpiration, even when soil water content was nearly zero. As a result, both instantaneous water use efficiency (WUE(i)) and leaf carbon isotope composition (δ(13)C) indicated that this clone was less affected by drought than both hybrids at maximal drought stress. However, this clone shed its leaves when the drought threshold was exceeded, which implied a greater loss of productivity. The B?×?M hybrid showed a relatively conservative response to water stress, with the greatest decrease in transpiring versus absorbing surface (total leaf area to root biomass ratio). This clone was also the only one to develop new leaves after rewatering, and its total biomass production was not significantly decreased by drought. Among the two hybrids, clone B?×?T was the most vigorous, with the greatest transpiration (E(i)) and net CO(2) assimilation (A) rates, allowing for high biomass production. However, it had a more risky strategy under drought conditions by keeping its stomata open and high E(i) rates under moderate drought, resulting in a lower recovery rate after rewatering. The opposite drought response strategies of the two hybrids were reflected by clone B?×?T having lower WUE(i) values than clone B?×?M at maximal drought, with a very low Ψ(min) value of -3.2 MPa, despite closed stomata and stopped photosynthetic activity. Positive linear relationships between A and g(s) for the three hybrids indicated strong stomatal control of photosynthesis. Moreover, the three poplar clones showed anisohydric behaviour for stomatal control and their use under long-term drought should be of interest, especially the B?×?M clone.     

竹节树无性系叶片生长特性与叶绿素荧光

对优选的6个竹节树(Carallia brachiata)无性系叶片性状(叶片干质量、叶面积、比叶重)、叶绿素相对含量和叶绿素荧光参数进行测定,并作单因素方差分析和聚类分析.结果表明:竹节树不同无性系叶片的干质量和叶面积差异不显著,叶绿素相对含量具有显著的差异性,不同无性系具有不同的光能吸收和利用能力;不同的无性系其叶绿素荧光的部分参数(Fo、Fv/Fm、Fv/Fo、ФPSⅡ、ETR)差异显著;竹节树无性系C-15植株与其他无性系相比具有较高水平的实际光化学效率(ΦPSⅡ)值,而非化学猝灭系数(qN)较低,同时具有较高的表观电子传递速率(ETR),表明竹节树无性系C-15有较高的光呼吸;相关分析表明,叶片生长较好的无性系其叶绿素相对含量较高,同时也具有较高的PSⅡ光化学效率;无性系C-15号具有较高的叶绿素相对含量及较强的光化学特性,可进行大力推广研发.     

Leaf photosynthetic characteristics of silver birch during three years of exposure to elevated concentrations of CO2 and O3 in the field

Effects of elevated concentrations of carbon dioxide ([CO2]) and ozone ([O3]) on photosynthesis and related biochemistry of two European silver birch (Betula pendula Roth) clones were studied under field conditions during 1999-2001. Seven-year-old trees of Clones 4 and 80 were exposed for 3 years to the following treatments in an open-top chamber experiment: outside control (OC), chamber control (CC), 2x ambient [CO2] (EC), 2x ambient [O3] (EO) and 2x ambient [CO2] + 2x ambient [O3] (EC+EO). During the experiment, gas exchange, chlorophyll fluorescence, amount and activity of Rubisco, concentrations of chlorophyll, soluble protein, soluble sugars, starch, nitrogen (N) and carbon:nitrogen (C:N) ratio were determined in short- and long-shoot leaves. Elevated [CO2] increased photosynthetic rate by around 30% when measurements were made at the growth [CO2]. When measured at ambient [CO2], photosynthesis was around 15% lower in EC trees than in CC trees. This was related to a approximately 10% decrease in total leaf N, to 26 and 20% decreases in the amount and activity of Rubisco, respectively, and to a 49% increase in starch concentration in elevated [CO2]. Elevated [O3] had no significant effect on gas exchange parameters and its effect on biochemistry was small in both clones. However, elevated [O3] decreased the proportion of Rubisco in total soluble proteins and the apparent quantum yield of photosystem II (PSII) photochemistry in light and increased non-photochemical quenching in 2000. The interactive effect of CO2 and O3 was variable. Elevated [O3] decreased chlorophyll concentration only in EO trees, and the EC+EO treatment decreased the total activity of Rubisco and increased the C:N ratio more than the EO treatment alone. The small effect of elevated [O3] on photosynthesis indicates that these young silver birches were fairly tolerant to annual [O3] exposures that were 2-3 times higher than the AOT40 value of 10 ppm.h. set as a critical dose for forest trees.     

Ozone impairs autumnal resorption of nitrogen from birch (Betula pendula) leaves, causing an increase in whole-tree nitrogen loss through litter fall

Saplings of one half-sib family of birch, Betula pendula Roth, were exposed to three ozone concentrations (non-filtered air (NF); non-filtered air + 10-20 nmol O(3) mol(-1) (NF+); non-filtered air + 40-60 nmol O(3) mol(-1) (NF++)) in open-top chambers during two growing seasons from 1997 to 1998. Shed leaves were collected regularly during both growing seasons and, in 1998, the dry mass (DM) and nitrogen (N) concentrations ([N]) of the shed leaves were measured to quantify the total amount of N lost through litter fall. Dry mass and [N] were also determined in mid-August for attached, mature and non-senescent leaves, in order to estimate autumnal leaf N resorption efficiency and proportional leaf DM decrease. Net photosynthetic capacity was measured during August and September 1998, in a population of leaves that emerged in mid-July. Photosynthesis declined with increasing leaf age in the NF++ treatment, whereas it remained high throughout the measurement period in the NF and NF+ treatments. In both years, leaves abscised prematurely in the NF++ treatment, whereas this effect was only significant in 1998 in the NF+ treatment. There was a strong linear relationship between proportional leaf shedding and daylight ozone exposure above a threshold of 40 nmol mol(-1) (daylight AOT40) during the growing season. The resorption of N was significantly impaired by ozone, and the smaller autumnal decrease in leaf DM in elevated ozone concentrations suggested that the bulk resorption of leaf DM was also inhibited. Nitrogen resorption efficiencies were 81, 73 and 63% and leaf mass decreases were 45, 36 and 30% in the NF, NF+ and NF++ treatments, respectively. Compared with the NF treatment, total N loss through litter fall was increased by 16 and 122% in the NF+ and NF++ treatments, respectively. We conclude that ozone impaired N resorption from birch leaves before abscission, causing a substantial increase in whole-tree N loss through litter fall.