Photosynthetic productivity of aspen clones varying in sensitivity to tropospheric ozone

Rooted cuttings from three aspen (Populus tremuloides Michx.) clones (216, 271 and 259, classified as high, intermediate and low in O(3) tolerance, respectively) were exposed to either diurnal O(3) profiles simulating those of Michigan's Lower Peninsula (episodic treatments), or diurnal square-wave O(3) treatments in open-top chambers in northern Michigan, USA. Ozone was dispensed in chambers ventilated with charcoal-filtered (CF) air. In addition, seedlings were compared to rooted cuttings in their response to episodic O(3) treatments. Early in the season, O(3) caused decreased photosynthetic rates in mature leaves of all clones, whereas only the photosynthetic rates of recently mature leaves of the O(3)-sensitive Clone 259 decreased in response to O(3) exposure. During midseason, O(3) caused decreased photosynthetic rates of both recently mature and mature leaves of the O(3)-sensitive Clone 259, but it had no effect on the photosynthetic rate of recently mature leaves of the O(3)-tolerant Clone 216. Late in the season, however, photosynthetic rates of both recently mature and mature leaves of Clone 216 were lower than those of the control plants maintained in CF air. Ozone decreased the photosynthetic rate of mature leaves of Clone 271, but it increased or had no effect on the photosynthetic rate of recently mature leaves. Photosynthetic response patterns of seedlings to O(3) treatment were similar to those of the clones, but total magnitude of the response was less, perhaps reflecting the diverse genotypes of the seedling population. Early leaf abscission was observed in all clones exposed to O(3); however, Clones 216 and 259 lost more leaf area than Clone 271. By late August, leaf area in the highest O(3) treatment had decreased relative to the controls by 26, 24 and 9% for Clones 216, 259 and 271, respectively. Ozone decreased whole-tree photosynthesis in all clones, and the decrease was consistently less in Clone 271 (23%) than in Clones 216 (56%) and 259 (56%), and was accompanied by declines in total biomass of 19, 28 and 47%, respectively. The relationship between biomass and whole-tree photosynthesis indicates that the negative impact of O(3) on biomass in the clones was determined largely by lower photosynthetic productivity of the foliage, rather than by potential changes in the carbon relations of other plant organs.     

Diurnal changes in leaf chemical constituents and (14)C partitioning in cottonwood

Diurnal changes in concentrations of leaf chemical fractions and partitioning of photosynthetically fixed (14)C within the plant and among chemical fractions were studied in rapidly growing cottonwood (Populus deltoides Bartr. ex Marsh.) seedlings. During the light period, leaf weight (mg cm(-2)) increased by about 25% primarily as a result of the accumulation of starch and sucrose, and to a lesser extent because of an increase in the content of amino acids and the chloroform fraction (pigments plus lipids). In contrast, reducing sugars and organic acids decreased in concentration. The partitioning of (14)C within the plant also changed during the light period. Acropetal transport to developing leaves and stem decreased from 81 to 55% of the total (14)C translocated from a source leaf in 4 hours, whereas basipetal transport to stem and roots increased from 13 to 37%. Although assimilation rate ((14)C fixed in 0.5 h) remained constant during the light period, the percentage of fixed (14)C translocated out of the source leaf in 4 h decreased from 27 to 9%. This change in transport rate of recently fixed (14)C was caused by a shift in (14)C partitioning from transport sucrose to storage starch. During the light period, the incorporation ratio ((14)C-sugar/(14)C-starch) decreased from 40 at 0700 h to 2 at 1900 h. The partitioning of carbon to different chemical fractions within the source leaf and the interactions or feedback between different sinks and the source leaf have a major influence on plant growth and development. Control of this carbon partitioning is located in both source and sink leaves.     

Distribution of (14)C-labeled photosynthate in loblolly pine (Pinus taeda) seedlings as affected by season and time after exposure

Distribution of (14)C-labeled photosynthate was determined in field-grown loblolly pine (Pinus taeda L.) seedlings on August 9 and October 15, 1984 and January 15 and March 12, 1985. Leaves on a lateral branch fixed (14)C photosynthetically and amounts of (14)C in seven biochemical fractions in each of six plant parts were determined 8, 24, and 72 h later. In all treatments, (14)C uptake was approximately 96% of that originally presented. Respiratory loss of (14)C ranged from 22 to 87% of uptake and increased sharply with increasing time after exposure and as the seedlings grew larger later in the study. Most (14)C was found in exposed leaves and very little occurred above the exposed branch. Amounts of (14)C decreased in the exposed leaves and increased in the roots with time after exposure and date. Sugars were generally the most heavily labeled fraction. Labeled sugar content of exposed leaves decreased by more than half between 8 and 72 h as sugars were metabolized and translocated to other parts, primarily the roots. In roots, the labeling of starch and residue (structural compounds) increased greatly with transport time and season. In all plant parts, proteins and amino acids contained very little (14)C regardless of date or time.     

Boron remobilization at low boron supply in olive (Olea europaea) in relation to leaf and phloem mannitol concentrations

For plant species in which a considerable portion of the photoassimilates are translocated in the phloem as sugar alcohols, boron is freely translocated from mature organs to growing tissues. However, the effects of decreased plant boron status on boron remobilization are poorly understood. We conducted a growth chamber experiment (CE) and a field experiment (FE) to study the effects of low boron supply on boron remobilization in olive (Olea europaea L.), a species that transports considerable amounts of mannitol in the phloem. For the CE, several physiological parameters were compared between control (B+) and boron-deficient olive plants (B-) during the expansion of new leaves. Boron remobilization was assessed by measuring boron content of selected leaves at the beginning and at the end of the CE. As expected, boron was remobilized from mature leaves to young leaves of B+ plants; however, considerable boron remobilization was also observed in B- plants, suggesting a mechanism whereby olive can sustain a minimum boron supply for growth of new tissues despite an insufficient external boron supply. Boron deficiency caused inhibition of new growth but had no effect on photosynthetic capacity per unit leaf surface area of young and mature leaves, thereby altering the carbon utilization pattern and resulting in carbon allocation to structures within the source leaves and accumulation of soluble carbohydrates. Specifically, in mature B- leaves in the CE and in B- leaves in the FE, mannitol concentration on a leaf water content basis increased by 48 and 27% respectively, compared with controls. Carbon export ability (assessed by both phloem anatomy and phloem exudate composition of FE leaves) was enhanced at low boron supply. We conclude that, at low boron supply, increased mannitol concentrations maintain boron remobilization from source leaves to boron-demanding sink leaves.     

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.     

Effects of elevated concentrations of ozone and carbon dioxide on the electrical impedance of leaves of silver birch (Betula pendula) clones

Effects of elevated concentrations of tropospheric ozone ([O3]) and carbon dioxide ([CO2]) on leaves of two silver birch (Betula pendula Roth) clones were monitored for three growing seasons (1998, 1999, 2000) by means of electrical impedance spectroscopy (EIS). The field trial with open-top chambers (OTCs) was conducted on two clones (Clone 4 and Clone 80) with five treatments and four independent replicates. Treatments were: (1) outside control, (2) chamber control, (3) 2x ambient [O3], (4) 2x ambient [CO2] and (5) 2x ambient [CO2] + 2x ambient [O3]. Fumigations started in 1999 and continued in 2000. Measurements were made in 1998 before the fumigations and thereafter EIS was carried out four times in each season. The impedance spectra of about 10 leaves from each tree at each time were measured at 42 frequencies between 80 and 1 MHz. Leaf spectra were modeled by a distributed circuit element model (DCE) (one DCE in series with a resistor), which yields the extracellular and intracellular resistances, the relaxation time and the distribution coefficient of the relaxation time. The EIS properties of the leaves changed significantly during the growing season when new leaves were expanding. The clones differed in their EIS properties. Clone 4 had a significantly higher extracellular resistance and distribution coefficient than Clone 80. The clones responded similarly to the fumigation treatments. Differences between treatments emerged especially during the second fumigation season in 2000. Elevated [O3] reduced both the relaxation time and the extracellular resistance, indicating cell membrane damage. Elevated [CO2] increased the intracellular resistance, indicating changes in symplastic composition. The biological interpretation of the EIS parameters in birch leaves is discussed.     

Silver birch and climate change: variable growth and carbon allocation responses to elevated concentrations of carbon dioxide and ozone

We studied the effects of elevated concentrations of carbon dioxide ([CO2]) and ozone ([O3]) on growth, biomass allocation and leaf area of field-grown O3-tolerant (Clone 4) and O3-sensitive clones (Clone 80) of European silver birch (Betula pendula Roth) trees during 1999-2001. Seven-year-old trees of Clones 4 and 80 growing outside in open-top chambers were exposed for 3 years to the following treatments: outside control (OC); chamber control (CC); 2 x ambient [CO2] (EC); 2 x ambient [O3] (EO); and 2 x ambient [CO2] + 2 x ambient [O3] (EC+EO). When the results for the two clones were analyzed together, elevated [CO2] increased tree growth and biomass, but had no effect on biomass allocation. Total leaf area increased and leaf abscission was delayed in response to elevated [CO2]. Elevated [O3] decreased dry mass of roots and branches and mean leaf size and induced earlier leaf abscission in the autumn; otherwise, the effects of elevated [O3] were small across the clones. However, there were significant interactions between elevated [CO2] and elevated [O3]. When results for the clones were analyzed separately, stem diameter, volume growth and total biomass of Clone 80 were increased by elevated [CO2] and the stimulatory effects of elevated [CO2] on stem volume growth and total leaf area increased during the 3-year study. Clone 80 was unaffected by elevated [O3]. In Clone 4, elevated [O3] decreased root and branch biomass by 38 and 29%, respectively, whereas this clone showed few responses to elevated [CO2]. Elevated [CO2] significantly increased total leaf area in Clone 80 only, which may partly explain the smaller growth responses to elevated [CO2] of Clone 4 compared with Clone 80. Although we observed responses to elevated [O3], the responses to the EC+EO and EC treatments were similar, indicating that the trees only responded to elevated [O3] under ambient [CO2] conditions, perhaps reflecting a greater quantity of carbohydrates available for detoxification and repair in elevated [CO2].     

Stomatal and leaf growth responses to partial drying of root tips in willow

Root tips of intact willow (Salix dasyclados Wimm., Clone 81-090) plants were partially dried by exposure to ambient greenhouse air and then kept in water-vapor-saturated air for up to 3 days. The drying treatment increased abscisic acid (ABA) concentrations in both the root tips subjected to drying and in the xylem sap, while it reduced leaf stomatal conductance and leaf extension rate. Despite the decrease in stomatal conductance, leaf water potentials were unaffected by the root drying treatment, indicating that the treatment reduced hydraulic conductivity between roots and foliage. After roots subjected to drying were returned to a nutrient solution or excised, ABA concentrations in the remaining roots and in the xylem sap, stomatal conductance of mature leaves and extension rate of unfolding leaves all returned to values observed in control plants. The 4-fold increase in xylem sap ABA concentration following the root drying treatment was not solely the result of reduced sap flow, and thus may be considered a potential cause, not merely a consequence, of the observed reduction in stomatal conductance.     

Growth and carbohydrate status of coppice shoots of hybrid poplar following shoot pruning

Fifteen, 1-year-old Populus maximowiczii Henry x P. nigra L. 'MN9' trees were decapitated and allowed to sprout. After 8 weeks, all had 6 to 10 coppice shoots. All shoots, except the tallest (dominant) shoot, were removed from five of the trees (pruned treatment), and shoot growth, gas exchange and carbohydrate status were compared in the pruned and unpruned trees. Although photosynthetic rate of recently mature leaves of pruned trees was approximately 50% greater than that of leaves on the dominant shoot of unpruned trees, and the dry weight of leaves of pruned trees was 37% greater than that of the leaves on the dominant shoot of unpruned trees, the shoot dry matter relative growth rate did not differ between treatments. Concentrations of water-soluble carbohydrates and starch in the uppper stem and leaves of the dominant shoot were similar in pruned and unpruned trees. However, relative to that of the dominant shoot in unpruned trees, the lower stem in pruned trees was depleted in both soluble carbohydrates and starch. Starch deposition, assessed as the quantity of (14)C-starch in tissues 24 h after a fully expanded source leaf was labeled with (14)CO(2), was 3.9 times greater in roots of pruned trees than in roots of unpruned trees. We conclude that early removal of all but the dominant shoot reduces the carbohydrate status of the roots and the lower portion of the stem by eliminating the excised shoots as a source of photosynthate.     

Quantification of two-year-old hybrid poplar root systems: morphology, biomass, and (14)C distribution

Root morphology, biomass, and (14)C distribution were studied in two 2-year-old Populus trichocarpa x P. deltoides hybrids, which originated from hardwood cuttings, to determine the pattern of root distribution in a plantation and to refine methods for root recovery. The trees were labeled with (14)CO(2) and harvested after a 72-hour chase period. Roots attached to each labeled tree were analyzed for morphological traits at the time of harvest. Detached roots from within a 1-m(3) volume of soil surrounding each tree were separated from the soil and sorted on the basis of rooting depth and root diameter. Lateral roots > 2 mm in diameter had a largely horizontal orientation at their point of origin from the cutting and extended horizontally up to 4 m from the cutting. This resulted in considerable overlap of root systems in the plantation. Results from (14)C labeling indicated that 24 +/- 4% (+/- SD) of the carbon exported from branches-labeled within two weeks after branch budset-was translocated to the root system. Dilution of the root (14)C label indicated that from 0 (> 5 mm diameter roots) to 75% (< 2 mm diameter roots) of the roots recovered from within the 1-m(3) volume of soil surrounding a harvested tree originated from other trees. Total root biomass was 6 +/- 1 Mg ha(-1) for both hybrids. Sixty percent of the root biomass was recovered directly from excavation, 16% from coarse-sieving excavated soil, and 24% from re-sorting sieved soil. The study indicated that root growth of hybrid poplars may be rapid and extensive and that detailed sorting of soil subsamples substantially improves the recovery of fine roots < 2 mm in diameter.     

Simulating the response of mature yellow poplar and loblolly pine trees to shifts in peak ozone periods during the growing season using the TREGRO model

Multiple TREGRO simulations were conducted with meteorological data files containing different growing season peak ozone (O(3)) episodes at O(3) exposures of 1.0 and 2.0 x ambient O(3) to assess the relationship between O(3) response and the phenology of mature yellow poplar (Liriodendron tulipifera L.) and loblolly pine (Pinus taeda L.) trees. Regardless of O(3) exposure and peak O(3) episode occurrence, a peak O(3) episode in August caused the greatest reduction in carbon (C) gain in yellow poplar, whereas a peak O(3) episode in July caused the greatest reduction in C gain of loblolly pine. In both species, timing of the greatest simulated O(3) effect corresponded with the completion of the annual foliage production phenophase. Simulated C gain of yellow poplar (total tree, coarse root, and total nonstructural carbohydrate) was reduced by O(3) to a greater extent than the corresponding compartments in loblolly pine, but the opposite was true for fine roots. This differential sensitivity to O(3) reflects the fact that both C assimilation and the O(3) response of the species were parameterized according to observed field measurements of each species. The differential sensitivity to O(3) of these species may have long-term implications for species composition in southeastern USA forests.     

马尾松幼苗光合产物的运输与分配特征

【目的】解析光合产物在马尾松幼苗植株中的运输和分配规律,揭示马尾松生产力形成过程,为探讨不同环境干扰下马尾松光合产物分配过程的变化提供参考依据。【方法】采用 13 C同位素脉冲标记法,对1.5年生马尾松幼苗进行标记,标记结束后第0、2、5、17、24、72、120、168、216和 360 h ,按不同部位对马尾松幼苗进行全收获取样,测定 13 C含量,以监测近期合成的光合产物在马尾松幼苗中的运输和分配规律,同时测定光合产物全碳、非结构性碳水化合物(NSC)在各个器官的积累量。【结果】1)标记的光合产物在针叶中合成后,向各库器官的运输量随着时间延长由多逐渐减少,具体表现为标记结束后0~24 h内最多,24~216 h逐渐减少,216 h之后运输完成,且59%以上标记的光合产物在0~24 h内运输到各库器官。2)光合产物运输趋于稳定后,在各器官的分配大小依次为1年生叶>当年生叶>根>茎干>1年生枝>当年生枝,与生物量的分配大小一致,但与库活力大小不同,其库活力依次为当年生叶>当年生枝>1年生叶>根>1年生枝>茎干。3)各器官全碳和NSC积累量的分配与近期合成光合产物的分配大小一致,依次为1年生叶>当年生叶>根>茎干>1年生枝>当年生枝。【结论】马尾松幼苗光合产物运输速率大于0.1 m ·h -1 ,59%以上标记的光合产物在合成后的0~ 24 h 内完成向各个库器官输出。新合成的光合产物在各器官中的积累量表现为功能器官(叶和根)居多,这一分配规律有利于马尾松幼苗阶段的生长。     

Carbon translocation patterns associated with new root proliferation during episodic growth of transplanted Quercus rubra seedlings

Patterns of carbon allocation in northern red oak (Quercus rubra L.), characterized by episodic growth through recurrent single-season flushing, vary by growth stage. To examine post-transplant timing and carbohydrate sources for new root growth, dormant, bare-root, half-sibling northern red oak seedlings were transplanted to pots and placed in a favorable growth chamber environment. Unlabeled seedlings were harvested at transplant and at the bud swell stage. After leaf emergence, seedlings were exposed to (14)CO(2) at the linear shoot, linear leaf or lag growth stages. Seedlings were then placed in a growth room for 48 h to allow for translocation of (14)C-labeled current photosynthate and its stabilization in sink component plant parts. Seedlings were subsequently harvested and tissue (14)C:(12)C ratio analyzed. New root growth began during the linear shoot growth stage. However, no increase in (14)C:(12)C ratio was found in new roots until the linear leaf and lag growth stages, indicating a downward shift in translocation of current photosynthate to fuel new root growth. In old roots, (14)C:(12)C ratio increased at the lag stage. Our results indicate that both stored carbohydrates and current photosynthate contribute to new root growth of transplanted northern red oak seedlings; stored carbohydrates promote initial new root proliferation, whereas current photosynthate assumes a greater role as new leaves mature and the flush terminates. Optimizing nursery practices to increase carbohydrate reserves may reduce the time required to establish root-soil contact and facilitate early post-planting survival.     

Metabolic responses to water deficit in two Eucalyptus globulus clones with contrasting drought sensitivity

We compared the metabolic responses of leaves and roots of two Eucalyptus globulus Labill. clones differing in drought sensitivity to a slowly imposed water deficit. Responses measured included changes in concentrations of soluble and insoluble sugars, proline, total protein and several antioxidant enzymes. In addition to the general decrease in growth caused by water deficit, we observed a decrease in osmotic potential when drought stress became severe. In both clones, the decrease was greater in roots than in leaves, consistent with the observed increases in concentrations of soluble sugars and proline in these organs. In roots of both clones, glutathione reductase activity increased significantly in response to water deficit, suggesting that this enzyme plays a protective role in roots during drought stress by catalyzing the catabolism of reactive oxygen species. Clone CN5 has stress avoidance mechanisms that account for its lower sensitivity to drought compared with Clone ST51.     

Microautoradiographic localization of phosphate and carbohydrates in mycorrhizal roots of Populus tremula x Populus alba and the implications for transfer processes in ectomycorrhizal associations

Microautoradiographic studies were carried out to examine the distribution and exchange of phosphate and labeled carbohydrates in mycorrhizal roots of Populus tremula x Populus alba L. following application of 33P-orthophosphate (Pi) and 14CO2. Labeled Pi was not homogeneously distributed along the mycorrhizal longitudinal axis. The fungal sheath and the Hartig net contained more 33Pi in the median parts of the root than in the apical or basal root zones, indicating that uptake and transfer of Pi to the host plant was localized mainly in this area. The Pi was translocated by the Hartig net and the interfacial apoplast to the host plant. It was distributed by way of the stele within the plant. Young leaves and meristematic tissue in the shoot tip were the main sinks for Pi. In plants that were left in the dark for 5 days before 33Pi application, the reduced carbohydrate supply caused a decrease in Pi absorption by mycorrhizal roots. Microautoradiography of mycorrhizal roots after assimilation of 14CO2 revealed that: (1) the fungal partner had a high capacity to attract photosynthates; (2) the main transfer of carbohydrates was localized in the median zone of a mycorrhizal root; (3) carbohydrates that were absorbed by the mycorrhizal fungus were translocated to the fungal sheath and were homogeneously distributed; and (4) in the main exchange zone, cortical cell nuclei showed a high sink capacity, indicating increased metabolic activity in these cells. We postulate that (1) the phosphate demand of the host plant regulates absorption of Pi by the fungus, and (2) a bidirectional transfer of carbohydrates and Pi occurs across the same interface structure in ectomycorrhizal roots of Populus.     

Abscisic acid in leaves and roots of willow: significance for stomatal conductance

Excised leaves and roots of willow (Salix dasyclados Wimm.) accumulated abscisic acid (ABA) in response to desiccation. The accumulation of ABA was greater in young leaves and roots than in old leaves and roots. In mature leaves, ABA accumulation was related to the severity and duration of the desiccation treatment. Water loss equal to 12% of initial fresh weight caused the ABA content of mature leaves to increase measurably within 30 min and to double in 2.5 h. The drying treatment caused significant (P = 0.05) reductions in leaf water potential and stomatal conductance. Recovery of leaf water potential to the control value occurred within 10 min of rewatering the dehydrated leaves, but recovery of stomatal conductance took an hour or longer, depending on the interval between dehydration and rewatering. The addition of ABA to the transpiration stream of well-watered excised leaves was sufficient to cause partial stomatal closure within 1 h and, depending on ABA concentration, more or less complete stomatal closure within 3 h. When the ABA solution was replaced with water, stomatal conductance increased at a rate inversely related to the concentration of the ABA solution with which the leaves had been supplied.     

Mild ozone exposure alters (14)C dynamics in foliage of Pinus taeda L

To explore the physiological mechanisms underlying ozone-induced growth reductions in loblolly pine (Pinus taeda L.), seedlings were exposed to sub-ambient (charcoal-filtered), ambient or twice-ambient ozone in open-top chambers for three growing seasons. In the final year of exposure, current-year needle fascicles were labeled with (14)CO(2) and the incorporation of (14)C into biochemical fractions was followed for 48 hours. Irrespective of ozone treatment, losses of (14)C-assimilates from foliage to respiration and translocation were minimal during the first 3 hours, whereas more than 60% of the label was lost during the next 45 hours. Radiolabel in sugar decreased rapidly after a lag period, roughly paralleling the pattern of total (14)C loss. The amount of (14)C label in starch and lipids plus pigments remained constant throughout the 48-hour chase period, whereas the amount of (14)C label in other fractions showed a net decrease over the 48-hour chase period. Ozone treatments altered foliar carbon dynamics in two ways: (1) ozone exposure increased foliar (14)C retention up to 21% for the first 5 hours after labeling, but not thereafter, and (2) ozone exposure decreased partitioning of (14)C into starch and increased partitioning of (14)C into organic acids, residue, and lipids plus pigments, indicating an intensified partitioning of carbon to injury and repair processes. Both short-term carbon retention and diversion of carbon from storage compounds to repair processes are foliar mechanisms by which ozone exposure could decrease growth in loblolly pine seedlings.     

Interactive effects of elevated ozone and temperature on carbon allocation of silver birch (Betula pendula) genotypes in an open-air field exposure

In the present experiment, the single and combined effects of elevated temperature and ozone (O(3)) on four silver birch genotypes (gt12, gt14, gt15 and gt25) were studied in an open-air field exposure design. Above- and below-ground biomass accumulation, stem growth and soil respiration were measured in 2008. In addition, a (13)C-labelling experiment was conducted with gt15 trees. After the second exposure season, elevated temperature increased silver birch above- and below-ground growth and soil respiration rates. However, some of these variables showed that the temperature effect was modified by tree genotype and prevailing O(3) level. For instance, in gt14 soil respiration was increased in elevated temperature alone (T) and in elevated O(3) and elevated temperature in combination (O(3) + T) treatments, but in other genotypes O(3) either partly (gt12) or totally nullified (gt25) temperature effects on soil respiration, or acted synergistically with temperature (gt15). Before leaf abscission, all genotypes had the largest leaf biomass in T and O(3) + T treatments, whereas at the end of the season temperature effects on leaf biomass depended on the prevailing O(3) level. Temperature increase thus delayed and O(3) accelerated leaf senescence, and in combination treatment O(3) reduced the temperature effect. Photosynthetic : non-photosynthetic tissue ratios (P : nP ratios) showed that elevated temperature increased foliage biomass relative to woody mass, particularly in gt14 and gt12, whereas O(3) and O(3) + T decreased it most clearly in gt25. O(3)-caused stem growth reductions were clearest in the fastest-growing gt14 and gt25, whereas mycorrhizal root growth and sporocarp production increased under O(3) in all genotypes. A labelling experiment showed that temperature increased tree total biomass and hence (13)C fixation in the foliage and roots and also label return was highest under elevated temperature. Ozone seemed to change tree (13)C allocation, as it decreased foliar (13)C excess amount, simultaneously increasing (13)C excess obtained from the soil. The present results suggest that warming has potential to increase silver birch growth and hence carbon (C) accumulation in tree biomass, but the final magnitude of this C sink strength is partly counteracted by temperature-induced increase in soil respiration rates and simultaneous O(3) stress. Silver birch populations' response to climate change will also largely depend on their genotype composition.     

Nitrogen absorption,translocation and distribution from urea applied in autumn to leaves of young potted apple (Malus domestica) trees

We studied the absorption, assimilation, translocation and distribution of nitrogen (N) from urea applied in autumn to leaves of 1-year-old potted Fuji/M26 apple (Malus domestica Borkh) trees. In early October, all leaves of each tree were painted with either 3% urea (enriched to 10 atom % with 15N) or water (control trees). Four trees were harvested before the treatment and N and amino acid contents were determined. Four trees from each treatment were harvested at 2, 4, 7, 10, 15 and 20 days after urea or water application. Total N, amino acids and 15N in leaves, bark, xylem, shank and roots were analyzed to determine uptake and mobilization of N from urea. Most uptake of 15N by leaves occurred during the first 2 days following application of urea. The mean rate of absorption during these 2 days was 0.29 g m-2 day-1. Amino acids in leaves, bark and roots increased significantly after urea application compared with control values. The highest concentrations of amino acids in leaves and bark occurred 4 days after application, whereas the highest concentrations of amino acids in roots occurred 10 days after application. Total 15N content in leaves peaked 2 days after urea application and then decreased, whereas 15N content in roots and bark increased throughout the experiment. Total 15N content in xylem and shank was low. Leaves absorbed 35% of the 15N applied as urea, and 63.6% of absorbed 15N was translocated out of leaves within 20 days after urea application. We conclude that N from urea was converted to amino acids in leaves after foliar application in autumn, and roots and bark were the main sinks of N from urea applied to leaves.