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  • 1
    ISSN: 1365-3040
    Source: Blackwell Publishing Journal Backfiles 1879-2005
    Topics: Biology
    Notes: Stomatal density, anatomy and nutrient concentrations of Scots pine (Pinus sylvestris L.) needles were studied during 3 years of growth at elevated CO2 (693 ± 30 µmol mol−1), at elevated temperature (ambient +2·8–6·2 °C depending on the time of the year) and in a combination of elevated CO2 and temperature in closed-top chambers. The treatments were started in August 1996. At elevated temperature, the needles that were grown in the first year (i.e. the 1997 cohort) were thinner, had thinner mesophyll in the abaxial side, thinner vascular cylinder and lower stomatal density than those grown at ambient temperature. The proportion of mesophyll area occupied by vascular cylinder or intercellular spaces were not changed. Lower stomatal density apparently did not lead to decreased use of water, as these needles had higher concentrations of less mobile nutrients (Ca, Mg, B, Zn and Mn), which could indicate increased total transpiration. In the 1997 and 1998 cohorts, elevation of temperature decreased concentrations of N, P, K, S and Cu. In the 1999 cohort, contradictory, higher concentrations of N and S at elevated temperature may be related to increased nutrient mineralization in the soil. Elevation of CO2 did not affect stomatal density, needle thickness, thickness of epidermis or hypodermis, vascular cylinder or intercellular spaces. Concentrations of N, P, S and Cu decreased at elevated CO2. Reductions were transient and most distinct in the 1997 cohort. The effects of CO2 and temperature were in some cases interactive, which meant that in the combined treatment stomatal density decreased less than at elevated temperature, and concentrations of nutrients decreased less than expected on the basis of separate treatments, whereas the thickness of the epidermis and hypodermis decreased more than in the separate treatments. In conclusion, alterations in the anatomy and stomatal density of Scots pine needles were more distinct at elevated temperature than at elevated CO2. Both elevated CO2 and temperature-induced changes in nutrient concentrations that partly corresponded to the biochemical and photosynthetic alterations in the same cohorts (Luomala et al. Plant, Cell and Environment26, 645–660, 2003) Reductions in nutrient concentrations and alterations in the anatomy were transient and more evident in the needle cohort that was grown in the first treatment year.
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  • 2
    Electronic Resource
    Electronic Resource
    Oxford, UK : Blackwell Science Ltd
    Global change biology 11 (2005), S. 0 
    ISSN: 1365-2486
    Source: Blackwell Publishing Journal Backfiles 1879-2005
    Topics: Biology , Energy, Environment Protection, Nuclear Power Engineering , Geography
    Notes: Atmospheric change may affect plant phenolic compounds, which play an important part in plant survival. Therefore, we studied the impacts of CO2 and O3 on the accumulation of 27 phenolic compounds in the short-shoot leaves of two European silver birch (Betula pendula Roth) clones (clones 4 and 80). Seven-year-old soil-grown trees were exposed in open-top chambers over three growing seasons to ambient and twice ambient CO2 and O3 concentrations singly and in combination in central Finland.Elevated CO2 increased the concentration of the phenolic acids (+25%), myricetin glycosides (+18%), catechin derivatives (+13%) and soluble condensed tannins (+19%) by increasing their accumulation in the leaves of the silver birch trees, but decreased the flavone aglycons (−7%) by growth dilution. Elevated O3 increased the concentration of 3,4′-dihydroxypropiophenone 3-β-d-glucoside (+22%), chlorogenic acid (+19%) and flavone aglycons (+4%) by inducing their accumulation possibly as a response to increased oxidative stress in the leaf cells. Nevertheless, this induction of antioxidant phenolic compounds did not seem to protect the birch leaves from detrimental O3 effects on leaf weight and area, but may have even exacerbated them. On the other hand, elevated CO2 did seem to protect the leaves from elevated O3 because all the O3-derived effects on the leaf phenolics and traits were prevented by elevated CO2. The effects of the chamber and elevated CO2 on some compounds changed over time in response to the changes in the leaf traits, which implies that the trees were acclimatizing to the altered environmental conditions. Although the two clones used possessed different composition and concentrations of phenolic compounds, which could be related to their different latitudinal origin and physiological characteristics, they responded similarly to the treatments. However, in some cases the variation in phenolic concentrations caused by genotype or chamber environment was much larger than the changes caused by either elevated CO2 or O3.
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  • 3
    ISSN: 1365-2486
    Source: Blackwell Publishing Journal Backfiles 1879-2005
    Topics: Biology , Energy, Environment Protection, Nuclear Power Engineering , Geography
    Notes: Impacts of ozone and CO2 enrichment, alone and in combination, on leaf anatomical and ultrastructural characteristics, nutrient status and cell wall chemistry in two European silver birch (Betula pendula Roth) clones were studied. The young soil-growing trees were exposed in open-top chambers over three growing seasons to 2 × ambient CO2 and/or ozone concentrations in central Finland. The trees were measured for changes in altogether 35 variables of leaf structure, nutrients and cell wall chemistry of green leaves, and 20 of the measured variables were affected by CO2 and/or O3. Elevated CO2 increased the size of chloroplasts and starch grains, number of mitochondria, P : N ratio, and contents of cell wall hemicellulose. Elevated CO2 decreased the total leaf thickness, specific leaf area, concentrations of N, K, Cu, S and Fe, and contents of cell wall α-cellulose, uronic acids, acid-soluble lignin and acetone-soluble extractives. Elevated ozone led to thinner leaves, higher palisade to spongy ratio, increased number of peroxisomes and mitochondria, reduced content of Mn, Zn, Cu, hemicellulose and uronic acids, and lower Mn : N and Zn : N ratios. In the combined exposure, interactions were antagonistic. Ultrastructural changes became more evident towards the end of the exposure. Young leaves were tolerant against ozone-caused oxidative stress, whereas oxidative H2O2 accumulation was found in older leaves. CO2 enrichment improved ozone tolerance not only through increased photosynthesis rates, but also through changes in cell wall chemistry (hemicellulose, in particular). However, nutrient imbalances due to ozone and/or CO2 may predispose the trees to other biotic and abiotic stresses. Down-regulation and up-regulation of photosynthesis under elevated CO2 through anatomical changes is discussed.
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  • 4
    ISSN: 1365-2486
    Source: Blackwell Publishing Journal Backfiles 1879-2005
    Topics: Biology , Energy, Environment Protection, Nuclear Power Engineering , Geography
    Notes: In the present open-top chamber experiment, two silver birch clones (Betula pendula Roth, clone 4 and clone 80) were exposed to elevated levels of carbon dioxide (CO2) and ozone (O3), singly and in combination, and soil CO2 efflux was measured 14 times during three consecutive growing seasons (1999–2001). In the beginning of the experiment, all experimental trees were 7 years old and during the experiment the trees were growing in sandy field soil and fertilized regularly. In general, elevated O3 caused soil CO2 efflux stimulation during most measurement days and this stimulation enhanced towards the end of the experiment. The overall soil respiration response to CO2 was dependent on the genotype, as the soil CO2 efflux below clone 80 trees was enhanced and below clone 4 trees was decreased under elevated CO2 treatments. Like the O3 impact, this clonal difference in soil respiration response to CO2 increased as the experiment progressed. Although the O3 impact did not differ significantly between clones, a significant time × clone × CO2× O3 interaction revealed that the O3-induced stimulation of soil respiration was counteracted by elevated CO2 in clone 4 on most measurement days, whereas in clone 80, the effect of elevated CO2 and O3 in combination was almost constantly additive during the 3-year experiment. Altogether, the root or above-ground biomass results were only partly parallel with the observed soil CO2 efflux responses. In conclusion, our data show that O3 impacts may appear first in the below-ground processes and that relatively long-term O3 exposure had a cumulative effect on soil CO2 efflux. Although the soil respiration response to elevated CO2 depended on the tree genotype as a result of which the O3 stress response might vary considerably within a single tree species under elevated CO2, the present experiment nonetheless indicates that O3 stress is a significant factor affecting the carbon cycling in northern forest ecosystems.
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  • 5
    ISSN: 1365-2486
    Source: Blackwell Publishing Journal Backfiles 1879-2005
    Topics: Biology , Energy, Environment Protection, Nuclear Power Engineering , Geography
    Notes: Field-growing silver birch (Betula pendula Roth) clones (clone 4 and 80) were exposed to elevated CO2 and O3 in open-top chambers for three consecutive growing seasons (1999–2001). At the beginning of the OTC experiment, all trees were 7 years old. We studied the single and interaction effects of CO2 and O3 on silver birch below-ground carbon pools (i.e. effects on fine roots and mycorrhizas, soil microbial communities and sporocarp production) and also assessed whether there are any clonal differences in these below-ground CO2 and O3 responses. The total mycorrhizal infection level of both clones was stimulated by elevated CO2 alone and elevated O3 alone, but not when elevated CO2 was used in fumigation in combination with elevated O3. In both clones, elevated CO2 affected negatively light brown/orange mycorrhizas, while its effect on other mycorrhizal morphotypes was negligible. Elevated O3, instead, clearly decreased the proportions of black and liver-brown mycorrhizas and increased that of light brown/orange mycorrhizas. Elevated O3 had a tendency to decrease standing fine root mass and sporocarp production as well, both of these O3 effects mainly manifesting in clone 4 trees. CO2 and O3 treatment effects on soil microbial community composition (PLFA, 2- and 3-OH-FA profiles) were negligible, but quantitative PLFA data showed that in 2001 the PLFA fungi : bacteria-ratio of clone 80 trees was marginally increased because of elevated CO2 treatments. This study shows that O3 effects were most clearly visible at the mycorrhizal root level and that some clonal differences in CO2 and O3 responses were observable in the below-ground carbon pools. In conclusion, the present data suggests that CO2 effects were minor, whereas increasing tropospheric O3 levels can be an important stress factor in northern birch forests, as they might alter mycorrhizal morphotype assemblages, mycorrhizal infection rates and sporocarp production.
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  • 6
    Electronic Resource
    Electronic Resource
    Copenhagen : Munksgaard International Publishers
    Physiologia plantarum 113 (2001), S. 0 
    ISSN: 1399-3054
    Source: Blackwell Publishing Journal Backfiles 1879-2005
    Topics: Biology
    Notes: Infiltration of reduced ascorbate (ASC) into the leaves of Betula pendula Roth and subsequent measurement of its loss therein after incubation allowed us to follow ascorbate transport from apoplast to symplast in intact leaves. All of the ascorbate extracted from the native apoplast was in fully oxidized form, dehydroascorbate (DHA). When 5 mM of ASC was infiltrated into the leaves, its intense decay occurred, but only 55% of ASC lost was recovered in apoplast as DHA. When ASC was added to the freshly extracted intercellular washing fluid (IWF), ASC oxidation occurred as well. However, all oxidized ASC was recovered as DHA, indicating that further decomposition of DHA did not occur. Similarly, all of the ASC infiltrated into the leaves was found therein either as ASC or DHA after incubation of leaves for up to 60 min. On this base the ascorbate infiltrated into the leaves and not recovered in the IWF was interpreted as ascorbate taken up into the symplast. The calculated uptake rates of ascorbate at different ASC concentrations followed saturation kinetics with the maximum uptake rate of 300 nmol m−2 plasma membrane (PM) area min−1 and Michaelis constant of 12.8 mM. The uptake of ascorbate was significantly inhibited by the addition of dithiothreitol or by PM H+ ATPase inhibitor erythrosin B. Thus, our results support the previous observations that DHA is preferably transported from the apoplastic to the cytoplasmic side of the membrane and show that this process is dependent upon PM proton gradient.
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  • 7
    ISSN: 1399-3054
    Source: Blackwell Publishing Journal Backfiles 1879-2005
    Topics: Biology
    Notes: Leaf development and senescence were studied in greenhouse-grown silver birch (Betula pendula Roth) seedlings over a period of 7 weeks. Prior to the experiment, leaves from 100 seedlings were marked for five sampling dates. Timing of the developmental phases was studied with biochemical analyses of total soluble protein, Rubisco protein, chlorophyll concentration and at the level of gene expression related to photosynthesis, energy metabolism, ethylene synthesis and protein degradation. During the sampling period, an initial increase in photosynthetic capacity could be seen, when expression of the Rubisco small subunit gene (RbcS) and Rubisco protein (EC 4.1.1.39) were examined. Down-regulation of photosynthesis, visible as a decrease in Rubisco protein and RbcS mRNA, started soon after full expansion of the leaves and processes related to senescence followed. mRNA accumulation for the ethylene-forming enzyme 1-aminocyclopropane-1-carboxylic acid oxidase increased first during the onset of senescence. Protein degradation was observed as a loss of soluble proteins, with a simultaneous increase in the leucine aminopeptidase (EC 3.4.11.1) mRNA levels. The mRNA levels of ribonuclease-like pathogenesis-related protein 10 also increased clearly during senescence, but the mitochondrial phosphate translocator mRNA showed only a slight increase. Chlorophyll concentration of the leaves decreased after the mRNA levels of these senescence-related genes had become more abundant.
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  • 8
    ISSN: 1365-2486
    Source: Blackwell Publishing Journal Backfiles 1879-2005
    Topics: Biology , Energy, Environment Protection, Nuclear Power Engineering , Geography
    Notes: The aim of this study was to examine the effects of elevated carbon dioxide [CO2] and ozone [O3] and their interaction on wood chemistry and anatomy of five clones of 3-year-old trembling aspen (Populus tremuloides Michx.). Wood chemistry was studied also on paper birch (Betula papyrifera Marsh.) and sugar maple (Acer saccharum Marsh.) seedling-origin saplings of the same age. Material for the study was collected from the Aspen Free-Air CO2 Enrichment (FACE) experiment in Rhinelander, WI, USA, where the saplings had been exposed to four treatments: control (C; ambient CO2, ambient O3), elevated CO2 (560 ppm during daylight hours), elevated O3 (1.5 × ambient during daylight hours) and their combination (CO2+O3) for three growing seasons (1998–2000). Wood chemistry responses to the elevated CO2 and O3 treatments differed between species. Aspen was most responsive, while maple was the least responsive of the three tree species. Aspen genotype affected the responses of wood chemistry and, to some extent, wood structure to the treatments. The lignin concentration increased under elevated O3 in four clones of aspen and in birch. However, elevated CO2 ameliorated the effect. In two aspen clones, nitrogen in wood samples decreased under combined exposure to CO2 and O3. Soluble sugar concentration in one aspen clone and starch concentration in two clones were increased by elevated CO2. In aspen wood, α-cellulose concentration changed under elevated CO2, decreasing under ambient O3 and slightly increasing under elevated O3. Hemicellulose concentration in birch was decreased by elevated CO2 and increased by elevated O3. In aspen, elevated O3 induced statistically significant reductions in distance from the pith to the bark and vessel lumen diameter, as well as increased wall thickness and wall percentage, and in one clone, decreased fibre lumen diameter. Our results show that juvenile wood properties of broadleaves, depending on species and genotype, were altered by atmospheric gas concentrations predicted for the year 2050 and that CO2 ameliorates some adverse effects of elevated O3 on wood chemistry.
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  • 9
    ISSN: 1365-2486
    Source: Blackwell Publishing Journal Backfiles 1879-2005
    Topics: Biology , Energy, Environment Protection, Nuclear Power Engineering , Geography
    Notes: The objective of the present study was to investigate the interactive effects of elevated [CO2] and soil nutrient availability on secondary xylem structure and chemical composition of 41-year-old Norway spruce (Picea abies (L.) Karst.) trees. The nonfertilized and irrigated-fertilized trees were, for 3 years, continuously exposed to elevated [CO2] in whole-tree chambers. Elevated [CO2] decreased concentrations of soluble sugars, acid-soluble lignin and nitrogen in stem wood, but the effects were not consistent between sampling height and/or fertilization. The effect of 2*ambient [CO2] on wood structure depended on the exposure year and/or fertilization. Radial lumen diameter decreased and annual ring width increased in the second year of exposure (1999) in elevated [CO2]. In the latter, the CO2 effect was significant only in the nonfertilized trees. Stem wood chemistry and structure were significantly affected by fertilization. Fertilization increased the concentrations of nitrogen and gravimetric lignin, annual ring width, and radial lumen diameter. Fertilization decreased C/N ratio, mean ring density, earlywood density, latewood density, cell wall thickness, cell wall index, and latewood percentage. We conclude that elevated [CO2] had only minor effects on wood properties while fertilization had more marked effects and thus may affect ecosystem processes and suitability of wood for different end-use purposes.
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  • 10
    Electronic Resource
    Electronic Resource
    Oxford, UK : Blackwell Publishing Ltd
    Physiologia plantarum 77 (1989), S. 0 
    ISSN: 1399-3054
    Source: Blackwell Publishing Journal Backfiles 1879-2005
    Topics: Biology
    Notes: Uptake of dissolved inorganic carbon (DIC) from a nutrient solution by willow roots was measured in light and darkness and the distribution in the plant of DIC taken up by the roots was determined. It was also studied whether the transport system could be activated by preincubation with dissolved inorganic carbon.Willow plants (Salix cv. Aquatica gigantea) grown in hydroponic culture media were preincubated for 2 days with or without 0.74 mM NaHCO3. After preincubation, either unlabelled or [14C]-labelled NaHCO3 was injected into the media and after 1, 5, 10 and 24 h either in light or in darkness the plants were harvested in pieces into liquid nitrogen, lyophilized and burned in a combustion chamber. 14C was transported through the roots to the shoots and leaves both in light and in darkness, although incorporation of 14C in darkness was only half of that in light at the end of the 24-h feeding period. Both in light and in darkness the amount of 14C increased in all parts of willow plants with time. In light the rate of labelling was highest into cuttings and shoots. In darkness more than half of the total label was detected in cuttings of both the non-activated and the activated treatments.In the shoots the middle part was most strongly labelled after 5 and 10 h, but after 24 h 14C moved towards the base of the shoot. In the leaves at all feeding times most radioactivity was incorporated into the young, fully open leaves on the upper part of the shoots. Preincubation of plants with unlabelled NaHCO3 in growth media had no clear effect on the rate of DIC uptake either in light or in darkness.
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