Publications by authors named "Elina Oksanen"

46 Publications

Ozone affects plant, insect, and soil microbial communities: A threat to terrestrial ecosystems and biodiversity.

Sci Adv 2020 Aug 12;6(33):eabc1176. Epub 2020 Aug 12.

Institute of Research on Terrestrial Ecosystems (IRET), National Research Council of Italy (CNR), Via Madonna del Piano 10, I-50019 Sesto Fiorentino, Italy.

Elevated tropospheric ozone concentrations induce adverse effects in plants. We reviewed how ozone affects (i) the composition and diversity of plant communities by affecting key physiological traits; (ii) foliar chemistry and the emission of volatiles, thereby affecting plant-plant competition, plant-insect interactions, and the composition of insect communities; and (iii) plant-soil-microbe interactions and the composition of soil communities by disrupting plant litterfall and altering root exudation, soil enzymatic activities, decomposition, and nutrient cycling. The community composition of soil microbes is consequently changed, and alpha diversity is often reduced. The effects depend on the environment and vary across space and time. We suggest that Atlantic islands in the Northern Hemisphere, the Mediterranean Basin, equatorial Africa, Ethiopia, the Indian coastline, the Himalayan region, southern Asia, and Japan have high endemic richness at high ozone risk by 2100.
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http://dx.doi.org/10.1126/sciadv.abc1176DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7423369PMC
August 2020

RPA-PCR couple: an approach to expedite plant diagnostics and overcome PCR inhibitors.

Biotechniques 2020 10 20;69(4):270-280. Epub 2020 Aug 20.

Department of Environmental & Biological Sciences, University of Eastern Finland, Kuopio 70211, Finland.

DNA extraction can be lengthy and sometimes ends up with amplification inhibitors. We present the potential of recombinase polymerase amplification (RPA) to replace plant DNA extraction. In our rapid 'RPA-PCR couple' concept, RPA is tuned to slower reaction kinetics to promote amplification of long targets. RPA primers amplify target and some flanking regions directly from simple plant macerates. Then PCR primers exponentially amplify the target directly from the RPA reaction. We present the coupling of RPA with conventional, TaqMan and SYBR Green PCR assays. We applied the concept to strawberry pathogens and the identification marker . We found RPA-PCR couple specific, sensitive and reliable. The approach may also benefit other difficult samples such as food, feces and ancient samples.
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http://dx.doi.org/10.2144/btn-2020-0065DOI Listing
October 2020

Insect herbivory dampens Subarctic birch forest C sink response to warming.

Nat Commun 2020 05 21;11(1):2529. Epub 2020 May 21.

Faculty of Biological and Environmental Sciences, University of Helsinki, Niemenkatu 73, 15140, Lahti, Finland.

Climate warming is anticipated to make high latitude ecosystems stronger C sinks through increasing plant production. This effect might, however, be dampened by insect herbivores whose damage to plants at their background, non-outbreak densities may more than double under climate warming. Here, using an open-air warming experiment among Subarctic birch forest field layer vegetation, supplemented with birch plantlets, we show that a 2.3 °C air and 1.2 °C soil temperature increase can advance the growing season by 1-4 days, enhance soil N availability, leaf chlorophyll concentrations and plant growth up to 400%, 160% and 50% respectively, and lead up to 122% greater ecosystem CO uptake potential. However, comparable positive effects are also found when insect herbivory is reduced, and the effect of warming on C sink potential is intensified under reduced herbivory. Our results confirm the expected warming-induced increase in high latitude plant growth and CO uptake, but also reveal that herbivorous insects may significantly dampen the strengthening of the CO sink under climate warming.
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http://dx.doi.org/10.1038/s41467-020-16404-4DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7242322PMC
May 2020

Differences in growth and gas exchange between southern and northern provenances of silver birch (Betula pendula Roth) in northern Europe.

Tree Physiol 2020 02;40(2):198-214

Department of Environmental and Biological Sciences, University of Eastern Finland, Yliopistokatu 7, PO Box 111, 80101 Joensuu, Finland.

Due to its ubiquity across northern latitudes, silver birch (Betula pendula Roth) is an attractive model species for studying geographical trait variation and acclimation capacity. Six birch provenances from 60 to 67°N across Finland were grown in a common garden and studied for provenance and genotype variation. We looked for differences in height growth, photosynthetic gas exchange and chlorophyll content index (CCI) and compared the gas exchange of early and late leaves on short and long shoots, respectively. The provenances stratified into southern and northern groups. Northern provenances attained less height growth increment and had higher stomatal conductance (gs) and lower intrinsic water-use efficiency (WUE, Anet/gs) than southern provenances, whereas net photosynthesis (Anet) or CCI did not show clear grouping. Short shoot leaves had lower gs and higher WUE than long shoot leaves in all provenances, but there was no difference in Anet between shoot types. The separation of the provenances into two groups according to their physiological responses might reflect the evolutionary history of B. pendula. Latitudinal differences in gas exchange and water use traits can have plausible consequences for global carbon and water fluxes in a warming climate.
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http://dx.doi.org/10.1093/treephys/tpz124DOI Listing
February 2020

Elevated temperature and ozone modify structural characteristics of silver birch (Betula pendula) leaves.

Tree Physiol 2020 04;40(4):467-483

Department of Environmental and Biological Sciences, University of Eastern Finland, PO Box 1627, Yliopistonranta 1, Kuopio FI-70211, Finland.

To study the effects of slightly elevated temperature and ozone (O3) on leaf structural characteristics of silver birch (Betula pendula Roth), saplings of four clonal genotypes of this species were exposed to elevated temperature (ambient air temperature +0.8-1.0 °C) and elevated O3 (1.3-1.4× ambient O3), alone and in combination, in an open-air exposure field over two growing seasons (2007 and 2008). So far, the impacts of moderate elevation of temperature or the combination of elevated temperature and O3 on leaf structure of silver birch have not been intensively studied, thus showing the urgent need for this type of studies. Elevated temperature significantly increased leaf size, reduced non-glandular trichome density, decreased epidermis thickness and increased plastoglobuli size in birch leaves during one or both growing seasons. During the second growing season, O3 elevation reduced leaf size, increased palisade layer thickness and decreased the number of plastoglobuli in spongy cells. Certain leaf structural changes observed under a single treatment of elevated temperature or O3, such as increase in the amount of chloroplasts or vacuole, were no longer detected at the combined treatment. Leaf structural responses to O3 and rising temperature may also depend on timing of the exposure during the plant and leaf development as indicated by the distinct changes in leaf structure along the experiment. Genotype-dependent cellular responses to the treatments were detected particularly in the palisade cells. Overall, this study showed that even a slight but realistic elevation in ambient temperature can notably modify leaf structure of silver birch saplings. Leaf structure, in turn, influences leaf function, thus potentially affecting acclimation capacity under changing climate.
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http://dx.doi.org/10.1093/treephys/tpz127DOI Listing
April 2020

Trichomes form an important first line of defence against adverse environment-New evidence for ozone stress mitigation.

Authors:
Elina Oksanen

Plant Cell Environ 2018 07 20;41(7):1497-1499. Epub 2018 Apr 20.

Department of Environmental and Biological Sciences, University of Eastern Finland, PO Box 111, 80101, Joensuu, Finland.

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http://dx.doi.org/10.1111/pce.13187DOI Listing
July 2018

Genome sequencing and population genomic analyses provide insights into the adaptive landscape of silver birch.

Nat Genet 2017 Jun 8;49(6):904-912. Epub 2017 May 8.

Umeå Plant Science Centre, Department of Plant Physiology, Umeå University, Umeå, Sweden.

Silver birch (Betula pendula) is a pioneer boreal tree that can be induced to flower within 1 year. Its rapid life cycle, small (440-Mb) genome, and advanced germplasm resources make birch an attractive model for forest biotechnology. We assembled and chromosomally anchored the nuclear genome of an inbred B. pendula individual. Gene duplicates from the paleohexaploid event were enriched for transcriptional regulation, whereas tandem duplicates were overrepresented by environmental responses. Population resequencing of 80 individuals showed effective population size crashes at major points of climatic upheaval. Selective sweeps were enriched among polyploid duplicates encoding key developmental and physiological triggering functions, suggesting that local adaptation has tuned the timing of and cross-talk between fundamental plant processes. Variation around the tightly-linked light response genes PHYC and FRS10 correlated with latitude and longitude and temperature, and with precipitation for PHYC. Similar associations characterized the growth-promoting cytokinin response regulator ARR1, and the wood development genes KAK and MED5A.
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http://dx.doi.org/10.1038/ng.3862DOI Listing
June 2017

Low vapor pressure deficit reduces glandular trichome density and modifies the chemical composition of cuticular waxes in silver birch leaves.

Tree Physiol 2017 09;37(9):1166-1181

Department of Environmental and Biological Sciences, University of Eastern Finland, PO Box 111, 80101 Joensuu, Finland.

Cuticular wax layer is the first barrier against the outside environment and the first defense encountered by herbivores and pathogens. The effects of environmental factors on cuticular chemistry, and on the formation of glandular trichomes that account for the storage and secretion of lipophilic compounds to the leaf surface are poorly understood. Low vapor pressure deficit (VPD) has shown to reduce the nitrogen (N) status of plants. Thus, we studied the effects of elevated air humidity, indicated as VPD, and the effect of N fertilization on cuticular waxes and glandular trichome density in silver birch (Betula pendula Roth). Experiments were carried out in growth chambers with juvenile plants and in a long-term field experiment with older trees. Low VPD reduced the glandular trichome density in both experiments, in chamber and in field. The contents of the major triterpenoid and flavonoid aglycones correlated positively with glandular trichome density, which supports the role of trichomes in the exudation of secondary compounds to the leaf surface. A closer examination of the cuticular wax chemistry in the chamber experiment revealed that low VPD and N supply affected the composition of cuticular waxes, but not the total wax content. The deposition of different wax compounds followed a co-ordinated pattern in birch leaves, but different compound groups varied in their responses to N fertilization and low VPD. Low VPD reduced the hydrophobicity of cuticular waxes, as demonstrated by lower alkane content and less hydrophobic flavonoid profile in low VPD than in high VPD. Reduced hydrophobicity of the wax layer is presumed to increase leaf wettability. Together with reduced trichome density in low VPD it may enhance the susceptibility of trees to fungal pathogens and herbivores. High N supply under low VPD reduced the effect of low VPD on the cuticular wax composition. Total fatty acid content and the expression of β-amyrin synthase were lower under high N supply than under moderate N supply irrespective of VPD treatment. Nitrogen availability and decreasing VPD will modify leaf surface properties in silver birch and thereby affect tree defence against abiotic and biotic stress factors that emerge under climate change.
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http://dx.doi.org/10.1093/treephys/tpx045DOI Listing
September 2017

Low vapour pressure deficit affects nitrogen nutrition and foliar metabolites in silver birch.

J Exp Bot 2016 07 3;67(14):4353-65. Epub 2016 Jun 3.

University of Eastern Finland, Department of Environmental and Biological Sciences, PO Box 111, 80101 Joensuu, Finland.

Air humidity indicated as vapour pressure deficit (VPD) is directly related to transpiration and stomatal function of plants. We studied the effects of VPD and nitrogen (N) supply on leaf metabolites, plant growth, and mineral nutrition with young micropropagated silver birches (Betula pendula Roth.) in a growth chamber experiment. Plants that were grown under low VPD for 26 d had higher biomass, larger stem diameter, more leaves, fewer fallen leaves, and larger total leaf area than plants that were grown under high VPD. Initially, low VPD increased height growth rate and stomatal conductance; however, the effect was transient and the differences between low and high VPD plants became smaller with time. Metabolic adjustment to low VPD reflected N deficiency. The concentrations of N, iron, chlorophyll, amino acids, and soluble carbohydrates were lower and the levels of starch, quercetin glycosides, and raffinose were higher in the leaves that had developed under low VPD compared with high VPD. Additional N supply did not fully overcome the negative effect of low VPD on nutrient status but it diminished the effects of low VPD on leaf metabolism. Thus, with high N supply, the glutamine to glutamate ratio and starch production under low VPD became comparable with the levels under high VPD. The present study demonstrates that low VPD affects carbon and nutrient homeostasis and modifies N allocation of plants.
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http://dx.doi.org/10.1093/jxb/erw218DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5301935PMC
July 2016

Artificially decreased vapour pressure deficit in field conditions modifies foliar metabolite profiles in birch and aspen.

J Exp Bot 2016 07 2;67(14):4367-78. Epub 2016 Jun 2.

University of Eastern Finland, Department of Environmental and Biological Sciences, PO Box 111, 80101 Joensuu, Finland.

Relative air humidity (RH) is expected to increase in northern Europe due to climate change. Increasing RH reduces the difference of water vapour pressure deficit (VPD) between the leaf and the atmosphere, and affects the gas exchange of plants. Little is known about the effects of decreased VPD on plant metabolism, especially under field conditions. This study was conducted to determine the effects of artificially decreased VPD on silver birch (Betula pendula Roth.) and hybrid aspen (Populus tremula L.×P. tremuloides Michx.) foliar metabolite and nutrient profiles in a unique free air humidity manipulation (FAHM) field experiment during the fourth season of humidity manipulation, in 2011. Long-term exposure to decreased VPD modified nutrient homeostasis in tree leaves, as demonstrated by a lower N concentration and N:P ratio in aspen leaves, and higher Na concentration and lower K:Na ratio in the leaves of both species in decreased VPD than in ambient VPD. Decreased VPD caused a shift in foliar metabolite profiles of both species, affecting primary and secondary metabolites. Metabolic adjustment to decreased VPD included elevated levels of starch and heptulose sugars, sorbitol, hemiterpenoid and phenolic glycosides, and α-tocopherol. High levels of carbon reserves, phenolic compounds, and antioxidants under decreased VPD may modify plant resistance to environmental stresses emerging under changing climate.
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http://dx.doi.org/10.1093/jxb/erw219DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5301936PMC
July 2016

Volatile organic compounds emitted from silver birch of different provenances across a latitudinal gradient in Finland.

Tree Physiol 2015 Sep 19;35(9):975-86. Epub 2015 Jun 19.

Department of Environmental Science, University of Eastern Finland, PO Box 127, Kuopio, Finland.

Climate warming is having an impact on distribution, acclimation and defence capability of plants. We compared the emission rate and composition of volatile organic compounds (VOCs) from silver birch (Betula pendula (Roth)) provenances along a latitudinal gradient in a common garden experiment over the years 2012 and 2013. Micropropagated silver birch saplings from three provenances were acquired along a gradient of 7° latitude and planted at central (Joensuu 62°N) and northern (Kolari 67°N) sites. We collected VOCs emitted by shoots and assessed levels of herbivore damage of three genotypes of each provenance on three occasions at the central site and four occasions at the northern site. In 2012, trees of all provenances growing at the central site had higher total VOC emission rates than the same provenances growing at the northern site; in 2013 the reverse was true, thus indicating a variable effect of latitude. Trees of the southern provenance had lower VOC emission rates than trees of the central and northern provenances during both sampling years. However, northward or southward translocation itself had no significant effect on the total VOC emission rates, and no clear effect on insect herbivore damage. When VOC blend composition was studied, trees of all provenances usually emitted more green leaf volatiles at the northern site and more sesquiterpenes at the central site. The monoterpene composition of emissions from trees of the central provenance was distinct from that of the other provenances. In summary, provenance translocation did not have a clear effect in the short-term on VOC emissions and herbivory was not usually intense at the lower latitude. Our data did not support the hypothesis that trees growing at lower latitudes would experience more intense herbivory, and therefore allocate resources to chemical defence in the form of inducible VOC emissions.
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http://dx.doi.org/10.1093/treephys/tpv052DOI Listing
September 2015

Searching for common responsive parameters for ozone tolerance in 18 rice cultivars in India: Results from ethylenediurea studies.

Sci Total Environ 2015 Nov 11;532:230-8. Epub 2015 Jun 11.

University of Eastern Finland, Department of Biology, POB 111, 80101 Joensuu, Finland.

Eighteen rice (Oryza sativa) cultivars were screened for ozone (O3) tolerance and for the most responsive parameters with ethylenediurea (EDU) treatments at two experimental sites experiencing high ambient O3 conditions in the Indo-Gangetic Plains (IGP) of India. EDU was applied at 15 day intervals until the final harvest phase as a foliar spray at 300 ppm in order to protect the plants from the adverse effects of O3. Antioxidant activity, malondialdehyde content (MDA), chlorophyll content, gas exchange, and chlorophyll fluorescence (Fv/Fm) at the vegetative and flowering phases and harvest-related parameters were studied, for a total of 24 parameters. Seven of the studied cultivars had higher than average grainweightplant(-1) in all site and treatment combinations and can be recommended for cultivation in areas suffering from high O3 concentrations. The most responsive parameters with EDU treatment in high O3 across all cultivars were superoxide dismutase (SOD) and catalase (CAT) activities, the contents of oxidised (GSSG) and reduced (GSH) glutathione and MDA, and shoot weight plant(-1). These results indicated that the O3 scavenging activity of EDU is mediated through an antioxidant defence system rather than a direct effect on physiological parameters, such as photosynthesis and stomatal conductance.
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http://dx.doi.org/10.1016/j.scitotenv.2015.05.040DOI Listing
November 2015

Genetic and environmental determinants of insect herbivore community structure in a Betula pendula population.

F1000Res 2014 31;3:34. Epub 2014 Jan 31.

Faculty of Science and Forestry, Department of Biology, University of Eastern Finland, FIN-80101 Joensuu, Finland.

A number of recent studies have shown that intraspecific genetic variation of plants may have a profound effect on the herbivorous communities which depend on them. However less is known about the relative importance of intraspecific variation compared to other ecological factors, for example environmental variation or the effects of herbivore damage. We randomly selected 22 Betula pendula genotypes from a local population (< 0.9 ha), cloned them and planted cloned seedlings on two study sites separated at a regional scale (distance between sites about 30 km) to examine an insect community of 23-27 species on these genotypes. B. pendula genotypes did not differ in their species richness, but the total mean abundance and the structure of the insect herbivore community was significantly affected by the genotype, which could account for up to 27% of the total variation in community structure. B. pendula genotype accounted for two to four times more variation in the arthropod community structure than did environmental (block) variation on a local scale, while on a regional scale, genotypic and environmental (site) variation accounted for 4-14% of the arthropod community structure. The genetic effects were modified by environmental variation on both a local and regional scale over one study year, and locally, the largest part of the variation (38%) could be explained by the genotype × environment (block) interactions. Suppression of insect herbivores during one growing season led to changed arthropod community structure in the following growing season, but this effect was minimal and could explain only 4% of the total variation in insect community structure. Our results suggest that both genetic and environmental factors are important determinants of the community structure of herbivorous insects. Together these mechanisms appear to maintain the high diversity of insects in B. pendula forest ecosystems.
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http://dx.doi.org/10.12688/f1000research.3-34.v1DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3962004PMC
April 2014

Ozone affects ascorbate and glutathione biosynthesis as well as amino acid contents in three Euramerican poplar genotypes.

Tree Physiol 2014 Mar;34(3):253-66

INRA, UMR1137 EEF, F-54280 Champenoux, France.

Ozone is an air pollutant that causes oxidative stress by generation of reactive oxygen species (ROS) within the leaf. The capacity to detoxify ROS and repair ROS-induced damage may contribute to ozone tolerance. Ascorbate and glutathione are known to be key players in detoxification. Ozone effects on their biosynthesis and on amino acid metabolism were investigated in three Euramerican poplar genotypes (Populus deltoides Bartr. × Populus nigra L.) differing in ozone sensitivity. Total ascorbate and glutathione contents were increased in response to ozone in all genotypes, with the most resistant genotype (Carpaccio) showing an increase of up to 70%. Reduced ascorbate (ASA) concentration at least doubled in the two most resistant genotypes (Carpaccio and Cima), whereas the most sensitive genotype (Robusta) seemed unable to regenerate ASA from oxidized ascorbate (DHA), leading to an increase of 80% of the oxidized form. Increased ascorbate (ASA + DHA) content correlated with the increase in gene expression in its biosynthetic pathway, especially the putative gene of GDP-l-galactose phosphorylase VTC2. Increased cysteine availability combined with increased expression of γ-glutamylcysteine synthetase (GSH1) and glutathione synthetase (GSH2) genes allows higher glutathione biosynthesis in response to ozone, particularly in Carpaccio. In addition, ozone caused a remobilization of amino acids with a decreased pool of total amino acids and an increase of Cys and putrescine, especially in Carpaccio. In addition, the expression of genes encoding threonine aldolase was strongly induced only in the most tolerant genotype, Carpaccio. Reduced ascorbate levels could partly explain the sensitivity to ozone for Robusta but not for Cima. Reduced ascorbate level alone is not sufficient to account for ozone tolerance in poplar, and it is necessary to consider several other factors including glutathione content.
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http://dx.doi.org/10.1093/treephys/tpu004DOI Listing
March 2014

Carbohydrate concentrations and freezing stress resistance of silver birch buds grown under elevated temperature and ozone.

Tree Physiol 2013 Mar 20;33(3):311-9. Epub 2013 Feb 20.

Finnish Forest Research Institute, FIN-77600 Suonenjoki, Finland.

The effects of slightly elevated temperature (+0.8 °C), ozone (O3) concentration (1.3 × ambient O3 concentration) and their combination on over-wintering buds of Betula pendula Roth were studied after two growing seasons of exposure in the field. Carbohydrate concentrations, freezing stress resistance (FSR), bud dry weight to fresh weight ratio, and transcript levels of cytochrome oxidase (COX), alternative oxidase (AOX) and dehydrin (LTI36) genes were studied in two clones (clones 12 and 25) in December. Elevated temperature increased the bud dry weight to fresh weight ratio and the ratio of raffinose family oligosaccharides to sucrose and the transcript levels of the dehydrin (LTI36) gene (in clone 12 only), but did not alter the FSR of the buds. Genotype-specific alterations in carbohydrate metabolism were found in the buds grown under elevated O3. The treatments did not significantly affect the transcript level of the COX or AOX genes. No clear pattern of an interactive effect between elevated temperature and O3 concentration was found. According to these data, the increase in autumnal temperatures and slightly increasing O3 concentrations do not increase the risk for freeze-induced damage in winter in silver birch buds, although some alterations in bud physiology occur.
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http://dx.doi.org/10.1093/treephys/tpt001DOI Listing
March 2013

Needle metabolome, freezing tolerance and gas exchange in Norway spruce seedlings exposed to elevated temperature and ozone concentration.

Tree Physiol 2012 Sep 29;32(9):1102-12. Epub 2012 Aug 29.

Finnish Forest Research Institute, FIN-77600 Suonenjoki, Finland.

Northern forests are currently experiencing increasing mean temperatures, especially during autumn and spring. Consequently, alterations in carbon sequestration, leaf biochemical quality and freezing tolerance (FT) are likely to occur. The interactive effects of elevated temperature and ozone (O(3)), the most harmful phytotoxic air pollutant, on Norway spruce (Picea abies (L.) Karst.) seedlings were studied by analysing phenology, metabolite concentrations in the needles, FT and gas exchange. Sampling was performed in September and May. The seedlings were exposed to a year-round elevated temperature (+1.3 °C), and to 1.4× ambient O(3) concentration during the growing season in the field. Elevated temperature increased the concentrations of amino acids, organic acids of the citric acid cycle and some carbohydrates, and reduced the concentrations of phenolic compounds, some organic acids of the shikimic acid pathway, sucrose, cyclitols and steroids, depending on the timing of the sampling. Although growth onset occurred earlier at elevated temperature, the temperature of 50% lethality (LT(50)) was similar in the treatments. Photosynthesis and the ratio of photosynthesis to dark respiration were reduced by elevated temperature. Elevated concentrations of O(3) reduced the total concentration of soluble sugars, and tended to reduce LT(50) of the needles in September. These results show that alterations in needle chemical quality can be expected at elevated temperatures, but the seedlings' sensitivity to autumn and spring frosts is not altered. Elevated O(3) has the potential to disturb cold hardening of Norway spruce seedlings in autumn, and to alter the water balance of the seedling through changes in stomatal conductance (g(s)), while elevated temperature is likely to reduce g(s) and consequently reduce the O(3)-flux inside the leaves.
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http://dx.doi.org/10.1093/treephys/tps072DOI Listing
September 2012

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

Tree Physiol 2012 Jun 23;32(6):737-51. Epub 2012 Feb 23.

Department of Environmental Science, University of Eastern Finland, PO Box 1627, FIN-70211 Kuopio, Finland.

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.
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http://dx.doi.org/10.1093/treephys/tps005DOI Listing
June 2012

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

Tree Physiol 2011 Aug 18;31(8):808-18. Epub 2011 Aug 18.

Department of Biology, University of Eastern Finland, PO Box 111, FI-80101 Joensuu, Finland.

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.
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http://dx.doi.org/10.1093/treephys/tpr075DOI Listing
August 2011

Leaf volatile emissions of Betula pendula during autumn coloration and leaf fall.

J Chem Ecol 2010 Oct 14;36(10):1068-75. Epub 2010 Sep 14.

Department of Environmental Science, University of Eastern Finland, Kuopio Campus, P.O. Box 1627, FIN-70210, Kuopio, Finland.

Deciduous trees remobilize the nitrogen in leaves during the process of autumn coloration, thus providing a high quality food source for aphids preparing to lay over-wintering eggs. It has been suggested that aphids may use volatile organic compounds (VOCs) to: (a) select leaves where nutrient remobilization has started and induced defenses are reduced; and (b) detect the time of leaf abscission. We analyzed VOCs emitted by the foliage of Betula pendula Roth. during autumn coloration and from leaf litter just after leaf fall. We tested the hypothesis that costly, photosynthesis-related terpenes and other herbivore-induced VOCs related to attraction of aphid parasitoids and predators are reduced during the coloration process. We also investigated if the VOC emission profile of abscising leaves is different from that of early stage yellowing leaves. Enemy-luring compounds (E)-β-ocimene, linalool, and (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT) were emitted only from the green foliage. Methyl salicylate (MeSa), known to recruit predatory bugs and attract migrant aphids, was emitted until the first stage of color change. Cis-3-hexenol, an indicator of cellular disintegration, became dominant in the emissions from abscising leaves and from fresh leaf litter. We discuss the ecological significance of the observed changes in birch leaf VOC profiles during the process of autumn senescence.
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http://dx.doi.org/10.1007/s10886-010-9857-4DOI Listing
October 2010

Volatile emissions and phenolic compound concentrations along a vertical profile of Populus nigra leaves exposed to realistic ozone concentrations.

Photosynth Res 2010 Apr 21;104(1):61-74. Epub 2010 Apr 21.

Consiglio Nazionale delle Ricerche (CNR), Istituto di Biologia Agroambientale e Forestale (IBAF), Rome, Italy.

Plants are exposed to increasing levels of tropospheric ozone concentrations. This pollutant penetrates in leaves through stomata and quickly reacts inside leaves, thus making plants valuable ozone sinks, but at the same time triggers oxidation processes which lead to leaf injuries. To counteract these negative effects, plants produce an array of antioxidants which react with ozone and reactive molecules which ozone generates in the leaf tissues. In this study, we measured the effect of an ozone concentration which is likely to be attained in many areas of the world in the near future (80 ppb) on leaves of the vertical profile of the widespread agroforestry species Populus nigra. Changes in (1) physiological parameters (photosynthesis and stomatal conductance), (2) ozone uptake, (3) emission of volatile organic compounds (VOCs, i.e. isoprene, methanol and other oxygenated compounds), (4) concentration of antioxidant surface compounds, and (5) concentration of phenolic compounds were assessed. The aim was to assess whether the defensive pathways leading to isoprenoids and phenolics formation were induced when a moderate and chronic increment of ozone is not able to damage photosynthesis. No visual injuries and minor changes in physiology and ozone uptake were observed. The emission of isoprene and oxygenated six-carbon (C6) volatiles were inhibited by ozone, whereas methanol emission was increased, especially in developing leaves. We interpret these results as suggesting an ontogenetic shift in ozone-treated leaves, leading to a slower development and a faster senescence. Most surface and phenolic compounds showed a declining trend in concentration from the youngest to the fully expanded leaves. Ozone reduced the concentrations of chlorogenic acid derivatives at the leaf surface, whereas in total leaf extracts a metabolic shift towards few phenolics with higher antioxidant capacity was observed.
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http://dx.doi.org/10.1007/s11120-010-9549-5DOI Listing
April 2010

Red clover (Trifolium pratense L.) isoflavones: root phenolic compounds affected by biotic and abiotic stress factors.

J Sci Food Agric 2010 Feb;90(3):418-23

Institute of Applied Biotechnology, Department of Biosciences, University of Kuopio, Box 1627, 70211 Kuopio, Finland.

Background: Phenolic compounds have recently received considerable attention for their ability to protect plant and human cells from oxidative stress-induced damage. Red clover (Trifolium pratense L.) is a rich source of isoflavonoids with multiple potential protective functions. The aim of this study was to identify and characterise phenolic compounds in red clover roots by high-performance liquid chromatography and mass spectrometry and to study the effects of stress factors and growth stage on root phenolics.

Results: A total of 28 phenolic compounds were tentatively identified in red clover roots. The most abundant phenolics in pot-grown roots were formononetin glycoside malonate (G-M) (1.51-4.26 mg g(-1)), formononetin (2.21-3.57 mg g(-1)) and biochanin A (1.73-2.17 mg g(-1)), whereas field-grown roots were rich in formononetin-G-M (3.90-4.27 mg g(-1)), maackiain-G-M (2.35-3.02 mg g(-1)) and pseudobaptigenin-G-M (1.80-2.58 mg g(-1)). Concentrations were affected by the growth stage. Ozone exposure slightly affected the total phenolic content in roots and also had minor effects on individual compounds.

Conclusion: Elevated ozone, cultivation regime and growth stage affected the levels of phenolics in red clover roots, suggesting sensitivity of root phenolics to biotic and abiotic stress conditions. The high levels of phenolics found in roots even in late autumn may be utilised in many applications.
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http://dx.doi.org/10.1002/jsfa.3831DOI Listing
February 2010

Elevation of night-time temperature increases terpenoid emissions from Betula pendula and Populus tremula.

J Exp Bot 2010 Jun 24;61(6):1583-95. Epub 2010 Feb 24.

Department of Environmental Science, University of Eastern Finland, PO Box 1627, FI-70211 Kuopio, Finland.

Volatile organic compounds (VOCs) are expected to have an important role in plant adaptation to high temperatures. The impacts of increasing night-time temperature on daytime terpenoid emissions and related gene expression in silver birch (Betula pendula) and European aspen (Populus tremula) clones were studied. The plants were grown under five different night-time temperatures (6, 10, 14, 18, and 22 degrees C) while daytime temperature was kept at a constant 22 degrees C. VOC emissions were collected during the daytime and analysed by gas chromatography-mass spectrometry (GC-MS). In birch, emissions per leaf area of the C11 homoterpene 4,8-dimethy1-nona-1,3,7-triene (DMNT) and several sesquiterpenes were consistently increased with increasing night-time temperature. Total sesquiterpene (SQT) emissions showed an increase at higher temperatures. In aspen, emissions of DMNT and beta-ocimene increased from 6 degrees C to 14 degrees C, while several other monoterpenes and the SQTs (Z,E)-alpha-farnesene and (E,E)-alpha-farnesene increased up to 18 degrees C. Total monoterpene and sesquiterpene emission peaked at 18 degrees C, whereas isoprene emissions decreased at 22 degrees C. Leaf area increased across the temperature range of 6-22 degrees C by 32% in birch and by 59% in aspen. Specific leaf area (SLA) was also increased in both species. The genetic regulation of VOC emissions seems to be very complex, as indicated by several inverse relationships between emission profiles and expression of several regulatory genes (DXR, DXS, and IPP). The study indicates that increasing night temperature may strongly affect the quantity and quality of daytime VOC emissions of northern deciduous trees.
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http://dx.doi.org/10.1093/jxb/erq034DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2852659PMC
June 2010

Impact of experimentally elevated ozone on seed germination and growth of Russian pine (Pinus sylvestris) and spruce (Picea spp.) provenances.

Ambio 2009 Dec;38(8):443-7

Institute of Ecological Problems in the North of Ural Branch of Russian Acedemy of Sciences, Arkhangelsk, Russia.

The impact of elevated ozone concentrations on early ontogenetic stages of pine (Pinus sylvestris) and spruce (Picea abies, Picea obovata, P. abies x P. obovata) seedlings originating from different provenances in Russia were studied in the open-field ozone fumigation system located in Kuopio, Finland, over a span of 2 y. The AOT40 value (accumulated ozone dose over the threshold 40 ppb during daylight hours) was 11 ppm hr per growing season, which was 1.4 times higher than the ambient air concentration. The plants were measured for germination rate; shoot increment; needle length; and dry mass of needles, shoots, and roots. Significant differences between pine and spruce provenance response to ozone were found in all parameters. Ozone stress immediately reduced the germination rate of Northern pine provenances, whereas biomass reductions became evident during the second year of the exposure in all pine provenances. Spruce species were more tolerant to elevated ozone concentrations. Our results indicate that seedling development is vulnerable to increasing ozone concentrations and that attention must be paid to the provenance selection.
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http://dx.doi.org/10.1579/0044-7447-38.8.443DOI Listing
December 2009

Rising atmospheric CO2 concentration partially masks the negative effects of elevated O3 in silver birch (Betula pendula Roth).

Ambio 2009 Dec;38(8):418-24

The Finnish Forest Research Institute, Suonenjoki Research Unit, Finland.

This review summarizes the main results from a 3-year open top chamber experiment, with two silver birch (Betula pendula Roth) clones (4 and 80) where impacts of 2x ambient [CO2] (EC) and [O3] (EO) and their combination (EC + EO) were examined. Growth, physiology of the foliage and root systems, crown structure, wood properties, and biological interactions were assessed to understand the effects of a future climate on the biology of silver birch. The clones displayed great differences in their reaction to EC and EO. Growth in clone 80 increased by 40% in EC and this clone also appeared O3-tolerant, showing no growth reduction. In contrast, growth in clone 4 was not enhanced by EC, and EO reduced growth with root growth being most affected. The physiological responses of the clones to EO were smaller than expected. We found no O3 effect on net photosynthesis in either of the clones, and many parameters indicated no change compared with chamber controls, suggesting active detoxification and defense in foliage. In EO, increased rhizospheric respiration over time and accelerated leaf senescence was common in both clones. We assumed that elevated O3 offsets the positive effects of elevated CO2 when plants were exposed to combined EC + EO treatment. In contrast, the responses to EC + EO mostly resembled the ones in EC, at least partly due to stomatal closure, which thus reduced O3 flux to the leaves. However, clear cellular level symptoms of oxidative stress were observed also in EC + EO treatment. Thus, we conclude that EC masked most of the negative O3 effects during long exposure of birch to EC + EO treatment. Biotic interactions were not heavily affected. Only some early season defoliators may suffer from faster maturation of leaves due to EO.
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http://dx.doi.org/10.1579/0044-7447-38.8.418DOI Listing
December 2009

Near-ambient ozone concentrations reduce the vigor of Betula and Populus species in Finland.

Ambio 2009 Dec;38(8):413-7

University of Joensuu, Faculty of Biosciences, Finland.

In this review the main growth responses of Finnish birch (Betula pendula, B. pubescens) and aspen species (Populus tremula and P. tremuloides x P. tremula) are correlated with ozone exposure, indicated as the AOT40 value. Data are derived from 23 different laboratory, open-top chamber, and free-air fumigation experiments. Our results indicate that these tree species are sensitive to increasing ozone concentrations, though high intraspecific variation exists. The roots are the most vulnerable targets in both genera. These growth reductions, determined from trees grown under optimal nutrient and water supply, were generally accompanied by increased visible foliar injuries, carbon allocation toward defensive compounds, reduced carbohydrate contents of leaves, impaired photosynthesis processes, disturbances in stomatal function, and earlier autumn senescence. Because both genera have shown complex ozone defense and response mechanisms, which are modified by variable environmental conditions, a mechanistically based approach is necessary for accurate ozone risk assessment.
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http://dx.doi.org/10.1579/0044-7447-38.8.413DOI Listing
December 2009

Differential gene expression in senescing leaves of two silver birch genotypes in response to elevated CO2 and tropospheric ozone.

Plant Cell Environ 2010 Jun 29;33(6):1016-28. Epub 2010 Jan 29.

Suonenjoki Research Unit, Finnish Forest Research Institute, 77600 Suonenjoki, Finland.

Long-term effects of elevated CO(2) and O(3) concentrations on gene expression in silver birch (Betula pendula Roth) leaves were studied during the end of the growing season. Two birch genotypes, clones 4 and 80, with different ozone growth responses, were exposed to 2x ambient CO(2) and/or O(3) in open-top chambers (OTCs). Microarray analyses were performed after 2 years of exposure, and the transcriptional profiles were compared to key physiological characteristics during leaf senescence. There were genotypic differences in the responses to CO(2) and O(3). Clone 80 exhibited greater transcriptional response and capacity to alter metabolism, resulting in better stress tolerance. The gene expression patterns of birch leaves indicated contrasting responses of senescence-related genes to elevated CO(2) and O(3). Elevated CO(2) delayed leaf senescence and reduced associated transcriptional changes, whereas elevated O(3) advanced leaf senescence because of increased oxidative stress. The combined treatment demonstrated that elevated CO(2) only temporarily alleviated the negative effects of O(3). Gene expression data alone were insufficient to explain the O(3) response in birch, and additional physiological and biochemical data were required to understand the true O(3) sensitivity of these clones.
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http://dx.doi.org/10.1111/j.1365-3040.2010.02123.xDOI Listing
June 2010

Real-time monitoring of herbivore induced volatile emissions in the field.

Physiol Plant 2010 Feb 9;138(2):123-33. Epub 2009 Nov 9.

Ionicon Analytik GmbH, Technikerstrasse 21a, 6020 Innsbruck, Austria.

When plants are damaged by herbivorous insects they emit a blend of volatile organic compounds (VOCs) which include a range or terpenoids and green leaf volatiles (GLVs) formed via different metabolic pathways. The precise timing of these emissions upon the onset of herbivore feeding has not been fully elucidated, and the information that is available has been mainly obtained through laboratory based studies. We investigated emissions of VOCs from Populus tremula L. xP. tremuloides Michx. during the first 20 h of feeding by Epirrita autumnata (autumnal moth) larvae in a field site. The study was conducted using Proton Transfer Reaction-Mass Spectrometry (PTR-MS) to measure emissions online, with samples collected for subsequent analysis by complementary gas chromatography-mass spectrometry for purposes of compound identification. GLV emission peaks occurred sporadically from the outset, indicating herbivore activity, while terpene emissions were induced within 16 h. We present data detailing the patterns of monoterpene (MT), GLV and sesquiterpene (SQT) emissions during the early stages of herbivore feeding showing diurnal MT and SQT emission that is correlated more with temperature than light. Peculiarities in the timing of SQT emissions prompted us to conduct a thorough characterization of the equipment used to collect VOCs and thus corroborate the accuracy of results. A laboratory based analysis of the throughput of known GLV, MT and SQT standards at different temperatures was made with PTR-MS. Enclosure temperatures of 12, 20 and 25 degrees C had little influence on the response time for dynamic measurements of a GLV or MT. However, there was a clear effect on SQT measurements. Elucidation of emission patterns in real-time is dependent upon the dynamics of cuvettes at different temperatures.
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http://dx.doi.org/10.1111/j.1399-3054.2009.01322.xDOI Listing
February 2010

Gene expression responses of paper birch (Betula papyrifera) to elevated CO2 and O3 during leaf maturation and senescence.

Environ Pollut 2010 Apr 3;158(4):959-68. Epub 2009 Nov 3.

Finnish Forest Research Institute, Suonenjoki, Finland.

Gene expression responses of paper birch (Betula papyrifera) leaves to elevated concentrations of CO(2) and O(3) were studied with microarray analyses from three time points during the summer of 2004 at Aspen FACE. Microarray data were analyzed with clustering techniques, self-organizing maps, K-means clustering and Sammon's mappings, to detect similar gene expression patterns within sampling times and treatments. Most of the alterations in gene expression were caused by O(3), alone or in combination with CO(2). O(3) induced defensive reactions to oxidative stress and earlier leaf senescence, seen as decreased expression of photosynthesis- and carbon fixation-related genes, and increased expression of senescence-associated genes. The effects of elevated CO(2) reflected surplus of carbon that was directed to synthesis of secondary compounds. The combined CO(2)+O(3) treatment resulted in differential gene expression than with individual gas treatments or in changes similar to O(3) treatment, indicating that CO(2) cannot totally alleviate the harmful effects of O(3).
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http://dx.doi.org/10.1016/j.envpol.2009.10.008DOI Listing
April 2010

Leaf phenolic compounds in red clover (Trifolium pratense L.) induced by exposure to moderately elevated ozone.

Environ Pollut 2010 Feb 18;158(2):440-6. Epub 2009 Sep 18.

University of Kuopio, Department of Biosciences, Institute of Applied Biotechnology, Box 1627, 70211 Kuopio, Finland.

Red clover (Trifolium pratense L.), an important feed crop in many parts of the world, is exposed to elevated ozone over large areas. Plants can limit ozone-induced damages by various defence mechanisms. In this work, changes in the concentrations of antioxidant phenolic compounds induced by slightly elevated levels of ozone were determined in red clover leaves by high-performance liquid chromatography and mass spectrometry. 31 different phenolics were identified and the most abundant isoflavones and flavonoids were biochanin A glycoside malonate (G-M), formononetin-G-M and quercetin-G-M. Elevated ozone (mean 32.4 ppb) increased the total phenolic content of leaves and also had minor effects on the concentrations of individual compounds. Elevated ozone increased the net photosynthesis rate of red clover leaves before visible injuries by 21-23%. This study thus suggests that the concentrations of phenolics in red clover leaves change in response to slightly elevated ozone levels.
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http://dx.doi.org/10.1016/j.envpol.2009.08.029DOI Listing
February 2010