Current research in air pollution and climate change effects on northern terrestrial ecosystems Photo Jouni Hyvärinen Symposium in the honour of professor Satu Huttunen 1-2 December 2011 Department of Biology, University of Oulu ORGANIZERS Dr. Jaana Bäck Dr. Sirkku Manninen Dr. Pasi Rautio Dr. Eero Kubin Local organizers Professor Juha Tuomi M. Sc. Ritva Hiltunen With financial support of: FEDERATION OF FINNISH LEARNED SOCIETIES FINNISH AIR POLLUTION PREVENTION SOCIETY 2 Dear Satu, dear colleagues and friends, We are happy to see you all gathered here to recognise the valuable contributions of professor Satu Huttunen as researcher and university teacher. The aim of this symposium is to cover the broad field of plant ecophysiological, ecological and environmental research which You, Satu, have been involved in. The scope of oral presentations and posters ranges from effects of individual air pollutants or climate change factors on invidual species, to those at the global scale. The speakers and the other guests are scientists whose ways have crossed with those of Satu’s during the course of long career. I am one of Satu’s students from the 1980’s, but her career in the field of air pollution effects on vegetation started already long before that. Satu’s Ph.D. thesis entitled ”Influence of air pollution on the forest vegetation around Oulu” defended already in 1975. You, Satu. wrote in the introduction of Your doctoral dissertation: ”Chronic long-term injuries due to air pollution are of particular importance in cold northern climate and they constitute a continual stress factor in the ecosystem near industrial processes and urban areas. Air pollution has a clear effect on the growth of conifers several years before first visible damage appears. The present research has served to indicate that the damage found in Oulu is attributable to the combined effects of a number of pollutants over a long period. It is continuous presence of relatively small amounts of toxic substances together with a powerful fertilizer component which has repeatedly led to winter forest vegetation damage. The actual extent and severity of the damage in any one year is dependent on weather conditions, but the fundamental cause is to be found in the continuous long-term fertilizer loading and accumulation of toxins. This long-term accumulation process is extremely difficult to demonstrate, as seasonal variations and other natural features cannot be directly linked with the content figures obtained from chemical and other analyses.” Following Your example, many of Your students did field and experimental studies on the effects of sulphur and nitrogen emissions, acid deposition and heavy metals on trees in the late 1980’s and early 1990’s. Then we began studies on the effects of elevated tropospheric ozone on northern tree provenaces, while some of Your students started to study first tree and later bryophyte responses to elevated UV-B radiation. During all these years, issues related to the effects of climate, especially low temperatures, on vegetation have been in Your scientific agenda. You, Satu, also became internationally active already in the 1970’s. For example, You have acted as air pollution scientific expert in the scope of Scientific-Technical Cooperation between Finland and the Soviet Union in Air Pollution Questions and as a Nominated expert of the UN/ECE Working Group on Effects since 1978. You have also been active within the IUFRO (International Union of Forest Research Organizations) in terms of both air pollution and climate change issues and chaired the COST Action E 6: EUROSILVA on Tree Physiology in the late 1990’s. The map of Your study and scientific congress visits etc. includes most of the European countries including the former Soviet Union as well as North and South America. 3 Although You have been internationally very active, You have always remained loyal to Oulu and its university, where You were appointed first as an acting professor in 1981 and as a full professor in the physiological ecology of plants and environmental ecology in 1989. After Your dissertation and before getting the professorship, You worked for almost 15 years in different researcher positions of the Academy of Finland. I remember You having been very busy during the 1980’s. In addition to running large Academy research projects and being a member of its Research Council of the Environmental Sciences, You were running several regional bioindicator studies in Finland, acting as a member of numerous Finnish state commissions, scientific associations and foundations, and as the president of the Finnish Association for Nature Conservation. During Your career You have published over 200 scientific papers, most of these in international refereed journals. You have supervised 23 doctoral dissertations at the University of Oulu and acted as co-supervisor for a number of doctoral dissertations at foreign universities. Your CV also includes 15 supervised licentiate dissertations and 78 M.Sc. theses. Hence You are one of the most productive teachers, if not the most productive, in the history of the University of Oulu in terms of supervision of student theses. The whole research field of plant ecophysiology at the University of Oulu was created and developed essentially due to Your activities in educating and training young scientists in the field. As a female professor, You have always been a role-model for your female students, though your achievements have been highly appreciated also by the society in general. You were elected as the Finnish Woman of the Year in 1986 and got the UNEP’s Global 500 award in 1987. With these words, we warmly welcome You, Satu, and all Your colleagues and friends to this symposium. We hope that You enjoy the presentations, discussions and our company. Helsinki and Rovaniemi, November, 2011 Sirkku Manninen, Jaana Bäck and Pasi Rautio Satu as an Academy researcher in the late 1970’s in her office. 4 Programme Thursday, 1 December 2011 13.00-14.00 Registration 14.00-14.15 Welcome Sirkku Manninen, University of Helsinki Seppo Saarela, Head of Department of Biology, University of Oulu Session I Carbon sequestration (Chair Pasi Rautio, Finnish Forest Research Institute) 14.15-14.50 Forest transition and the global trends of tree biomass Pekka Kauppi, University of Helsinki, Finland 14.50-15.25 Winter and spring in northern forests: dynamic features and their consequences to climate change Jaana Bäck, University of Helsinki, Finland 15.25-16.00 Mycorrhizal fungal symbionts of forest trees on carbon and nutrient gradients - implications for climate change Annamari Markkola, University of Oulu, Finland 16.00-16.30 Coffee and tea (posters) Session II Species and community responses (Chair Pasi Rautio, Finnish Forest Research Institute) 16.30-17.05 Dwarf shrubs under stress related to global change and air pollution Kari Taulavuori, University of Oulu, Finland 17.05-17.40 Polluted areas of the Kola Peninsula: effects of multiple stressors on ecosystem structure and functions Mikhail Kozlov, University of Turku, Finland 17.40-18.15 Climate change effects on reindeer pastures in Northern Scandinavia Minna Turunen, Arctic centre, Rovaniemi, Finland 18.15-18.35 General Discussion 19-21 Buffet dinner at Botanical garden 5 Friday, 2 December 2011 Session III Forest health (Chair Sirkku Manninen, University of Helsinki) 9.00-9.45 Forest health monitoring from the 1980 to 2010, achievements, problems and perspectives Marco Ferretti, TerraData environmetrics, spin off Accademico dell'Università di Siena, Italy 9.45-10.30 Decline and recovery of forests in the Black Triangle area of Europe Josef Krecek, Czech Technical University, Prague 10.30-11.00 Poster session Maija Salemaa (Finnish Forest Research Institute): Nitrogen fixation activity by cyanobacteria growing on forest mosses increase towards the north in Finland Minna Turunen (Arctic Centre/University of Lapland, Finland): Response of reindeer forage plant phenolics to UV radiation Anna Hyyryläinen (University of Oulu, Finland): Seasonal dynamics in Sphagnum chlorophylls under altered UVB and temperature 11.00-11.30 Coffee and tea (posters) Session IV Effects on bryophytes (Chair Sirkku Manninen, University of Helsinki) 11.30-12.05 Reactive nitrogen deposition: too much of a good thing for peatland ecosystems Lucy Sheppard, Centre for Ecology and Hydrology, Edinburgh, Scotland 12.05-12.40 Ultraviolet radiation and bryophytes Javier Martinez-Abaigar, University La Rioja, Spain 12.40-13.30 Lunch Session V Phenological and biochemical responses (Chair Kari, Taulavuori, University of Oulu) 13.30-14.05 Growth of primordial shoot in Norway spruce buds preceding visible bud burst in relation to accumulated temperature in the field Sirkka Sutinen, Finnish Forest Research Institute, Joensuu, Finland 14.05-14.40 Impact of geoclimate conditions on triterpenoid content of fruit and leaf cuticular waxes from Finnish and Polish Vaccinium vitisidaea and Vaccinium myrtillus Anna Szakiel, University of Warsaw, Poland 6 14.40- 15.10 Coffee and tea (posters) Session VI Ozone, nitrogen and climate effects (Chair Kari, Taulavuori, University of Oulu) 15.10-15.45 Plant acclimation under air pollution stress and changing climate Sirkku Manninen, University of Helsinki, Finland 15.45-16.20 Prospects for future ozone effects on vegetation in the Nordic countries with consideration of climate change Håkan Pleijel, Göteborg university, Sweden 16.20-16.45 General discussion and closing remarks – Emerita professor Satu Huttunen Photo Jouni Hyvärinen 7 ABSTRACTS ORAL PRESENTATIONS 8 FOREST TRANSITION AND THE GLOBAL TRENDS OF TREE BIOMASS Pekka E. Kauppi Department of Environmental Sciences, University of Helsinki, P.O. Box 56, FI-00014 Helsinki, Finland Contact: [email protected] Like humans, forest trees have a life span from birth to death. Tree populations in analogy to human populations can be characterized with a population pyramid. Unlike humans, trees keep accumulating biomass through their life time. Unlike even the largest humans, big trees are heavy. Changes in the demography and hence the carbon content forest vegetation significantly affects the carbon dioxide concentration of the atmosphere. ‘Forest transition’ refers to a shift from shrinking to expanding forests within a region, nation, continent, or globally. Alexander Mather of University of Aberdeen, Scotland, was a leading analyst documenting the diffusion of forest transition on Earth. Unfortunately, the World has not yet reached forest transition in terms of forest area. Deforestation continues. Detailed analyses of forest attributes, not surprisingly, paint a picture of changing demography of tree populations. A common direction of change is a shift toward aging populations, in analogy to Europeans, Japanese and the Finns “becoming gray”. In carbon sequestration analysis, the population pyramid of trees is described noting the biomass accumulation of each tree during the lifetime. Interestingly, the population of dead trees accumulates necromass and sequesters carbon significantly in boreal forests at the present time. Shifts in tree populations over time and space respond to direct and indirect drivers. An expansion of forest biomass persists in Europe, North America and many regions of Asia. The causes and consequences of this development are only partially understood even in countries like Finland, where ample data are available. The large and highly competent community of scientists working on forest ecosystems research can look forward to interesting future work. At the global level, there is an urgent need for improved observation systems, which are capable of detecting changes in tree demography. Reference Pan, Y., Birdsey, R.A., Fang, J., Houghton, R., Kauppi, P.E., Kurz, W.A., Phillips, O.L., Shvidenko, A., Lewis, S.L., Canadell, J.G., Ciais, P., Jackson, R.B., Pacala, S., McGuire, D., Piao, S., Rautiainen, A., Stitch, S., Hayes, D., 2011. A Large and Persistent Carbon Sink in the World’s Forests. Science 333: 988-993. 9 WINTER AND SPRING IN NORTHERN FORESTS: DYNAMIC FEATURES AND THEIR CONSEQUENCES TO CLIMATE CHANGE Jaana Bäck1, 2, Pertti Hari2, Albert Porcar-Castell2, Juho Aalto2, 3 1 2 Department of Physics, P.O. Box 64, FI-00014 University of Helsinki, Finland Department of Forest Sciences, P.O. Box 27, FI-00014 University of Helsinki, Finland 3 Hyytiälä Forestry Field Station, Hyytiäläntie 124, FI-35500 Korkeakoski, Finland Contact: [email protected] Strong environmental constraints limit the plant performance in northern areas. The photosynthesis and growth of trees is synchronised with the annual cycle of temperature. In winter and early spring the photosynthetic rate of evergreen foliage is very low. An actively operating photosynthetic system would be unable to tolerate the hard conditions in early spring with low temperatures and high light intensity, and therefore a downregulation of photosynthetic machinery needs to take place during the hardening process. In dehardening, the activity is recovered and the CO2 uptake capacity gradually increases. However, during the dehardening process, other sinks for excess energy may be needed if the photosystems are still partially in an inactive stage. Volatile organic compounds (VOCs), especially monoterpenes, are constitutively produced and emitted from evergreen foliage. Emissions have traditionally been linked to ambient temperatures in an exponential manner, and thus considered to be highest in summertime. However, field emission rate measurements in early spring show high peaks during the period of dehardening. In this presentation we’ll evaluate the potential mechanisms related to VOC emissions in conifer foliage, and the importance of phenomena related to spring recovery of photosynthesis and onset of growth for measured emissions. Results from continuous field measurements are presented, and the consequences to climate change discussed. 10 MYCORRHIZAL FUNGAL SYMBIONTS OF FOREST TREES ON CARBON AND NITROGEN GRADIENTS - IMPLICATIONS FOR CLIMATE CHANGE Annamari Markkola1, Karita Saravesi1, Pasi Rautio2, Oili Tarvainen3, Anna Liisa Ruotsalainen1, Maarit Kaukonen1, Piippa Wäli1 and Rauni Strömmer4 1 Department of Biology, University of Oulu, P.O. Box 3000, FI-90014 Oulu, Finland 2 Finnish Forest Research Institute, P.O. Box 16, FI-96301 Rovaniemi, Finland 3 Finnish Forest Research Institute, Kirkkosaarentie, FI-9500 Muhos, Finland 4 Department of Environmental Sciences, University of Helsinki, Niemenkatu 73, FI-15140 Lahti, Finland Contact: [email protected] Ectomycorrhizal (ECM) fungal symbiosis is obligatory for nutrient uptake of forest trees, although the symbiosis results in high costs for the hosts. ECM fungi form large carbon sinks: up to 20 % of net photosynthesis products assimilated by the tree is allocated to the fungal symbionts. Mycorrhizae and fungal mycelia are thus an important pathway for carbon sequestration into the forest soil. A typical feature for ECM fungal communities is their high diversity: one host tree may harbour several dozens of fungal species in their roots. Fungal symbionts are known to differ in their carbon demands, both for their metabolic and structural needs. Climate change affects carbon sequestration into boreal and subarctic forest soils firstly due to enhanced photosynthesis in elevated atmospheric CO2 environment, soil warming and consequent nutrient mineralization in soil. Belowground allocation is often reported to increase in elevated atmospheric CO2 with a consequent increase in tree root and mycorrhizal biomass, especially in low nutrient conditions. In contrast, severe carbon limitation in forest trees may be created periodically by massive herbivore outbreaks, which may become more common along with the climate change in boreal and subarctic forests. Consequently, tree root biomass will be declined. Moreover, severe defoliation and foliar damage of the host trees have been found to alter composition of ECM fungal communities: thus, a relative decrease in mycorrhizal morphotypes with abundant external fungal mycelia (potential high C demand) and an increase in morphotypes with low amount of mycelia (potential low C demand) has been reported under limited C availability of the host. This accelerates the decline in carbon sequestration in ECM fungal mycelia and into the soil. On the other hand, a decline in those ECM species producing abundantly and/or large sporocarps and high C demanding mycorrhizal morphotypes with abundant external mycelia, have been reported under high soil N availability, especially in coniferous forests. A possible mechanism for these parallel changes in ECM communities associated with forest trees may lie in the change of cost to benefit ratios along carbon and nitrogen gradients. It is discussed, whether the observed changes in ECM communities are reversible in a long run, and whether they in turn might affect carbon storage in boreal and subarctic forest soils. 11 ERICACEOUS DWARF SHRUBS UNDER STRESS RELATED TO GLOBAL CHANGE AND AIR POLLUTION Kari Taulavuori1, Erja Taulavuori1 and Kari Laine2 1 Department of Biology, University of Oulu, P.O.Box 3000, FI-90014 Oulu, Finland 2 Thule Institute, University of Oulu, P.O. Box 7300, FI-90014 Oulu, Finland Contact: [email protected] Ericaceous dwarf shrubs are dominant species in the field layer of boreal heath forests. Ecophysiological studies on the Vaccinium sp. have long traditions in Oulu since 1960’s, and currently these species could be understood as model species of boreal ecosystems. This presentation focuses on studies related to air pollution and global change in Oulu since 1990’s. The responses are studied mainly from the seasonality, frost hardiness and oxidative stress perspectives. From the air pollution point of view we have studied these responses in OTC experiment with CO2 and O3 fumigations, with and without nitrogen supplement. Some interests were also towards responses to heavy metals. From the global change point of view, main interest was in overwintering under warming climate. This interest broadened also to lack of snow in winter, and summer droughts. In addition, response to enhanced UV-radiation is one of the research topics. Overall conclusion is that ericaceous dwarf shrubs are relatively tolerant against the studied environmental stress. 12 POLLUTED AREAS OF THE KOLA PENINSULA: EFFECTS OF MULTIPLE STRESSORS ON ECOSYSTEM STRUCTURE AND FUNCTIONS Mikhail V. Kozlov, Vitali Zverev and Elena L. Zvereva Section of Ecology, University of Turku, FI-20014 Turku, Finland Contact: [email protected] Monchegorsk, an industrial town in Murmansk District of Russia, is well known to researchers studying pollution effects on biota. Over 2000 publications were produced by at least 50 teams that have been exploring various aspects of industrial pollution in this region since the 1980s. A recent research synthesis (Kozlov & Zvereva 2011) included 89 effect sizes calculated from studies conducted near Monchegorsk, while an average number of effect sizes per polluter was 5.3 only. We conclude that, due to a number of unique characteristics, the impact zone of the nickel-copper smelter at Monchegorsk is the best studied example of local pollution impact on terrestrial biota. The achieved level of knowledge allows uncovering some of mechanisms behind the observed changes in structure and functions of polluted ecosystems. Ground survey of 2007 demonstrated that 66 km2 of the previously forested land area around Monchegorsk is transformed to industrial barrens - bleak open landscapes with only small patches of vegetation surrounded by bare land. Life persisting in these habitats suffers not only from the toxicity of pollutants but also from other stressors including low nutritional quality of soils, soil erosion, high wind speed causing sandblast and snow abrasion, and altered (more harsh) microclimate. Many species have evolved adaptations to this extreme environment. In particular, mountain birch (dominant tree species in industrial barrens) demonstrated not only tolerance to heavy metals, but also heritable changes in crown architecture, such as an increased production of long shoots. However, study of demographic processes showed that due to absence of natural regeneration (hampered by soil toxicity) populations of many plant species persisting in industrial barrens are aging, which (in spite of the discovered adaptations) may soon lead to their extinction. Decrease of emissions that occurred about 15 years ago has not yet resulted in the improvement of the ecological situation in most contaminated areas, where both birch populations and field layer vegetation continue to decline. The very first signs of recovery, i.e., naturally emerging Scots pine seedlings, have recently (in 2009-2010) been observed only at the outer border of industrial barrens. Changes in species composition caused by pollution and associated stressors alter biotic interactions and consequently affect ecosystem functions and services. However, a vast majority of case studies address organism- and population-level effects; the reported community-level effects concern primarily changes in species richness, while studies exploring even the key ecosystem functions (such as net primary productivity) are scarce. Although the existing data suggest that all ecosystem services (provisioning, regulating, supporting, and cultural) are adversely affected by pollution, further research is badly needed to understand the causal links behind these effects and examine dose dependence in order to quantitatively predict the fate of polluted ecosystems under different economic and climatic scenarios. 13 CLIMATE CHANGE EFFECTS ON REINDEER PASTURES IN NORTHERN FENNOSCANDIA Minna Turunen1, Francoise Martz2, Terhi Vuojala-Magga1 1 Arctic Centre, University of Lapland, P.O. Box 122, FI-96101 Rovaniemi, Finland 2 FFRI, Rovaniemi Research Unit, P.O. Box 16, FI-96301 Rovaniemi, Finland Contact: [email protected] In Northern Fennoscandia, the reindeer graze freely on natural pastures during the spring, summer, autumn and early winter. Its pasture habitats range from boreal coniferous forests to subarctic mountain birch woodlands, tundra, open mountains, peatlands and riverbanks. Reindeer herding, which is a meat-producing economic activity and has cultural relevance to local and indigenous peoples, is affected by global change including changes in climate and the socio-political environment. The aim of this presentation is to give a review about our research on the effects of climate change on reindeer pastures, conducted within the projects of ECOREIN (The Ecological and Socio-economical Responses of Global Change on Reindeer Pastures, 2006-2011) and Reindeer Forage and Supplementary Feeding (2008-2010) (http://www.arcticcentre.org/?DeptID=2072). Climate change may have both positive and negative effects on reindeer pastures. The northward and upward movement of the treeline and gradual replacement of lichens with vascular plants associated with increasing temperatures and nutrient availability may change the reindeer pastures in Northern Fennoscandia. The productivity of reindeer forage will most probably increase, but their protein concentrations may decrease due to the dilution effect caused by increased net C production. Our studies showed, that the quality of reindeer forage plants may be reduced due to the UV-B-induced increase in the concentration of phenolics, decreasing forage quality and choice, but reindeer may adapt to increased phenolics. Increased winter precipitation, ground icing and deeper snow cover may reduce the availability and/or quality of reindeer forage, but prolongation of snowless periods might have the opposite effect. Our studies in Finnish Lapland showed a significant relationship between the reindeer calf percentage (calves per 100 females) and the day of snowmelt measured during 1970-2006. The calf percentage was highest when the snowmelt was early, indicating improved availability of forage after the snow disappeared. Ground icing and thicker snow cover may aggravate the digging conditions of reindeer, and increase their energy consumption. Decreased availability of winter forage is related to poor condition of reindeer and lower calving percentages. Consequently, the pressure for supplementary winter feeding of reindeer has increased not only due to reduced area and quality of natural winter pasture land and increased number of reindeer, but also due to more difficult winter conditions. We studied the impacts of reindeer winter feeding on the pinedominated sub-xeric heath forest by using a reindeer field feeding experiment (March – April 2009, 2010). A significant fertilizer effect, expressed as increased N concentration of dwarf shrubs (E. nigrum, V. myrtillus) and growth of wavy hair-grass (D. flexuosa) could be observed already after the first winter. This study showed that besides grazing and trambling, a great deal of the environmental impacts of reindeer winter feeding are caused by uneaten supplementary forage (grass silage, hay) left on the pasture land. 14 FOREST HEALTH MONITORING IN EUROPE ACHIEVEMENTS, PROBLEMS AND PERSPECTIVES Marco Ferretti TerraData environmetrics, spin-off company of the University of Siena, Via L. Bardelloni 19, 58025 Monterotondo M.mo, Grosseto, Italy Contact: [email protected] Much of our current knowledge and understanding of “forest health monitoring” has been developed since the 1980s. It was across the past 30 years that major international monitoring and research programmes have been initiated and developed – and in some cases terminated. It was a sort of learning-by-doing process, which shaped our current vision of the “science of monitoring”. This process took place along a time when also environmental priorities have changed, the easiest example being the shift of concern from acidic deposition to climate change. Today, many questions the value of monitoring programmes in general, and forest monitoring in particular is at risk because of lack of credibility and funds. The two are connected: the lack of credibility (which is rooted in the poor attention given to monitoring design, data quality, and communication) creates a lack of confidence in monitoring results, which in turn results into skepticisms in the whole system and in problems to obtain support. However, it is not possible to deny the achievements of forest monitoring, in terms of results and in terms of improvement in methods and quality. The science basis have been improved and strengthen, and the data generated by the forest monitoring programmes are being, and have been, used to address key environmental issues. The historical data series achieved in Europe have very few competitors in the world. It is therefore somewhat surprising the easiness with which funding agencies at national and international level may decide to quit supporting programmes (i.e., infrastructure, organization, expertize, data) that should better considered as a strategic asset for environmental and forest protection. Forest monitoring is now facing a number of challenges. They involve organization, ability to provide the desired information in due time, scientific defensibility, ability to connect with other strategic networks (e.g. national forest inventories, long-term ecological research sites), and costs. Under the current economical and financial situation, convincing answers to the above challenges are even more important than before. It is therefore essential and urgent to translate such challenges into operational perspectives, and put them into practice. 15 DECLINE AND RECOVERY OF FORESTS IN THE BLACK TRIANGLE AREA OF EUROPE Josef Krecek1, Zuzana Horicka2 and Jana Novakova3 1 Department of Hydrology, Czech Technical University in Prague, Thakurova 7. CZ-166 29 Prague 6, Czech Republic 2 Institute for Environmental Studies, Charles University, 3 Benátská 2, CZ-128 01 Prague 2, Czech Republic Institute of Applied Ecology, Czech Agricultural University, CZ-281 63, Kostelec n.C.l., Czech Republic Contact: [email protected] In the 1970-1980s, forests in the Jizera Mountains (North Bohemia, the Czech Republic) declined as a consequence of the acid atmospheric deposition (namely sulphate from the lignite combustion) and commercial forestry practices (spruce plantations of a low stability, extensive clear-cut, application of heavy mechanisation, non-effective control of insect epidemics, and difficult reforestation). Environmental aspects of those processes included the genesis of catchment run-off, and, particularly, the drop in water quality in water courses and reservoirs. The infiltration capacity of soils decreased from 150 to 40 mm/h, and the network of skid-roads and periodical drainage increased from 1.3 to 4.7 km/km2. The direct (fast) run-off raised from 50 to 70 % of the annual water yield, and soil erosion from 0.01 to 1.34 mm/year. In surface waters, low pH (4-5), high content of toxic metals (namely aluminium, 1-2 mg/l), extinction of fish and drastically reduced zooplankton, phytoplankton and benthic fauna were observed. A recent recovery of surface waters in the Jizera Mts. (an increase in mean annual pH values to 5-6, a drop in aluminium concentrations to 0.2-0.5 mg/l, successful reintroduction of brook char, Salvelinus fontinalis) results from the decreased air pollution (in the 1990s, the deposition of sulphate decreased to ca. 40% in comparison with the year 1987), but also by reduced leaf area of mature spruce plantations, and by liming of both reservoirs and watersheds. The traditional forestry practices (skidding timber by horses or cables, respecting riparian zones, seasonal skidding or manual labour reforestation) have contributed to watershed stabilisation. However, there is a relatively long delay between the drop in the atmospheric load and progress in the biota. Environmental indicators show a delay of almost ten years. The composition of algal mats and fish populations in surface waters take even longer to respond to the environmental changes there. 16 REACTIVE NITROGEN DEPOSITION: TOO MUCH OF A GOOD THING FOR PEATLAND ECOSYSTEMS Lucy J Sheppard CEH Edinburgh, Bush Estate, Penicuik EH26 0QB, Midlothian, Scotland Contact: [email protected] Reactive nitrogen deposition (Nr) will be the third main driver of species change after land use and climate change and it is important to know which ecosystems are most sensitive and why. Likewise since the two forms of Nr have different origins, agriculture (reduced N) versus combustion, energy production and transport (oxidized N), regulatory bodies need to know whether the two forms act similarly or differently in the environment. This is also relevant because the deposition of these two Nr forms differs spatially and temporally. Nr is a fertiliser and has the capacity to transform ecosystems, by changing the competitive abilities of species, but it can also affect many above and below ground processes through its effects on pH, generally causing acidification in plants and soils. Peatlands are an important ecosystem where the effects of eutrophication are likely to be detrimental not just via the plants but also through impacts on biogeochemical cycling and ultimately climate change through effects on greenhouse gas emissions. At CEH Edinburgh we have a unique field Nr manipulation on an ombrotrophic bog in the Scottish Borders, which has been running since 2002 (Leith et al. 2004, Sheppard et al. 2004). The Whim bog experiment addresses not just N deposition/loads but also the form of N: ammonia, ammonium and nitrate. Above and below ground biogeochemical processes are being followed, together with species change in response to treatment, and interactions between the N treatments and abiotic and biotic stress. These results will be discussed in the context of other N manipulation studies on peatlands and peatland vegetation. The largest effects above ground and perhaps not surprisingly below ground were driven by the deposition of ammonia. After nearly 10 years the effects for deposition of N loads approaching the Critical Load for bogs (5-10 kg N ha-1y-1) on keystone species were highly visual: Eriophorum vaginatum is replacing Calluna and the green pigmented form of Sphagnum capillifolium together with the lichen Cladonia portentosa have almost disappeared from the 60 m NH3 free air release transect (Sheppard et al. 2011). By comparison changes in response to wet deposition have been much slower requiring a greater cumulative N load to cause effects, none of which have yet matched the ferocity of the ammonia effects. Many of the below ground changes in soil nutrients show strong coupling with effects above-ground, on the vegetation. Ammonia deposition especially, has the potential to increase emissions of the greenhouse gases methane and nitrous oxide but effects on CO2 are more difficult to evaluate due to the detrimental effects on the most productive species Calluna which has been replaced by the methane conducting Eriophorum spp. Ammonia can be a real issue for peatlands in northern latitudes due to the significant emissions from large mammal colonies. This presentation will highlight areas of concern. References Leith ID, Sheppard LJ, Fowler D et al. (2004). Quantifying dry NH3 deposition to an ombrotrophic bog from an automated NH3 release system. Water, Air, and Soil Pollution: Focus 4, 207-218. Sheppard LJ, Crossley A, Leith ID et al. (2004). An automated wet deposition system to compare the effects of reduced and oxidised N on ombrotrophic bog species: practical considerations. Water, Air, and Soil Pollution: Focus 4, 197–205. Sheppard LJ, Leith ID, Mizunuma T, Cape JN, Crossley A, Leeson S, Sutton MA, Van Dijk N, Fowler D. (2011). Dry deposition of ammonia gas drives species change faster than wet deposition of ammonium ions: evidence from a long-term field manipulation. Global Change Biology 17, 3589-3607. 17 ULTRAVIOLET RADIATION AND BRYOPHYTES Javier Martínez-Abaigar and Encarnación Núñez-Olivera Complejo Científico-Tecnológico, Universidad de La Rioja, Avda. Madre de Dios 51, 26006 Logroño (La Rioja), Spain Contact: [email protected] Ultraviolet radiation (UVR) is an ecological factor that has accompanied life since its origins and whose study has increased greatly since the discovery of the stratospheric ozone reduction. This reduction leads to an increase in UV-B radiation at ground level, and enhanced UV-B may damage photosynthetic organisms. Some habitats frequently dominated by bryophytes may be particularly affected by enhanced UV-B, such as circumpolar bogs and mountain streams above the timberline. In this context, we describe the effects of UVR on bryophytes, on the basis of laboratory and field experiments. In the laboratory, UVR stress may be preferentially indicated by a decrease in the maximum quantum yield of photosystem II (Fv/Fm), chlorophylls/phaeopigments ratios, chlorophyll a/b quotient, and net photosynthesis rates. Thus, the effects of UVR on bryophytes do not seem to be very different to those found in other photosynthetic organisms. However, the responses of bryophytes to UVR are still poorly understood, and thus further study is recommended, taking also into account that these responses depend not only on the species but also on the interaction with environmental factors. The bryophytes studied mostly seem to be UVR-tolerant, in some cases due to the accumulation of protecting UV-absorbing compounds. Liverworts seem to have higher amounts of both constitutive and inducible UV-absorbing compounds than mosses. This difference could support the recent finding that both groups are more phylogenetically distant than previously thought. Given that bryophytes have been used as bioindicators of numerous pollution processes and environmental changes, we propose their use as bioindicators of UVR. In particular, certain individual UV-absorbing compounds of the liverwort Jungermannia exsertifolia subsp. cordifolia may be good biomarkers of the UV-B enhancement due to stratospheric ozone reduction. 18 GROWTH OF PRIMORDIAL SHOOT IN NORWAY SPRUCE BUDS PRECEDING VISIBLE BUD BURST IN RELATION TO ACCUMULATED TEMPERATURE IN THE FIELD Sirkka Sutinen1, Jouni Partanen2, Anneli Viherä-Aarnio3 and Risto Häkkinen3 1 2 Finnish Forest Research Institute, P.O. Box 68, FI-80101 Joensuu, Finland Finnish Forest Research Institute, Finlandiantie 18, FI-58450 Punkaharju, Finland 3 Finnish Forest Research Institute, P.O. Box 18, FI-01301 Vantaa, Finland Contact: [email protected] Timing of bud development in ecodormancy is critical for the trees growing in boreal and temperate regions with seasonally alternating climates. The predicted global warming over the mid and high latitudes of northern continents can make the boreal forests more sensitive to frost damage due to temperature fluctuations in early spring. Phenological models on timing of bud development are needed as a part of simulation studies for predicting the risks entailed by the global climate warming. However, at present these models are based on observations of externally visible bud burst only. In order to get a more precise picture of the timing of bud development before bud burst, we followed the growth of primordial shoot in Norway spruce buds during three consecutive springs in the field. The aim of the study was to analyse, how the primordial shoot length is related to accumulated temperature sums calculated with different starting dates, threshold temperature values and time steps (hour/day). The dependence of microscopic primordial shoot length on accumulated temperature was found to be highly linear. The best coefficients of determination (R2) of different regression models were around 0.95. With the temperature threshold values 0 °C or +1 °C the R2 of all models were close to 0.95, i.e. not depending on the starting time or time step used in temperature accumulation. With lower and higher threshold values R2 varied according to starting date and time step used. Our results from the primordial shoot growth in relation to the temperature sums support the present phenological bud burst models used. Measuring and estimating the externally invisible growth of primordial shoots combined with the tests of bud sensitivity to frost might offer a valuable tool for further studies for predicting the risks of untimely bud development due to climate warming. 19 IMPACT OF GEOCLIMATIC CONDITIONS ON TRITERPENOID CONTENT OF FRUIT AND LEAF CUTICULAR WAXES FROM FINNISH AND POLISH VACCINIUM MYRTILLUS AND VACCINIUM VITIS-IDAEA Anna Szakiel1, Cezary Pączkowski1, Heini Koivuniemi2 and Satu Huttunen2 1 Department of Plant Biochemistry, Faculty of Biology, University of Warsaw, ul. Miecznikowa 1, 02-096 Warszawa, Poland 2 Botany Division, Department of Biology, University of Oulu, P.O. Box 3000, FI-90014 Oulu, Finland Contact: [email protected] Plants of boreal forest understorey are influenced by many abiotic and biotic factors, e.g. light, temperature, growing season length, soil fertility and moisture, winter snow cover, pollution; canopy composition, interactions with herbivores, neighboring plants, microbial pathogens or allies. This complex external impact is regarded to be one of possible reasons of “chemodiversity” among plants of the same species growing in different geoclimates. Such a chemical variability can be particularly expected in relation to plant surface cuticle, which provides the first protective barrier against harmful factors but also forms an interactive flexible interface between the plant and its environment. The aim of this research was to compare the triterpenoid content of cuticular wax layer of berries and leaves of two dwarf shrubs characteristical of forest plant communities of the northern hemisphere: deciduous Vaccinium myrtillus and evergreen Vaccinium vitis-idaea. Plant material was collected in natural forest habitats in Finland (65°066 N; 25°458 E) and Poland (52°455 N, 21°332 E); obtained chloroform-soluble cuticular waxes were fractionated by adsorption chromatography and subjected to GC-MS/FID analysis. Triterpenoid profiles differ markedly between the two plants, with wider range of triterpenes in V. vitis-idaea, and steroids in V. myrtillus. Triterpenoid content of fruit waxes is more abundant in V. vitis-idaea (approx. 70% and 40% of total mass of wax extracts from Finnish and Polish berries) than V. myrtillus (23% in both Finnish and Polish berries), moreover, some differences exist between plants of both origin in composition of triterpene alcohols and ketons. Observed quantitative differences in triterpenoid content of V. vitis-idaea leaf waxes (3-fold higher level of triterpene alcohols and ketones in Finnish plants, 2-fold higher level of triterpene acids in Polish plants) combined with specific seasonal changes (quick accumulation of triterpenoids in waxes of young leaves of Finnish plants during summer, versus their slowler but continuous accumulation in leaf waxes of Polish plants until late autumn) seem to be a consequence of different leaf development pattern evolved in an answer to climatic factors including the longer growth season in Poland and the thicker and longerlasting winter snow cover in Finland. The significance of adaptation to winter conditions is confirmed by striking quantitative similarity between triterpenoid content in cuticular waxes from leaves of Finnish and Polish V. myrtillus plants (about 15% of wax extract mass in leaves of both origin, with almost identical ratio of fractions of triterpene alcohols and ketones, acids, and steroids). Thus, the obtained results imply that the difference of growth strategy between deciduous V. myrtillus and evergreen V. vitis-idaea, as well as variability of geoclimatic and environmental conditions between Finland and Poland, can have an influence on triterpenoid content of fruit and leaf cuticular waxes of investigated plants. Acknowledgement. The study was carried out with the support of the project financing agreement POIG.02.02.00-14-024/08-00. 20 ACCLIMATION OF MOSSES AND TREES UNDER AIR POLLUTION STRESS AND CHANGING CLIMATE Sirkku Manninen Department of Environmental Sciences, University of Helsinki, P.O. Box 56, FI-00014 Helsinki, Finland Contact: [email protected] Acclimatisation or acclimation is a process by which individual plants adjust morphologically, physiologically or biochemically to a gradual change in their environment. Plants can also acclimate against air pollution stress. For example, higher leaf production rate (at the expense of root growth) to compensate acceletared leaf senescence and increased concentration of apoplastic ascorbate have been considered as acclimations to elevated ozone (O3). Many oxidative stress factors occur simultaneously. For example, high temperature and high sunlight radiation are correlated and can induce water stress and the formation of O3. Plants can cope under such conditions with the help of so called cross-resistance, where a defence mechanism to a stress factor is usually capable of inducing resistance to other stressors as well. One way of achieving cross-resistance is to react to a stress with morphological changes that are part of a more general acclimation strategy. However, our understanding of the interactive effects of air pollutants and changing climate in terms of plant acclimation is still poor. Water content and metabolic activity of ectohydric bryophytes such as Sphagnum capillifolium and Pleurozium schreberi depend on the surrounding water conditions. Recent results show that S. capillifolium acclimates to open light-exposed conditions by producing sphagnorubin and robust, thick shoots - usually with more than one capitulum and longer branches than in the green type, which grows under Calluna. The dense clumps are apparently the means by which the open-grown S. capillifolium maximises its biomass per ground surface area, given the important role of the canopy interior branches in water storage (Manninen et al. 2011). Similarly, P. schreberi has a higher biomass per ground surface area in open urban forest stands than in rural forests with higher tree canopy cover (Manninen et al., unpublished). Scots pine (Pinus sylvestris) is a climax species which is adapted to open, dry habitats, while European aspen (Populus tremula) is a pioneer species which thrives in both dry and mesic sites. Experimental results on Scots pine and field results on European aspen suggest increased amounts of epicuticular waxes after exposure to elevated O3 and in an urban environment, respectively (Manninen et al. 2009). In European aspen, the amount of epicuticular waxes showed annual variation and was positively correlated with maximum 1-h O3 concentration and negatively with rainfall (Nikula et al. 2010). Increased amounts of epicuticular waxes is considered as acclimation to increased drought stress under elevated O3 due to impaired stomatal closure. The results will be discussed in terms of moss response to elevated nitrogen (N) deposition and the role of epicuticular waxes in plant response to abiotic and biotic stresses. References Manninen S., Huttunen S., Tømmervik H., Hole L.R., Solberg S. (2009). Northern plants and ozone. Ambio 38, 406-413; Manninen S., Woods C., Leith I.D., Sheppard L.J. (2011). Physiological and morphological effects of long-term ammonium or nitrate deposition on green and red (shade and open grown) Sphagnum capillifolium. Environ. Exp. Bot. 72, 140-148; Nikula, S.,Vapaavuori, E., Manninen, S. (2010). Urbanization-related changes in European aspen (Populus tremula L.): Leaf traits and litter decomposition. Environ. Pollut. 158, 2132-2142. 21 PROSPECTS FOR FUTURE OZONE EFFECTS IN THE NORDIC COUNTRIES WITH CONSIDERATION OF CLIMATE CHANGE H. Pleijel1, J. Klingberg1, J. Uddling1, M Engardt2, P.E. Karlsson3 1 DPES, University of Gothenburg, P.O. Box 461, SE-40530 Göteborg, Sweden 2 SMHI, SE-60176 Norrköping, Sweden 3 Swedish Environmental Research Institute, P.O. Box 5302, SE-40014 Göteborg, Sweden Contact: [email protected] The negative impacts of surface ozone (O3) on vegetation depend on external exposure, leaf gas exchange and plant antioxidant defence capacity, all influenced by climate and CO2 concentrations. In this study the influence of climate change on simulated stomatal O3 uptake of a generic crop and a generic deciduous tree at ten European sites, ranging from North Finland to South Spain, was investigated using the LRTAP Mapping Manual stomatal flux model. O3 concentrations were calculated by a chemistry transport model (MATCH) for three 30-year periods (1961-1990, 20212050, 2071-2100), with constant precursor emissions (year 2000) and meteorology from a regional climate model (RCA3). RCA model runs for the IPCC A2 and B2 scenarios were used. Comparing the AOT40 index (integrating the exceedance of the O3 concentration 40 ppb) with the POD (Phytotoxic Ozone Dose) resulted in very a different geographical distribution of estimated O3 risk for vegetation over Europe. Despite substantially increased modelled future O3 concentrations in central and south Europe, the flux-based risk for O3 damage to vegetation was predicted to remain unchanged or decrease at most sites, mainly as a result of projected reductions in stomatal conductance under rising CO2 concentrations. The CO2 effect is uncertain, especially for trees, but will in principle promote a reduced stomatal uptake of O3 at a certain level of O3. If the CO2 effect will turn out to become small, many areas in Europe will experience an increasing risk for vegetation damage. Drier conditions (higher vapour pressure deficit, lower soil moisture) in southern Europe, limiting stomatal O3 uptake, were also important. At northern latitudes, the current parameterisation of the stomatal conductance model suggest O3 uptake to be mainly limited by temperature. Here, an earlier onset of the growing season may become very important for O3 effects on vegetation in the future. The study demonstrates the importance of accounting for the influences by meteorological variables and rising CO2 on stomatal O3 uptake, and of developing their representation in models used for O3 risk assessment under climate change. A further aspect to consider for O3 exposure in high latitudes is the potential change in the dynamics of O3 occurrence through the year. In northern Fennoscandia there is a more pronounced and early spring peak in O3 compared to the southern parts, while during the summer, when vegetation is most active, there are higher O3 concentrations in the south. The development of O3 concentrations in the north will depend on the large-scale background of O3 (indicated to increase), emissions of O3 precursors in Northern Europe (which have declined) and the influence of climate change. The latter can in principle enhance O3 formation, but may also lead to larger deposition if the duration of the period with snow cover declines. The potential consequences of these changes are discussed. 22 ABSTRACTS POSTERS 23 NITROGEN FIXATION ACTIVITY BY CYANOBACTERIA GROWING ON FOREST MOSSES INCREASES TOWARDS THE NORTH IN FINLAND Maija Salemaa1, Sanna Leppänen2, Raisa Mäkipää1, Aino Smolander1 and Marja Tiirola2 1 Finnish Forest Research Institute, Vantaa Research Unit, P.O. Box 18, FI-01301 Vantaa, Finland 2 Biological and Environmental Science, University of Jyväskylä, P.O. Box 35, FI-40014 University of Jyväskylä, Finland Contact: [email protected] In nitrogen limited boreal ecosystems, biological nitrogen (N2) fixation is an important source of N. We studied biological N2 fixation of cyanobacteria-moss associations on 10 intensively monitored forest ecosystem plots (ICP Forests Level II) along a latitudinal gradient in Finland in 2010. Acetylene reduction (ARA) and stable isotope (15N) methods were compared using the two common forest moss species, Hylocomium splendens, and Pleurozium schreberi. The 15N method was more sensitive than the ARA to find signals of cyanobacterial activity. According to the calibrating equation from a northern spruce plot (Pallasjärvi), 3 moles of reduced acetylene corresponds 1 mole of fixed N2. N2 fixation rate associated to mosses increased towards the north and was at highest 0.6–1 kgN/ha/year (based on the bryophyte biomass in the field). In southern Finland, only few signals of N2 fixation were found. The upper parts of the moss shoots showed 2–3 times higher N2-fixing activity than the lower parts, but there were not big differences between H. splendens and P. schreberi. The moisture level of mosses regulated strongly the rate of N2-fixing activity. The results emphasize that the bryophyte layer has a significant effect on the N input in northern forests. 24 RESPONSE OF REINDEER FORAGE PHENOLICS TO UV RADIATION Minna Turunen1 and Françoise Martz1, 2 1 Arctic Centre, University of Lapland, P.O. Box 122, FI-96101 Rovaniemi, Finland 2 FFRI, Rovaniemi Research Unit, P.O. Box 16, FI-96301 Rovaniemi, Finland Contact: [email protected] Reindeer feed on a wide variety of plants, altogether 200-300 species in Northern Fennoscandia. The availability of different species depends greatly on the season. The aim of the present study was to investigate the effects of long-term UV radiation on the total concentration and chemical composition of methanol-extractable compounds in the leaves of Menyanthes trifoliata L. (buckbean), Eriophorum russeolum Fries ex Hartman (russet cotton grass) and Epilobium angustifolium L. (willow herb), which are important reindeer forage plants in Northern Fennoscandia. We studied these effects by conducting a UV-B exclusion experiment (in oligotrophic flark fen) and two UV-B enhancement experiments (in mesotrophic flark fen and sub-xeric heath forest) in northern Finland (68ºN). In all the studied species, seasonal changes in methanol extractable compounds could be observed. The most important species-specific results were the following: Menyanthes trifoliata: Lower minimum temperatures decreased the flavonol content of M. trifoliata. UV exclusion did not affect the total content of methanol extractable compounds, but it significantly decreased the proportion of flavonols concomitantly with an increase in iridoids. Lower content of flavonols under UV-B exclusion can be explained by lower UV-induced oxidative stress. Iridoids, known for their bitter taste, may have important consequences for forage selection and digestion in reindeer and elk. Eriophorum russeolum: Significant UV effects were detected in E. russeolum leaves in a developmental-specific manner. At the end of the growing season, the proportion of total soluble phenolics was higher in leaves exposed to enhanced UV-A and UV-B radiation than in the control leaves, but the phenolic composition was not significantly modified. Epilobium angustifolium: No UV treatment effect was detected in the content of or composition of E. angustifolium leaf soluble phenolics. These studies showed that the phenolic response of the plants is strongly dependent on the plant species, developmental stage of the plant, the compound in question and the type of the experiment (UV exclusion vs. UV enhancement). Although high phenolic contents are known to deteriorate the quality of forage and decrease the feed intake of ruminants the importance of an increased concentration of phenolics for forage selection and digestion in reindeer is not known. It is likely that reindeer would avoid plant tissues rich in phenolics despite their high nutritive value. Reindeer may, however, be able to balance their diet and minimize the harmful effect of phenolics by favouring mixtures of different plants in the summer when there is a large selection of forage plants and when getting enough highly nutritious nourishment is not a problem. 25 SEASONAL DYNAMICS IN SPHAGNUM CHLOROPHYLL UNDER ALTERED UVB AND TEMPERATURE Anna Hyyryläinen1,3, Minna Turunen2, Pasi Rautio3, Françoise Martz2,3 and Satu Huttunen1 1 Department of Biology, University of Oulu, P.O.Box 3000, FI-90014 Oulu, Finland 2 Arctic Centre, University of Lapland, P.O. Box 122, FI-96101 Rovaniemi, Finland 3 Finnish Forest Research Institute, P.O. Box 16, FI-96301 Rovaniemi, Finland Contact: anna.koivulehto(at)gmail.com Solar UV-B radiation is an important ecological factor regulating the growth, morphology and chemical constituents of numerous plant species. It has been shown to affect each species in a specific way. Little is known about the effects of UV-B on seasonal variation of photosynthetic pigments in different species of Sphagnum mosses. Alteration of other environmental factors, such as temperature, pH, water regime etc., may modulate the impact of solar UV-B radiation on plants. To find out about this impact, three species of peat mosses (Sphagnum balticum, S. jensenii and S. lindbergii) were studied in an open field UV-B attenuation experiment in an oligotrophic tall-sedge Sphagnum flark fen in Northern Finland. Throughout the experiment (2002-2010), the plants were exposed to: 1) attenuated solar UV-B radiation, enhanced temperature (UV-B treatment), 2) ambient solar UV-B radiation, enhanced temperature (control treatment) or 3) ambient solar UV-B radiation and temperature (ambient conditions). The temperature and UV-B were manipulated using plastic filters (polyester for UV-B treatment and cellulose acetate for control plots). Each treatment was replicated ten times. Seasonal and interannual changes in chlorophyll a and b in the peat mosses were monitored. For that, the samples were collected four times in both 2007 (S. lindbergii) and 2008 (all the three species). Solar UV-B radiation, pH of the substrate and the temperature were registered at each plot during the growing seasons. The content of chlorophyll a and b was measured spectrophotometrically. In this poster, we present the preliminary results from the sampling in 2008. Each Sphagnum species expressed individual pattern of seasonal changes in chlorophyll a and b content and ratio, as well as species-specific responses to simulated changes in the environment. The total content of chlorophyll was significantly higher in S. lindbergii and S. jensenii compared to S. balticum. In both S. balticum and S. jensenii, the amount of chlorophyll a decreased by the end of the growing season. In the dominating S. lindbergii it increased in the end of summer in ambient conditions and under UV-B treatment, and decreased in control conditions. In both S. lindbergii and S. jensenii a/b ratio remained stable in all treatments starting the midsummer, but in S. balticum it grew substantially in the end of the season. Treatment effect varied between species and within the growing season. 26 Participants Dr. Jaana Bäck, University of Helsinki, Finland Dr. Marco Ferretti, TerraData environmetrics , University of Siena, Italy M.Sc. Ritva Hiltunen, University of Oulu, Finland Emerita professor Satu Huttunen, University of Oulu, Finland Ph.D. student Anna Hyyryläinen, University of Oulu, Finland Dr. Anne Jokela, University of Oulu, Finland Phil.Lic. Marketta Karhu, City of Oulu, Finland Mr. Antti Kulmala, Ministry of Environment, Finland Professor Pekka Kauppi, University of Helsinki, Finland Professor Riitta Keiski, University of Oulu, Finland Dr. Josef Krecek, Czech Technical University, Prague, Czech Republic Dr. Mikhail Kozlov, University of Turku, Finland Dr. Eero Kubin, Finnish Forest Research Institute, Muhos, Finland Professor Kari Laine, Thule Institute, University of Oulu, Finland Dr. Annamari Markkola, University of Oulu, Finland Dr. Sirkku Manninen, University of Helsinki, Finland Dr. Päivi Merilä, Finnish Forest Research Institute, Rovaniemi, Finland Dr. Javier Martinez- Abaigar, Universidad de La Rioja, Spain Professor Jari Oksanen, University of Oulu, Finland M.Sc. Tuulikki Pakonen, University of Oulu, Finland Professor Håkan Pleijel, University of Gothenburg, Sweden Dr. Jarmo Poikolainen, Finnish Forest Research Institute,, Muhos, Finland Dr. Pasi Rautio, Finnish Forest Research Institute, Rovaniemi, Finland Dr. Anna-Liisa Ruotsalainen, University of Oulu, Finland Professor Seppo Saarela, University of Oulu, Finland Dr. Maija Salemaa, Finnish Forest Research Institute, Vantaa, Finland Dr. Lucy Sheppard, Centre for Ecology and Hydrology , Edinburgh, Scotland Dr. Marja-Liisa Sutinen, Finnish Forest Research Institute , Rovaniemi, Finland Dr. Sirkka Sutinen, Finnish Forest Research Institute , Joensuu, Finland Dr. Anna Szakiel, University of Warsaw, Poland Dr. Kari Taulavuori, University of Oulu, Finland Professor Juha Tuomi, University of Oulu, Finland Dr. Minna Turunen, Arctic Centre, University of Lapland, Finland Phil.Lic. Minna Vanhatalo, University of Oulu, Finland Dr. Piippa Wäli, University of Oulu, Finland 27
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