Long term effects of ionising radiation in the Chernobyl Exclusion zone on DNA integrity and chemical defence systems of Scots pine (Pinus sylvestris).

The Chernobyl Nuclear Power Plant (ChNPP) accident in 1986 resulted in extremely high levels of acute ionising radiation, that killed or damaged Scots pine (Pinus sylvestris) trees in the surrounding areas. Dead trees were cleared and buried, and new plantations established a few years later. Today, more than three decades later, gamma and beta-radiation near the ChNPP is still elevated compared with ambient levels but have decreased by a factor of 300 and 100, respectively. In the present work, Scots pine-trees growing at High (220 µGy h-1), Medium (11 µGy h-1), and Low (0.2 µGy h-1) total (internal + external) dose rates of chronically elevated ionising radiation in the Chernobyl Exclusion zone were investigated with respect to possible damage to DNA, cells and organelles, as well as potentially increased levels of phenolic and terpenoid antioxidants. Scots pine from the High and Medium radiation sites had elevated levels of DNA damage in shoot tips and needles as shown by the COMET assay, as well as increased numbers of resin ducts and subcellular abnormalities in needles. Needles from the High radiation site showed elevated levels of monoterpenes and condensed tannins compared with those from the other sites. In conclusion, more than three decades after the ChNPP accident substantial DNA damage and (sub)cellular effects, but also mobilisation of stress-protective substances possessing antioxidant activity were observed in Scots pine trees growing at elevated levels of ionising radiation. This demonstrates that the radiation levels in the Red Forest still significantly impact the plant community.

Legacies of invertebrate exclusion and tree secondary metabolites control fungal communities in dead wood.

During decomposition of organic matter, microbial communities may follow different successional trajectories depending on the initial environment and colonizers. The timing and order of the species arrival (assembly history) can lead to divergent communities through priority effects. We explored how assembly history and resource quality affected fungal communities and decay rate of decomposing wood, 1.5 and 4.5 years after tree felling. Additionally, we investigated the effect of invertebrate exclusion during the first two summers. We measured initial resource quality of bark and wood of aspen (Populus tremula) logs and surveyed the fungal communities by DNA metabarcoding at different times during succession. We found that gradients in fungal community composition were related to resource quality and we discuss how this may reflect different fungal life history strategies. As with previous studies, the initial amount of bark tannins was negatively correlated with wood decomposition rate over 4.5 years. The initial fungal community explained variation in community composition after 1.5, but not 4.5, years of succession. Although the assembly history of initial colonizers may cause alternative trajectories in successional communities, our results indicate that the communities may converge with the arrival of secondary colonizers. We also identified a strong legacy of invertebrate exclusion on fungal communities, even after 4.5 years of succession, thereby adding crucial knowledge on the importance of invertebrates in affecting fungal community development. By measuring and manipulating aspects of assembly history and resource quality that have rarely been studied, we expand our understanding of the complexity of fungal community dynamics.

Fungal community dynamics across a forest-alpine ecotone.

Climate change is causing upward shift of forest lines worldwide, with consequences for soil biota and carbon (C) sequestration. We here analyse compositional changes in the soil biota across the forest line ecotone, an important transition zone between different ecosystems. We collected soil samples along transects stretching from subalpine mountain birch forests to alpine heath. Soil fungi and micro-eukaryotes were surveyed using DNA metabarcoding of the ITS2 and 18S markers, while ergosterol was used to quantify fungal biomass. We observed a strong shift in the soil biota across the forest line ecotone: Below the forest line, there were higher proportions of basidiomycetes and mucoromycetes, including ectomycorrhizal and saprotrophic fungi. Above it, we observed relatively more root-associated ascomycetes, including Archaeorhizomycetes, ericoid mycorrhizal fungi and dark septate endophytes. Ergosterol and percentage C content in soil correlated strongly and positively with the abundance of root-associated ascomycetes. The predominance of ectomycorrhizal and saprotrophic fungi below the forest line probably promote high C turnover, while root-associated ascomycetes above the forest line may enhance C sequestration. With further rise in forest lines, there will be a corresponding shift in the below-ground biota, probably leading to enhanced release of soil C.

Establishment of spruce plantations in native birch forests reduces soil fungal diversity.

Plantations of Norway spruce have been established well beyond its natural range in many parts of the world, potentially impacting native microbial ecosystems and the processes they mediate. In this study, we investigate how the establishment of spruce plantations in a landscape dominated by native birch forests in western Norway impacts soil properties and belowground fungal communities. Soil cores were collected from neighboring stands of planted spruce and native birch forests. We used DNA metabarcoding of the rDNA internal transcribed spacer 2 region and ergosterol measurements to survey the fungal community composition and its biomass, respectively. In the two investigated soil layers (litter and humus), fungal community composition, diversity and biomass were strongly affected by the tree species shift. Native birch stands hosted markedly richer fungal communities, including numerous fungi not present in planted spruce stands. In contrast, the spruce stands included higher relative abundance of ectomycorrhizal fungi as well as higher fungal biomass. Hence, establishing plantations of Norway spruce in native birch forests leads to significant losses in diversity, but increase in biomass of ectomycorrhizal fungi, which could potentially impact carbon sequestration processes and ecosystem functioning.

Accumulation of phenolics and growth of dioecious Populus tremula (L.) seedlings over three growing seasons under elevated temperature and UVB radiation.

Accumulation of secondary metabolites may exhibit developmentally regulated variation in different plant organs. Moreover, prevailing environmental conditions may interact with development-related variations in plant traits. In this study, we examined developmentally regulated variation in phenolic accumulation in the twigs of dioecious Populus tremula (L.) and how the effects of elevated temperature and ultraviolet B (UVB) radiation on growth and phenolics accumulation varied as the plants get older. In an open-field experiment, six female and six male genotypes were exposed to single and combined elevated temperature and UVB radiation treatments for three consecutive growing seasons. The concentrations of low molecular weight phenolics and condensed tannins did not show age-dependent variation in the twigs. In temperature-treated plants, diameter growth rate decreased, and concentration of condensed tannins increased as plants aged; there were no cumulative effects of elevated UVB radiation on growth and phenolic accumulation. Females maintained a higher concentration of low molecular weight phenolics throughout the experimental period; however, growth and phenolic concentration did not vary over time in females and males. Our results suggest that phenolic accumulation in perennial plants may not necessarily always exhibit age-dependent variation and the effects of elevated temperature on growth and phenolic may diminish as plants get older.

Responses in growth and phenolics accumulation to lateral bud removal in male and female saplings of Populus tremula (L.) under simulated climate change.

The responses in growth and defense after tissue damage are highly variable in plants depending on species, damaged-tissue type and the intensity of damage. The prevailing abiotic conditions can also influence these responses. In this study, our aim was to examine how the removal of lateral vegetative buds affects the growth and accumulation of phenolics in saplings of the dioecious Populus tremula grown under simulated climate change. For three growing seasons, the saplings were grown under ambient conditions (control), elevated temperature (+2°C) and elevated UV radiation (30%) (UVB and UVA as its control), or a combination of these. In the fourth growing season, all saplings were grown under ambient conditions. The bud removal was performed twice - in summer and autumn - in the third year. The responses of growth and the accumulation of phenolics to the bud removal were measured at the end of the fourth growing season. Removal of 5% of the lateral buds resulted in higher leaf, stem and total plant biomass in both sexes of P. tremula saplings, compared to intact plants. The effects were greater in the temperature-treated plants, especially in the temperature-treated females. The concentrations of flavonoids and condensed tannins were higher in the bud-removed individuals. The concentration of condensed tannins was also higher in the males than in the females, opposite to the concentration of phenolic acids. There was no significant interaction between bud removal and UVB treatment on either growth or phenolics. Our results suggest that plants can allocate resources to both growth and defense simultaneously in response to tissue loss, and that global warming can modify the responses to some extent.

Ultraviolet radiation accelerates photodegradation under controlled conditions but slows the decomposition of senescent leaves from forest stands in southern Finland.

Depending on the environment, sunlight can positively or negatively affect litter decomposition, through the ensemble of direct and indirect processes constituting photodegradation. Which of these processes predominate depends on the ecosystem studied and on the spectral composition of sunlight received. To examine the relevance of photodegradation for litter decomposition in forest understoreys, we filtered ultraviolet radiation (UV) and blue light from leaves of Fagus sylvatica and Betula pendula at two different stages of senescence in both a controlled-environment experiment and outdoors in four different forest stands (Picea abies, Fagus sylvatica, Acer platanoides, Betula pendula). Controlling for leaf orientation and initial differences in leaf chlorophyll and flavonol concentrations; we measured mass loss at the end of each experiment and characterised the phenolic profile of the leaf litter following photodegradation. In most forest stands, less mass was lost from decomposing leaves that received solar UV radiation compared with those under UV-attenuating filters, while in the controlled environment UV-A radiation either slightly accelerated or had no significant effect on photodegradation, according to species identity. Only a few individual phenolic compounds were affected by our different filter treatments, but photodegradation did affect the phenolic profile. We can conclude that photodegradation has a small stand- and species-specific effect on the decomposition of surface leaf litter in forest understoreys during the winter following leaf fall in southern Finland. Photodegradation was wavelength-dependent and modulated by the canopy species filtering sunlight and likely creating different combinations of spectral composition, moisture, temperature and snowpack characteristics.

No evidence of a protective or cumulative negative effect of UV-B on growth inhibition induced by gamma radiation in Scots pine (Pinus sylvestris) seedlings.

Exposure to ambient UV-B radiation may prime protective responses towards various stressors in plants, though information about interactive effects of UV-B and gamma radiation is scarce. Here, we aimed to test whether UV-B exposure could prime acclimatisation mechanisms contributing to tolerance to low-moderate gamma radiation levels in Scots pine seedlings, and concurrently whether simultaneous UV-B and gamma exposure may have an additive adverse effect on seedlings that had previously not encountered either of these stressors. Responses to simultaneous UV-B (0.35 W m-2) and gamma radiation (10.2-125 mGy h-1) for 6 days with or without UV-B pre-exposure (0.35 W m-2, 4 days) were studied across various levels of organisation, as compared to effects of either radiation type. In contrast to UV-B, and regardless of UV-B presence, gamma radiation at ≥42.9 mGy h-1 caused increased formation of reactive oxygen species and reduced shoot length, and reduced root length at 125 mGy h-1. In all experiments there was a gamma dose rate-dependent increase in DNA damage at ≥10.8 mGy h-1, generally with additional UV-B-induced damage. Gamma-induced growth inhibition and gamma- and UV-B-induced DNA damage were still visible 44 days post-irradiation, even at 20.7 mGy h-1, probably due to genomic instability, but this was reversed after 8 months. In conclusion, there was no evidence of a protective effect of UV-B on gamma-induced growth inhibition and DNA damage in Scots pine, and no additive adverse effect of gamma and UV-B radiation on growth in spite of the additional UV-B-induced DNA damage.

The influence of spectral composition on spring and autumn phenology in trees.

Several recent reviews highlight the molecular mechanisms that underpin phenological responses to temperature and photoperiod; however, these have mostly overlooked the influence of solar radiation and its spectral composition on these processes. For instance, solar radiation in the blue and ultraviolet (UV) regions of the spectrum, as well as the red/far-red (R:FR) ratio, can influence spring and autumn phenology. Solar radiation reaching the Earth changes diurnally and seasonally; however, rising global temperatures, latitudinal range shifts and light pollution are likely to produce novel combinations of phenological cues for tree species. Here, we review the literature on phenological responses to spectral composition. Our objective was to explore the natural variation in spectral composition using radiative transfer models and to reveal any species-specific or ecotype-specific responses relating to latitudinal origin. These responses are likely to be most pronounced at high latitudes where spectral composition varies most throughout the year. For instance, trees from high latitudes tend to be more sensitive to changes in R:FR than those from low latitudes. The effects of blue light and UV radiation on phenology have not been studied as much as those of R:FR, but the limited results available suggest both could be candidate cues affecting autumn leaf colouration and senescence. Failure of more-southern species and ecotypes to adapt and use spectral cues during northwards range shifts could result in mistimed phenology, potentially resulting in frost damage, reduced fitness and limited range expansion. Future areas for research should look to establish how consistently different functional types of tree respond to spectral cues and identify photoreceptor-mediated mechanisms that allow plants to combine information from multiple light cues to coordinate the timing of phenological events. It should then be feasible to consider the synchronous or sequential action of light cues within a hierarchy of environmental factors regulating phenology.

Elevated air humidity increases UV mediated leaf and DNA damage in pea (Pisum sativum) due to reduced flavonoid content and antioxidant power.

Growth in high relative air humidity (RH, >85%) affects plant morphology and causes diminished response to stomatal closing signals. Many greenhouses are prone to high RH conditions, which may negatively affect production and post-harvest quality. UV radiation induces stomatal closure in several species, and facilitates disease control. We hypothesised that UV exposure may trigger stomatal closure in pea plants (Pisum sativum) grown in high RH, thereby restoring stomatal function. The effects of UV exposure were tested on plants grown in moderate (60%) or high (90%) RH. UV exposure occurred at night, according to a disease control protocol. Lower stomatal conductance rates were found in UV-exposed plants, though UV exposure did not improve the rate of response to closing stimuli or desiccation tolerance. UV-exposed plants showed leaf curling, chlorosis, necrosis, and DNA damage measured by the presence of cyclobutane pyrimidine dimers (CPD), all of which were significantly greater in high RH plants. These plants also had lower total flavonoid content than moderate RH plants, and UV-exposed plants had less than controls. Plants exposed to UV had a higher content of cuticular layer uronic compounds than control plants. However, high RH plants had a higher relative amount of cuticular waxes, but decreased proteins and uronic compounds. Plants grown in high RH had reduced foliar antioxidant power compared to moderate RH. These results indicate that high RH plants were more susceptible to UV-induced damage than moderate RH plants due to reduced flavonoid content and oxidative stress defence.

Spruce and beech as local determinants of forest fungal community structure in litter, humus and mineral soil.

Beech forests reaches its native distribution limit in SE Norway, but is expected to expand substantially northwards due to climate warming. This may potentially result in a fundamental transformation of contemporary Northern European forests, with tentative effects on the associated belowground fungi. Fungal communities mediate vital ecosystem processes such as ecosystem productivity and carbon sequestration in boreal forests. To investigate how soil fungi is affected by the vegetation transition from spruce to beech forest, we sampled litter, humus and mineral soil in a forest landscape dominated by beech, spruce or a mixture of these. The fungal communities in the soil samples were analyzed by DNA metabarcoding of the rDNA ITS2 region. Although soil layers were the most important structuring gradient, we found clear differences in fungal species composition between spruce and beech plots. The differences in fungal community composition were most evident in the litter and least in the mineral soil. Decomposers, most notably Mycena, dominated the litter layer while various mycorrhizal fungi dominated the humus and mineral layers. Some ectomycorrhizal taxa, such as Cenoccocum and Russula, were more abundant in spruce forests. Differences in fungal community composition between forest types can potentially have large impacts on carbon sequestration rates.

Fertilization Changes Chemical Defense in Needles of Mature Norway Spruce (Picea abies).

Nitrogen availability limits growth in most boreal forests. However, parts of the boreal zone receive significant levels of nitrogen deposition. At the same time, forests are fertilized to increase volume growth and carbon sequestration. No matter the source, increasing nitrogen in the boreal forest ecosystem will influence the resource situation for its primary producers, the plants, with possible implications for their defensive chemistry. In general, fertilization reduces phenolic compound concentrations in trees, but existing evidence mainly comes from studies on young plants. Given the role of the phenolic compounds in protection against herbivores and other forest pests, it is important to know if phenolics are reduced with fertilization also in mature trees. The evergreen Norway spruce is long-lived, and it is reasonable that defensive strategies could change from the juvenile to the reproductive and mature phases. In addition, as the needles are kept for several years, defense could also change with needle age. We sampled current and previous year needles from an N fertilization experiment in a Norway spruce forest landscape in south-central Norway to which N had been added annually for 13 years. We analyzed total nitrogen (N) and carbon (C), as well as low-molecular phenolics and condensed tannins. Needles from fertilized trees had higher N than those from controls plots, and fertilization decreased concentrations of many flavonoids, as well as condensed tannins in current year needles. In previous year needles, some stilbenes and condensed tannins were higher in fertilized trees. In control trees, the total phenolic concentration was almost five times as high in previous year needles compared with those from the current year, and there were great compositional differences. Previous year needles contained highest concentrations of acetophenone and stilbenes, while in the current year needles the flavonoids, and especially coumaroyl-astragalins dominated. Condensed tannins did not differ between current and previous year needles from control trees. In conclusion, the phenolic defense of current year needles of mature P.abies trees was strongly changed upon fertilization. This may imply that nitrogen deposition and forest fertilization leave forests less robust in a time when pests may take advantages of a changing climate.

Elevated temperature and CO2 affect responses of European aspen (Populus tremula) to soil pyrene contamination.

High northern latitudes are climatic sensitive areas, and are also regions to which polycyclic aromatic hydrocarbons (PAHs) easily transport and accumulate with potential risk to natural ecosystems. However, the effect of PAHs on northern woody plant growth and defense under climate change is very little studied. Here, we conducted a unique experiment in greenhouses to investigate sex-related responses of the dioecious Populus tremula to pyrene (50mgkg-1) and residue of pyrene in soils under ambient and elevated temperature (+1.8°C on average) and CO2 (740ppm). Pyrene decreased stem biomass and leaf area by 9% and 6%, respectively under ambient conditions, and the reduction of leaf area was more severe under elevated temperature (38%), elevated CO2 (37%), and combined T+CO2 (42%). Other growth parameters were unchanged by pyrene. Pyrene did not affect the concentration of leaf total phenolics under ambient conditions, but increased it by 16%, 1%, and 20% compared to controls under elevated temperature, elevated CO2, and T+CO2, respectively. Pyrene had only minor sex-specific effects on plant growth and phenolics. The concentration of residual pyrene in pyrene-spiked soils was higher under elevated CO2 than under ambient, elevated temperature, and combined T+CO2. The results suggest that both sexes of P. tremula have the capacity to regulate growth and metabolism to adjust to the stress of the tested pyrene contamination under elevated temperature and CO2, but potential risk of pyrene to plants still exists in the future changing climate.

A Seven-Year Study of Phenolic Concentrations of the Dioecious Salix myrsinifolia.

In boreal woody plants, concentrations of defensive phenolic compounds are expected to be at a high level during the juvenile phase and decrease in maturity, although there is variation between plant species. Females of dioecious species, like most of the Salicaceae, are expected to invest their resources in defense and reproduction, while males are expected to be more growth-oriented. We studied age- and sex-dependent changes in leaf and stem phenolics, and in height and diameter growth in a dioecious Salix myrsinifolia plants over a seven-year time period. In addition, we registered flowering as well as rust damage in the leaves. From the first year and throughout ontogenetic development from juvenile to adult phases, there was no significant change in the concentrations of any of the studied compounds in the leaves of S. myrsinifolia. In the stems, the concentrations of six out of 43 identified compounds decreased slightly with age, which may be partly explained by dilution caused by the increment in stem diameter with age. The fairly steady chemistry level over seven years, accompanied by moderate genotypic phenolic variation, indicates important roles of chemical defenses against herbivory for this early-successional species.

Sex-related responses of European aspen (Populus tremula L.) to combined stress: TiO2 nanoparticles, elevated temperature and CO2 concentration.

The combined effects of climate change and chemical contaminants on plant performance are still not well understood. Especially, whether different sexes of dioecious plants respond differently to combined stresses is unknown. In order to study the sex-related responses of European aspen to soil nTiO2 contamination (0, 50, 300 mg kg-1) under elevated temperature (+1.6 °C) and CO2 (730 ppm), we conducted a study in greenhouses. Ti accumulated in roots exposed to nTiO2 (1.1-3.3 and 2.7-21.1 mg kg-1 in 50 and 300 mg kg-1 treatments, respectively). Elevated CO2 had no effects on Ti uptake, while elevated temperature increased it in the 300 mg kg-1 treatment. Males grew taller than females under ambient conditions, but females had greater height and biomass increment under elevated temperature. In all climate treatments, nTiO2 increased leaf phenolics in females by 12-19% and 15-26% at 50 and 300 mg kg-1, respectively. Leaf phenolics decreased under elevated temperature, but increased under elevated CO2 in both sexes. Results suggest that females have better chemical defense against nTiO2 than males under future climate conditions. In the longer run, this may cause changes in the competitive abilities of both sexes, which again may affect sex ratios and genetic variation in nature.

Effects of long-term UV-exposure and plant sex on the leaf phenoloxidase activities and phenolic concentrations of Salix myrsinifolia (Salisb.).

The accumulation of flavonoids on the leaf surface is a well-characterized protective mechanism against UV-B radiation. Other protective mechanisms, such as the induction of antioxidative enzymes and peroxidase-mediated lignification may also be important. The effects of UV-B radiation have mainly been considered in short-term studies, whereas ecologically more relevant long-term field studies are still rare. Here we examined the effects of long-term exposure to enhanced UV-B radiation on the activities of two antioxidative enzymes, polyphenol oxidase (PPO; EC 1.10.2.2 and EC 1.14.18.1) and guaiacol peroxidase (POD; EC 1.11.1.7), as well as the phenolic concentrations in two sexes of the dioecious species, Salix myrsinifolia. After three consecutive growth seasons with enhanced UV-B radiation, we found that PPO activity was decreased by UV radiation in male plants, which might explain their lower UV-B tolerance when compared to female plants. In addition, male plants had higher specific activity than did female plants under ambient conditions, supporting the idea that males of S. myrsinifolia are generally more growth-oriented than females. By contrast, neither UV treatment nor sex had significant effects on the POD activities of willows. Gender differences in the concentrations of phenolic compounds are in line with the general concept that males are less well defended than females. We suggest that the inability to increase PPO and POD activity, along with lower accumulation of UV-B absorbing compounds under UV-B exposure, might be one of the reasons why males had thinner leaves and were less tolerant of UV-B than were females.

Effects of soil pyrene contamination on growth and phenolics in Norway spruce (Picea abies) are modified by elevated temperature and CO2.

With the constant accumulation of polycyclic aromatic hydrocarbons (PAHs) in soil and increasing temperature and CO2 levels, plants will inevitably be exposed to combined stress. Studies on the effects of such combined stresses are needed to develop mitigation and adaptation measures. Here, we investigated the effects of soil pyrene contamination (50 mg kg-1) on growth and phenolics of 1-year-old Norway spruce seedlings from five different origins in Finland at elevated temperature (+ 2 °C) and CO2 (+ 360 ppm). Pyrene significantly decreased spruce height growth (0-48%), needle biomass (0-44%), stem biomass (0-43%), and total phenolic concentrations in needles (2-13%) and stems (1-19%) compared to control plants. Elevated temperature alone did not affect growth but led to lower concentrations of total phenolics in needles (5-29%) and stems (5-18%) in both soil treatments. By contrast, elevated CO2 led to higher needle biomass (0-39%) in pyrene-spiked soils and higher concentrations of stem phenolics (0-18%) in pyrene-spiked and control soils compared to ambient treatments. The decrease in height growth and phenolic concentrations caused by pyrene was greater at elevated temperature, while elevated CO2 only marginally modified the response. Seedlings from different origins showed different responses to the combined environmental stressors. The changes in growth and in the quantity and quality of phenolics in this study suggest that future climate changes will aggravate the negative influence of soil pyrene pollution on northern conifer forest ecosystems.

Moose selecting for specific nutritional composition of birch places limits on food acceptability.

Despite decades of intense research, it remains largely unsolved which nutritional factors underpin food selection by large herbivores in the wild. We measured nutritional composition of birch foliage (Betula pubescens) available to, and used by, moose (Alces alces) in natural settings in two neighboring regions with contrasting animal body mass. This readily available food source is a staple food item in the diet of moose in the high-fitness region, but apparently underutilized by moose in the low-fitness region. Available birch foliage in the two regions had similar concentrations of macronutrients (crude protein [CP], fiber fractions, and water-soluble carbohydrates [WSC]), although a notably lower variation of WSC in the low-fitness region. For minerals, there were several area differences: available birch foliage in the low-fitness region had less Mg (depending on year) and P, but more Ca, Zn, Cu, and Mn. It also had higher concentrations of some plant secondary metabolites: chlorogenic acids, quercetins, and especially MeOH-soluble condensed tannins. Despite the area differences in available foliage, we found the same nutritional composition of birch foliage used in the two regions. Compared to available birch foliage, moose consistently used birch foliage with more CP, more structural fiber (mainly hemicellulose), less WSC, higher concentrations of several minerals (Ca, Zn, K, Mn, Cu), and lower concentrations of some secondary metabolites (most importantly, MeOH-soluble condensed tannins). Our study conceptually supports the nutrient-balancing hypothesis for a large herbivore: within a given temporal frame, moose select for plant material that matches a specific nutritional composition. As our data illustrate, different moose populations may select for the same composition even when the nutritional composition available in a given food source varies between their living areas. Such fastidiousness limits the proportion of available food that is acceptable to the animal and has bearings on our understanding and application of the concept of carrying capacity.

Effect of climate change on bud phenology of young aspen plants (Populus tremula. L).

Boreal tree species are excellent tools for studying tolerance to climate change. Bud phenology is a trait, which is highly sensitive to environmental fluctuations and thus useful for climate change investigations. However, experimental studies of bud phenology under simulated climate change outdoors are deficient. We conducted a multifactorial field experiment with single (T, UVA, UVB) and combined treatments (UVA+T, UVB+T) of elevated temperature (T, +2°C) and ultraviolet-B radiation (+30% UVB) in order to examine their impact on both male and female genotypes of aspen (Populus tremula L.). This study focuses on the effect of the treatments in years 2 and 3 after planting (2013, 2014) and follows how bud phenology is adapting in year 4 (2015), when the treatments were discontinued. Moreover, the effect of bud removal was recorded. We found that elevated temperature played a key role in delaying bud set and forcing bud break in intact individuals, as well as slightly delaying bud break in bud-removed individuals. UVB delayed the bud break in bud-removed males. In addition, both UVA and UVB interacted with temperature in year 3 and even in year 4, when the treatments were off, but only in male individuals. Axillary bud removal forced both bud break and bud set under combined treatments (UVA+T, UVB+T) and delayed both under individual treatments (T, UVB). In conclusion, male aspens were more responsive to the treatments than females and that effect of elevated temperature and UV radiation on bud set and bud break of aspen is not disappearing over 4-year study period.

Correction to: Compositional Changes in Foliage Phenolics with Plant Age, a Natural Experiment in Boreal Forests.

The article Compositional Changes in Foliage Phenolics with Plant Age, a Natural Experiment in Boreal Forests, written by Hilde Karine Wam, Caroline Stolter and Line Nybakken, was originally published electronically on the publisher's internet portal (currently SpringerLink) on August 29, 2017 without open access.