Anne Kasurinen

Chemical Composition and Decomposition of Silver Birch Leaf Litter Produced under Elevated CO2 and O3

Two field-growing silver birch (Betula pendula Roth) clones (clone 4 and 80) were exposed to elevated CO2 and O3 over three growing seasons (1999–2001). In each year, the nutrients and cell wall chemistry of naturally abscised leaf litter were analyzed in order to determine the possible CO2- and O3-induced changes in the litter quality. Also CO2 and O3 effects on the early leaf litter decomposition dynamics (i.e. decomposition before the lignin decay has started) were studied with litter-bag experiments (Incubation 1 with 1999 leaf litter, Incubation 2 with 2000 leaf litter, and Incubation 3 with 2001 leaf litter) in a nearby silver birch forest. Elevated CO2 decreased N, S, C:P and α-cellulose concentrations, but increased P, hemicellulose and lignin+polyphenolic concentrations, C:N and lignin+polyphenolic:N in both clones. CO2 enrichment decreased the subsequent decomposition of leaves of clone 4 transiently (in Incubations 1 and 2), whereas elevated CO2 effects on the subsequent leaf decomposition of clone 80 were inconsistent. In contrast to CO2, O3 decreased P concentrations and increased C:P, but both of these trends were visible in elevated O3 treatment only. O3-induced decreases in Mn, Zn and B concentrations were observed also, but O3 effects on the cell wall chemistry of leaf litter were minor. Some O3-induced changes either became more consistent in leaf litter collected during 2001 (decrease in B concentrations) or appeared only in this litter lot (decrease in N concentrations, decrease in decomposition at the end of Incubation 3). In conclusion, in northern birch forests elevated CO2 and O3 levels have the potential to affect leaf litter quality, but consistent CO2 and O3 effects on the decomposition process remain to be validated.

Contrasting responses of silver birch VOC emissions to short- and long-term herbivory

There is a need to incorporate the effects of herbivore damage into future models of plant volatile organic compound (VOC) emissions at leaf or canopy levels. Short-term (a few seconds to 48 h) changes in shoot VOC emissions of silver birch (Betula pendula Roth) in response to feeding by geometrid moths (Erannis defoliaria Hübner) were monitored online by proton transfer reaction time-of-flight mass spectrometry (PTR-TOF-MS). In addition, two separate field experiments were established to study the effects of long-term foliage herbivory (FH, 30-32 days of feeding by geometrids Agriopis aurantiaria (Clerck) and E. defoliaria in two consecutive years) and bark herbivory (BH, 21 days of feeding by the pine weevil (Hylobius abietis L.) in the first year) on shoot and rhizosphere VOC emissions of three silver birch genotypes (gt14, gt15 and Hausjärvi provenance). Online monitoring of VOCs emitted from foliage damaged by geometrid larvae showed rapid bursts of green leaf volatiles (GLVs) immediately after feeding activity, whereas terpenoid emissions had a tendency to gradually increase during the monitoring period. Long-term FH caused transient increases in total monoterpene (MT) emissions from gt14 and sesquiterpene (SQT) emissions from Hausjärvi provenance, mainly in the last experimental season. In the BH experiment, genotype effects were detected, with gt14 trees having significantly higher total MT emissions compared with other genotypes. Only MTs were detected in the rhizosphere samples of both field experiments, but their emission rates were unaffected by genotype or herbivory. The results suggest that silver birch shows a rapid VOC emission response to short-term foliage herbivory, whereas the response to long-term foliage herbivory and bark herbivory is less pronounced and variable at different time points.

Rising Atmospheric CO2 Concentration Partially Masks the Negative Effects of Elevated O3 in Silver Birch (Betula pendula Roth)

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

The effect of warming and enhanced ultraviolet radiation on gender-specific emissions of volatile organic compounds from European aspen.

Different environmental stress factors often occur together but their combined effects on plant secondary metabolism are seldom considered. We studied the effect of enhanced ultraviolet (UV-B) (31% increase) radiation and temperature (ambient +2 °C) singly and in combination on gender-specific emissions of volatile organic compounds (VOCs) from 2-year-old clones of European aspen (Populus tremula L.). Plants grew in 36 experimental plots (6 replicates for Control, UV-A, UV-B, T, UV-A+T and UV-B+T treatments), in an experimental field. VOCs emitted from shoots were sampled from two (1 male and 1 female) randomly selected saplings (total of 72 saplings), per plot on two sampling occasions (June and July) in 2014. There was a significant UV-B×temperature interaction effect on emission rates of different VOCs. Isoprene emission rate was increased due to warming, but warming also modified VOC responses to both UV-A and UV-B radiation. Thus, UV-A increased isoprene emissions without warming, whereas UV-B increased emissions only in combination with warming. Warming-modified UV-A and UV-B responses were also seen in monoterpenes (MTs), sesquiterpenes (SQTs) and green leaf volatiles (GLVs). MTs showed also a UV × gender interaction effect as females had higher emission rates under UV-A and UV-B than males. UV × gender and T × gender interactions caused significant differences in VOC blend as there was more variation (more GLVs and trans-β-caryophyllene) in VOCs from female saplings compared to male saplings. VOCs from the rhizosphere were also collected from each plot in two exposure seasons, but no significant treatment effects were observed. Our results suggest that simultaneous warming and elevated-UV-radiation increase the emission of VOCs from aspen. Thus the contribution of combined environmental factors on VOC emissions may have a greater impact to the photochemical reactions in the atmosphere compared to the impact of individual factors acting alone.

The Significance of Ectomycorrhizas in Chemical Quality of Silver Birch Foliage and Above-Ground Insect Herbivore Performance

We tested whether the ectomycorrhizal (ECM) infection level of roots of silver birch (Betula pendula) affects performance of above-ground insect herbivores by increasing available plant biomass, by enhancing availability of nutrients, or by modifying concentration of defense compounds, i.e., phenolics, in birch foliage. Insect performance was determined for a phloem-feeding generalist (Lygus rugulipennis, the European tarnished plant bug), a phloem-feeding specialist (Calaphis flava, the birch aphid), and a chewing generalist (Epirrita autumnata, the autumnal moth larva). Silver birch plantlets had either natural ECM infection level (on average 24% of short roots with ECM), reduced ECM infection level with fungicide (F−, 9% ECM), or enhanced ECM infection level after inoculation with the fungus Paxillus involutus (PI+, 45% ECM) or Leccinum versipelle (LV+, 42% ECM). In general, the most pronounced effect of ECM was observed on growth of plantlets, i.e., stem growth decreased. In PI+-treated plants, leaf biomass also decreased. The effect of mycorrhizal colonization on the host plant’s nitrogen (N) and phosphorous (P) concentration was dependent on the mycorrhizal species and experiment. Fungicide treatment did not cause a consistent decrease in nutrients. Finally, defense of birch against herbivory, expressed as foliar phenolic concentration in plantlets, was not modified by ECM. However, E. autumnata had a significantly higher relative growth rate on PI+ plantlets with high leaf N concentration than on LV+ plantlets with low leaf N concentration. The birch aphid C. flava produced significantly less nymphs on birches with enhanced ECM infection levels (PI+ and LV+ plantlets) than on controls. In summary, our data show that the ECM infection level mainly affects the growth parameters of plantlets, whereas effects on leaf chemical quality are minor. Our data show that effects of ECM infection of birch roots on aboveground herbivores are multifaceted and depend on the fungal species forming ectomycorrhiza and also on the degree of specialization and feeding guild of insects.

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

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

Impact of warming, moderate nitrogen addition and bark herbivory on BVOC emissions and growth of Scots pine (Pinus sylvestris L.) seedlings

The changing climate will expose boreal forests to rising temperatures, increasing soil nitrogen (N) levels and an increasing risk of herbivory. The single and interaction effects of warming (+2 °C increase), moderate N addition (30 kg ha-1 year-1) and bark herbivory by large pine weevil (Hylobius abietis L.) on growth and emissions of biogenic volatile organic compounds (BVOCs) from shoots of Scots pine (Pinus sylvestris L.) seedlings were studied in growth chambers over 175 days. In addition, warming and N addition effects on shoot net photosynthesis (Pn) were measured. Nitrogen addition increased both shoot and root dry weights, whereas warming, in combination with herbivory, reduced stem height growth. Warming together with N addition increased current-year shoot Pn, whereas N effects on previous-year shoot Pn were variable over time. Warming decreased non-oxygenated monoterpene (MT) emissions in June and increased them in July. Of individual MT compounds, α-pinene, δ-3-carene, γ-terpinene and terpinolene were among the most frequently responsive compounds in warming treatments in the May-July period. Sesquiterpene emissions were observed only from warming treatments in July. Moderate N addition increased oxygenated monoterpenes in May, and MTs in June and September. However, N addition effect on MTs in June was clearer without warming than with warming. Bark herbivory tended to increase MT emissions in combination with warming and N addition 3 weeks after the damage caused by weevils. Of individual compounds in other BVOC blends, herbivory increased the emissions of methyl-benzene, benzene and hexanal in July. Hence, though both warming and N addition have a potential to change BVOC emissions from Scots pines, the N effect may also be partly cancelled by warming. Furthermore, herbivory pressure in combination with climate warming and N addition may, at least periodically, increase BVOC release to the atmosphere from young Scots pine seedlings.

Transfer of elements relevant to nuclear fuel cycle from soil to boreal plants and animals in experimental meso- and microcosms

Uranium (U), cobalt (Co), molybdenum (Mo), nickel (Ni), lead (Pb), thorium (Th) and zinc (Zn) occur naturally in soil but their radioactive isotopes can also be released into the environment during the nuclear fuel cycle. The transfer of these elements was studied in three different trophic levels in experimental mesocosms containing downy birch (Betula pubescens), narrow buckler fern (Dryopteris carthusiana) and Scandinavian small-reed (Calamagrostis purpurea ssp. Phragmitoides) as producers, snails (Arianta arbostorum) as herbivores, and earthworms (Lumbricus terrestris) as decomposers. To determine more precisely whether the element uptake of snails is mainly via their food (birch leaves) or both via soil and food, a separate microcosm experiment was also performed. The element uptake of snails did not generally depend on the presence of soil, indicating that the main uptake route was food, except for U, where soil contact was important for uptake when soil U concentration was high. Transfer of elements from soil to plants was not linear, i.e. it was not correctly described by constant concentration ratios (CR) commonly applied in radioecological modeling. Similar nonlinear transfer was found for the invertebrate animals included in this study: elements other than U were taken up more efficiently when element concentration in soil or food was low.

Interactions of ectomycorrhizas and above-ground insect herbivores on silver birch.

Mycorrhizas are mostly beneficial to host plant growth and survival, e.g., due to improved water and nutrient uptake and enhanced pathogen protection, but also a significant amount of host plant carbon is allocated below-ground to support the mycorrhizal growth. These facts and on the other hand the possibility of mycorrhizas to mediate changes in above-ground defensive chemistry may affect performance of above-ground insect herbivores with different feeding guilds. To see the functionality of complex interaction between mycorrhizal status of plants, host plant chemical quality and insect herbivore performance in a wider ecological scale, studies should be conducted in field conditions and factors contributing to global climate change, such as elevated tropospheric ozone (O3), should also be considered. We recently demonstrated in laboratory study, that insect herbivore response to ectomycorrhizal status of birches was dependent on the fungal species forming ectomycorrhiza and the degree of specialization and feeding guild of insects. In this addendum we provide results from the field study where silver birch (Betula pendula Roth.) seedlings were, in addition to mycorrhizal manipulation, exposed to low-level O3 and the host plant growth and chemical quality was analysed as well as the performance of two insect herbivores with different feeding guilds was tested.