Acclimation of subarctic vegetation to warming and increased cloudiness.

Subarctic ecosystems are exposed to elevated temperatures and increased cloudiness in a changing climate with potentially important effects on vegetation structure, composition, and ecosystem functioning. We investigated the individual and combined effects of warming and increased cloudiness on vegetation greenness and cover in mesocosms from two tundra and one palsa mire ecosystems kept under strict environmental control in climate chambers. We also investigated leaf anatomical and biochemical traits of four dominant vascular plant species (Empetrum hermaphroditum, Vaccinium myrtillus, Vaccinium vitis-idaea, and Rubus chamaemorus). Vegetation greenness increased in response to warming in all sites and in response to increased cloudiness in the tundra sites but without associated increases in vegetation cover or biomass, except that E. hermaphroditum biomass increased under warming. The combined warming and increased cloudiness treatment had an additive effect on vegetation greenness in all sites. It also increased the cover of graminoids and forbs in one of the tundra sites. Warming increased leaf dry mass per area of V. myrtillus and R. chamaemorus, and glandular trichome density of V. myrtillus and decreased spongy intercellular space of E. hermaphroditum and V. vitis-idaea. Increased cloudiness decreased leaf dry mass per area of V. myrtillus, palisade thickness of E. hermaphroditum, and stomata density of E. hermaphroditum and V. vitis-idaea, and increased leaf area and epidermis thickness of V. myrtillus, leaf shape index and nitrogen of E. hermaphroditum, and palisade intercellular space of V. vitis-idaea. The combined treatment caused thinner leaves and decreased leaf carbon for V. myrtillus, and increased leaf chlorophyll of E. hermaphroditum. We show that under future warmer increased cloudiness conditions in the Subarctic (as simulated in our experiment), vegetation composition and distribution will change, mostly dominated by graminoids and forbs. These changes will depend on the responses of leaf anatomical and biochemical traits and will likely impact carbon gain and primary productivity and abiotic and biotic stress tolerance.

Volatile-mediated between-plant communication in Scots pine and the effects of elevated ozone.

Conifers are dominant tree species in boreal forests, but are susceptible to attack by bark beetles. Upon bark beetle attack, conifers release substantial quantities of volatile organic compounds known as herbivore-induced plant volatiles (HIPVs). Earlier studies of broadleaved plants have shown that HIPVs provide information to neighbouring plants, which may enhance their defences. However, the defence responses of HIPV-receiver plants have not been described for conifers. Here we advance knowledge of plant-plant communication in conifers by documenting a suite of receiver-plant responses to bark-feeding-induced volatiles. Scots pine seedlings exposed to HIPVs were more resistant to subsequent weevil feeding and received less damage. Receiver plants had both induced and primed volatile emissions and their resin ducts had an increased epithelial cell (EC) mean area and an increased number of cells located in the second EC layer. Importantly, HIPV exposure increased stomatal conductance and net photosynthesis rate of receiver plants. Receiver-plant responses were also examined under elevated ozone conditions and found to be significantly altered. However, the final defence outcome was not affected. These findings demonstrate that HIPVs modulate conifer metabolism through responses spanning photosynthesis and chemical defence. The responses are adjusted under ozone stress, but the defence benefits remain intact.

Analysis of Transcriptome and Terpene Constituents of Scots Pine Genotypes Inherently Resistant or Susceptible to Heterobasidion annosum.

Root and stem rot caused by Heterobasidion annosum is a severe problem in boreal Scots pine. Dissecting the features of disease resistance is generally an essential step in resistance breeding in plants and forest trees. In this study, we explored inherent resistance factors of Scots pine against H. annosum. A total of 236 families consisting of 85 full-sib (FS), 35 half-sib population mix (HSpm), and 116 half-sib (HS) families of Scots pine seedlings were inoculated with a H. annosum isolate. We sampled needle tissues before inoculation for terpene measurements and RNA sequencing. Based on the lesion area, the extremes of 12 resistant and 12 susceptible families were selected for further analyses. Necrotic lesions resulting from fungal infection were in a weak to moderate relationship with the plant height. Monoterpenes were the principal terpene compounds observed in Scots pine seedlings. Concentrations of 3-carene were significantly higher in pine genotypes inherently resistant compared with susceptible seedlings. By contrast, susceptible genotypes had significantly higher proportions of α-pinene. Gene ontology analysis of differential expressed transcripts (DETs) revealed that response to biotic factors was enriched in resistant seedlings. Functional characterization of individual DETs revealed that higher expression of transcripts involved in response to abiotic stress was common in susceptible genotypes. This observation was supported by the annotation of hub genes in a key module that was significantly correlated with the lesion trait through weighted gene co-expression network analysis (WGCNA) of 16 HS and HSpm samples. These findings contribute to our understanding of constitutive resistance factors of Scots pine against Heterobasidion root and stem rot diseases.

Separating the effects of air and soil temperature on silver birch. Part II. The relation of physiology and leaf anatomy to growth dynamics.

The aboveground parts of boreal forest trees grow earlier in the growing season, the roots mostly later. The idea was to examine whether root growth followed soil temperature, or whether shoot growth also demanded most resources in the early growing season (soil temperature vs internal sink strengths for resources). The linkage between air and soil temperature was broken by switching the soil temperature. We aimed here to identify the direct effects of different soil temperature patterns on physiology, leaf anatomy and their interactions, and how they relate to the control of the growth dynamics of silver birch (Betula pendula Roth). Sixteen 2-year-old seedlings were grown in a controlled environment for two 14-week simulated growing seasons (GS1, GS2). An 8-week dormancy period interposed the GSs. In GS2, soil temperature treatments were applied: constant 10 °C (Cool), constant 18 °C (Warm), early growing season at 10 °C switched to 18 °C later (Early Cool Late Warm) and 18 °C followed by 10 °C (Early Warm Late Cool) were applied during GS2. The switch from cool to warm enhanced the water status, net photosynthesis, chlorophyll content index, effective yield of photosystem II (ΔF/Fm') and leaf expansion of the seedlings. Warm treatment increased the stomatal number per leaf. In contrast, soil cooling increased glandular trichomes. This investment in increasing the chemical defense potential may be associated with the decreased growth in cool soil. Non-structural carbohydrates were accumulated in leaves at a low soil temperature showing that growth was more hindered than net photosynthesis. Leaf anatomy differed between the first and second leaf flush of silver birch, which may promote tree fitness in the prevailing growing conditions. The interaction of birch structure and function changes with soil temperature, which can further reflect to ecosystem functioning.

Effects of elevated ozone and warming on terpenoid emissions and concentrations of Norway spruce depend on needle phenology and age.

Norway spruce (Picea abies (L.) Karst) trees are affected by ongoing climate change, including warming and exposure to phytotoxic levels of ozone. Non-volatile terpenoids and volatile terpenoids (biogenic organic volatile compounds, BVOCs) protect spruce against biotic and abiotic stresses. BVOCs also affect the atmosphere's oxidative capacity. Four-year-old Norway spruce were exposed to elevated ozone (EO) (1.4 × ambient) and warming (1.1 °C + ambient air) alone and in combination on an open-field exposure site in Central Finland. Net photosynthesis, needle terpenoid concentrations and BVOC emissions were measured four times during the experiment's second growing season: after bud opening in May, during the mid-growing season in June, and after needle maturation in August and September. Warming increased terpene concentrations in May due to advanced phenology and decreased them at the end of the growing season in matured current-year needles. Ozone enhanced these effects of warming on several compounds. Warming decreased concentrations of oxygenated sesquiterpenes in previous-year needles. Decreased emissions of oxygenated monoterpenes by warming and ozone alone in May were less prominent when ozone and warming were combined. A similar interactive treatment response in isoprene, camphene, tricyclene and α-pinene was observed in August when the temperature and ozone concentration was high. The results suggest long-term warming may reduce the terpenoid-based defence capacity of young spruce, but the defence capacity can be increased during the most sensitive growth phase (after bud break), and when high temperatures or ozone concentrations co-occur. Reduced BVOC emissions from young spruce may decrease the atmosphere's oxidative capacity in the warmer future, but the effect of EO may be marginal because less reactive minor compounds are affected.

Responses in growth and emissions of biogenic volatile organic compounds in Scots pine, Norway spruce and silver birch seedlings to different warming treatments in a controlled field experiment.

We investigated the responses in growth and emissions of biogenic volatile organic compounds (BVOCs) in Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies [L.] Karst.) and silver birch (Betula pendula Roth) seedlings to a simulated climate warming of +2 °C (T2) and +4 °C (T4), compared to the ambient conditions, during two growing seasons (2019-2020) in a controlled field experiment in central Finland. In all seedlings, height was measured weekly. Diameter was measured continuously for one seedling from each tree species per plot. For shoot and root biomass measurements, half of the seedlings were harvested at end of the first growing season and the rest at the end of the second growing season. Foliage BVOC emission rates were measured at the end of the second growing season. Biomass, height, and diameter growth of silver birch did benefit the most from warming in both growing seasons. In the Scots pine and Norway spruce seedlings, height and diameter growth increased with increasing temperature in the second growing season, more so in Scots pine. Overall, the shoot and root biomass of conifer seedlings increased with increasing temperature. In the conifer seedlings, warming increased biomass and diameter growth more than height growth, due to their predetermined height growth pattern. The warming increased BVOC emissions more clearly in silver birch, whilst the BVOC emissions were in conifers less sensitive to temperature variation. Based on our findings, silver birch seedlings could be expected to benefit the most from warmer growing conditions and Norway spruce the least.

Influence of increased nutrient availability on biogenic volatile organic compound (BVOC) emissions and leaf anatomy of subarctic dwarf shrubs under climate warming and increased cloudiness.

BACKGROUND AND AIMS: Climate change is subjecting subarctic ecosystems to elevated temperature, increased nutrient availability and reduced light availability (due to increasing cloud cover). This may affect subarctic vegetation by altering the emissions of biogenic volatile organic compounds (BVOCs) and leaf anatomy. We investigated the effects of increased nutrient availability on BVOC emissions and leaf anatomy of three subarctic dwarf shrub species, Empetrum hermaphroditum, Cassiope tetragona and Betula nana, and if increased nutrient availability modifies the responses to warming and shading. METHODS: Measurements of BVOCs were performed in situ in long-term field experiments in the Subarctic using a dynamic enclosure system and collection of BVOCs into adsorbent cartridges analysed by gas chromatography-mass spectrometry. Leaf anatomy was studied using light microscopy and scanning electron microscopy. KEY RESULTS: Increased nutrient availability increased monoterpene emission rates and altered the emission profile of B. nana, and increased sesquiterpene and oxygenated monoterpene emissions of C. tetragona. Increased nutrient availability increased leaf tissue thicknesses of B. nana and C. tetragona, while it caused thinner epidermis and the highest fraction of functional (intact) glandular trichomes for E. hermaphroditum. Increased nutrient availability and warming synergistically increased mesophyll intercellular space of B. nana and glandular trichome density of C. tetragona, while treatments combining increased nutrient availability and shading had an opposite effect in C. tetragona. CONCLUSIONS: Increased nutrient availability may enhance the protection capacity against biotic and abiotic stresses (especially heat and drought) in subarctic shrubs under future warming conditions as opposed to increased cloudiness, which could lead to decreased resistance. The study emphasizes the importance of changes in nutrient availability in the Subarctic, which can interact with climate warming and increased cloudiness effects.

Molecular and Chemical Screening for Inherent Disease Resistance Factors of Norway Spruce (Picea abies) Clones Against Conifer Stem Rot Pathogen Heterobasidion parviporum.

Root and stem rot of conifer trees caused by Heterobasidion annosum species complex leads to huge economic losses in Europe, yet not much is known about the molecular and chemical basis for host resistance. To identify inherent chemical or molecular markers in clones found to be either resistant or susceptible, we sampled needle tissues of all the clones before pathogen inoculation. We conducted a short-term resistance screening by using the pathogen H. parviporum to inoculate 70 Norway spruce clones. Based on lesion size, subsets of highly susceptible and resistant clones were further analyzed. Terpene detection and RNA sequencing were performed to explore inherent variations in genotypes differing in resistance to pathogenic challenge at chemical and transcriptional levels. A negative correlation emerged between resistance and growth. Terpene profiles of resistant clones showed higher content of monoterpenes and sesquiterpenes, with concomitant increased transcript abundance of genes involved in the terpenoid pathway. A set of upregulated genes relevant to flavonoid biosynthesis was observed in resistant genotypes, whereas higher transcripts of lignin biosynthetic genes were prevalent in susceptible clones. Genes involved in flavonoid and lignin biosynthesis as well as terpene content may have a role in facilitating resistance of Norway spruce against H. parviporum. Our results provide strong support on the feasibility of sampling needle tissues before pathogen inoculation, and the approach could be of value for large-scale screening of novel biomarkers for durable resistance. The additional insights could form a basis for further research on resistance screening in this pathosystem.

Does ozone exposure affect herbivore-induced plant volatile emissions differently in wild and cultivated plants?

Concentrations of tropospheric ozone have more than doubled in the Northern Hemisphere since pre-industrial times. Plant responses to single abiotic or biotic stresses, such as ozone exposure and herbivore-feeding, have received substantial attention, especially for cultivated plants. Modern cultivated plants have been subjected to selective breeding that has altered plant chemical defences. To understand how ozone might affect plant responses to herbivore-feeding in wild and cultivated plants, we studied the volatile emissions of brassicaceous plants after exposure to ambient (~ 15 ppb) or elevated ozone (80 ppb), with and without Plutella xylostella larvae-feeding. Results indicated that most of the wild and cultivated plants increased volatile emissions in response to herbivore-feeding. Ozone alone had a weaker and less consistent effect on volatile emissions, but appeared to have a greater effect on wild plants than cultivated plants. This study highlights that closely related species of the Brassicaceae have variable responses to ozone and herbivore-feeding stresses and indicates that the effect of ozone may be stronger in wild than cultivated plants. Further studies should investigate the mechanisms by which elevated ozone modulates plant volatile emissions in conjunction with biotic stressors.

Deposition of α-pinene oxidation products on plant surfaces affects plant VOC emission and herbivore feeding and oviposition.

White cabbage, Brassica oleracea, plants and artificial leaves covered with B. oleracea epicuticular wax were exposed to α-pinene and α-pinene oxidation products formed through the oxidation of α-pinene by ozone (O3) and hydroxyl (OH) radicals. O3 and OH-induced oxidation of α-pinene led to the formation of oxygenated volatile organic compounds (OVOCs) and secondary organic aerosol particles (SOA), referred to together as oxidation products (OP). Exposure of cabbage plants to O3 and OH-induced α-pinene OP led to the deposition and re-emission of gas-phase OP by exposed cabbage plants. In a series of 2-choice bioassays, the specialist cruciferous herbivore, Plutella xylostella adults deposited less eggs on artificial leaves exposed to α-pinene OP than on control plants exposed to clean filtered air. P. xylostella larvae did not show a specific feeding preference when offered leaves from different exposure treatments. However, the generalist Indian stick insect, Carausius morosus, fed more on control filtered air-exposed plants than on those exposed to α-pinene OP. Taken together, our results show that exposure to α-pinene oxidation products affects VOC emissions of B. oleracea and alters P. xylostella oviposition and C. morosus feeding responses.

Wildfire effects on BVOC emissions from boreal forest floor on permafrost soil in Siberia.

One of the effects of climate change on boreal forest will be more frequent forest wildfires and permafrost thawing. These will increase the availability of soil organic matter (SOM) for microorganisms, change the ground vegetation composition and ultimately affect the emissions of biogenic volatile organic compounds (BVOCs), which impact atmospheric chemistry and climate. BVOC emissions from boreal forest floor have been little characterized in southern boreal region, and even less so in permafrost soil, which underlies most of the northern boreal region. Here, we report the long-term effects of wildfire on forest floor BVOC emission rates along a wildfire chronosequence in a Larix gmelinii forest in central Siberia. We determined forest floor BVOC emissions from forests exposed to wildfire 1, 23 and > 100 years ago. We studied how forest wildfires and the subsequent succession of ground vegetation, as well as changes in the availability of SOM along with the deepened and recovered active layer, influence BVOC emission rates. The forest floor acted as source of a large number of BVOCs in all forest age classes. Monoterpenes were the most abundant BVOC group in all age classes. The total BVOC emission rates measured from the 23- and >100-year-old areas were ca. 2.6 times higher than the emissions from the 1-year-old area. Lower emissions were related to a decrease in plant coverage and microbial decomposition of SOM after wildfire. Our results showed that forest wildfires play an important indirect role in regulating the amount and composition of BVOC emissions from post-fire originated boreal forest floor. This could have a substantial effect on BVOC emissions if the frequency of forest wildfires increases in the future as a result of climate warming.

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

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

Secondary Organic Aerosol Formation from Healthy and Aphid-Stressed Scots Pine Emissions.

One barrier to predicting biogenic secondary organic aerosol (SOA) formation in a changing climate can be attributed to the complex nature of plant volatile emissions. Plant volatile emissions are dynamic over space and time, and change in response to environmental stressors. This study investigated SOA production from emissions of healthy and aphid-stressed Scots pine saplings via dark ozonolysis and photooxidation chemistry. Laboratory experiments using a batch reaction chamber were used to investigate SOA production from different plant volatile mixtures. The volatile mixture from healthy plants included monoterpenes, aromatics, and a small amount of sesquiterpenes. The biggest change in the volatile mixture for aphid-stressed plants was a large increase (from 1.4 to 7.9 ppb) in sesquiterpenes-particularly acyclic sesquiterpenes, such as the farnesene isomers. Acyclic sesquiterpenes had different effects on SOA production depending on the chemical mechanism. Farnesenes suppressed SOA formation from ozonolysis with a 9.7-14.6% SOA mass yield from healthy plant emissions and a 6.9-10.4% SOA mass yield from aphid-stressed plant emissions. Ozonolysis of volatile mixtures containing more farnesenes promoted fragmentation reactions, which produced higher volatility oxidation products. In contrast, plant volatile mixtures containing more farnesenes did not appreciably change SOA production from photooxidation. SOA mass yields ranged from 10.8 to 23.2% from healthy plant emissions and 17.8-26.8% for aphid-stressed plant emissions. This study highlights the potential importance of acyclic terpene chemistry in a future climate regime with an increased presence of plant stress volatiles.

Evaluation of potential genetic and chemical markers for Scots pine tolerance against Heterobasidion annosum infection.

MAIN CONCLUSION: Two terpene compounds and four genes were identified as potential biomarkers for further evaluation for Scots pine susceptibility or tolerance against Heterobasidion annosum. Scots pine (Pinus sylvestris) is one of the main sources of timber in the boreal zone of Eurasia. Commercial pine plantations are vulnerable to root and butt rot disease caused by the fungus Heterobasidion annosum. The pathogen affects host growth rate, causes higher mortality and decreases in timber quality, resulting in considerable economic losses to forest owners. Genetic and biochemical factors contributing to Scots pine tolerance against H. annosum infection are not well understood. We assessed the predictive values of a set of potential genetic and chemical markers in a field experiment. We determined the expression levels of 25 genes and the concentrations of 36 terpenoid compounds in needles of 16 Scots pine trees randomly selected from a natural population prior to artificial infection. Stems of the same trees were artificially inoculated with H. annosum, and the length of necrotic lesions was documented 5 months post inoculation. Higher expression level of four genes included in our analysis and encoding predicted α-pinene synthase (two genes), geranyl diphosphate synthase (GPPS), and metacaspase 5 (MC5), could be associated with trees exhibiting increased levels of necrotic lesion formation in response to fungal inoculation. In contrast, concentrations of two terpenoid compounds, ß-caryophyllene and α-humulene, showed significant negative correlations with the lesion size. Further studies with larger sample size will help to elucidate new biomarkers or clarify the potential of the evaluated markers for use in Scots pine disease resistance breeding programs.

Dual RNA-seq analysis provides new insights into interactions between Norway spruce and necrotrophic pathogen Heterobasidion annosum s.l.

BACKGROUND: Root and butt rot of conifer trees caused by fungi belonging to the Heterobasidion annosum species complex is one of the most economically important fungal diseases in commercial conifer plantations throughout the Northern hemisphere. We investigated the interactions between Heterobasidion fungi and their host by conducting dual RNA-seq and chemical analysis on Norway spruce trees naturally infected by Heterobasidion spp. We analyzed host and pathogen transcriptome and phenolic and terpenoid contents of the spruce trees. RESULTS: Presented results emphasize the role of the phenylpropanoid and flavonoid pathways in the chemical defense of Norway spruce trees. Accumulation of lignans was observed in trees displaying symptoms of wood decay. A number of candidate genes with a predicted role in the higher level regulation of spruce defense responses were identified. Our data indicate a possible role of abscisic acid (ABA) signaling in the spruce defense against Heterobasidion infection. Fungal transcripts corresponding to genes encoding carbohydrate- and lignin-degrading enzymes, secondary metabolism genes and effector-like genes were expressed during the host colonization. CONCLUSIONS: Our results provide additional insight into defense strategies employed by Norway spruce trees against Heterobasidion infection. The potential applications of the identified candidate genes as markers for higher resistance against root and butt rot deserve further evaluation.

Tissue Microbiome of Norway Spruce Affected by Heterobasidion-Induced Wood Decay.

Plants live in close association with microbial symbionts, which may affect the host fitness, productivity, and tolerance against biotic and abiotic stressors. The composition of plant microbial communities is influenced by many biotic and abiotic factors, but little is known about the effect of plant pathogens on the structure of these communities. In this study, we investigated the structure of bacterial communities associated with different tissues of asymptomatic and symptomatic (Heterobasidion-rotten) Norway spruce (Picea abies (L.) Karst.) trees. Our results demonstrated that each of the investigated anatomic tissues (root, bark, down stem, upper stem, and needles) harbored a unique bacterial assemblage. However, the health status of the host trees had little effect on the structure of bacterial communities, as the only significant differences among asymptomatic and symptomatic trees were found in the composition of the bacterial communities of needles. Proteobacteria was predominant in all anatomic regions with the highest abundance in needles (86.7%), whereas Actinobacteria showed an opposite trend, being more abundant in the woody tissues than in needles. Additionally, we performed profiling of terpenoid compounds present in spruce xylem and phloem. Total concentrations of monoterpenes and sesquiterpenes were considerably higher in asymptomatic trees. However, we found no significant correlations between terpenoid profiles of spruce trees and the composition of their bacterial communities. Our results provide an insight into the diversity of bacteria associated with Norway spruce tree tissues. At the same time, the health status and terpenoid content of host trees had a limited effect on the composition of bacterial communities in our survey.

Climate Change Effects on Secondary Compounds of Forest Trees in the Northern Hemisphere.

Plant secondary compounds (PSCs), also called secondary metabolites, have high chemical and structural diversity and appear as non-volatile or volatile compounds. These compounds may have evolved to have specific physiological and ecological functions in the adaptation of plants to their growth environment. PSCs are produced by several metabolic pathways and many PSCs are specific for a few plant genera or families. In forest ecosystems, full-grown trees constitute the majority of plant biomass and are thus capable of producing significant amounts of PSCs. We summarize older literature and review recent progress in understanding the effects of abiotic and biotic factors on PSC production of forest trees and PSC behavior in forest ecosystems. The roles of different PSCs under stress and their important role in protecting plants against abiotic and biotic factors are also discussed. There was strong evidence that major climate change factors, CO2 and warming, have contradictory effects on the main PSC groups. CO2 increases phenolic compounds in foliage, but limits terpenoids in foliage and emissions. Warming decreases phenolic compounds in foliage but increases terpenoids in foliage and emissions. Other abiotic stresses have more variable effects. PSCs may help trees to adapt to a changing climate and to pressure from current and invasive pests and pathogens. Indirect adaptation comes via the effects of PSCs on soil chemistry and nutrient cycling, the formation of cloud condensation nuclei from tree volatiles and by CO2 sequestration into PSCs in the wood of living and dead forest trees.

Understorey Rhododendron tomentosum and Leaf Trichome Density Affect Mountain Birch VOC Emissions in the Subarctic.

Subarctic vegetation is composed of mountain birch [Betula pubescens ssp. czerepanovii (MB)] forests with shrubs and other species growing in the understorey. The effects of the presence and density of one understorey shrub, Rhododendron tomentosum (RT), on the volatile emissions of MB, were investigated in a Finnish subarctic forest site in early and late growing season. Only MB trees with an RT-understorey emitted the RT-specific sesquiterpenoids, palustrol, ledol and aromadendrene. Myrcene, which is the most abundant RT-monoterpene was also emitted in higher quantities by MB trees with an RT-understorey. The effect of RT understorey density on the recovery of RT compounds from MB branches was evident only during the late season when sampling temperature, as well as RT emissions, were higher. MB sesquiterpene and total emission rates decreased from early season to late season, while monoterpene emission rate increased. Both RT and MB terpenoid emission rates were linked to density of foliar glandular trichomes, which deteriorated over the season on MB leaves and emerged with new leaves in the late season in RT. We show that sesquiterpene and monoterpene compounds emitted by understorey vegetation are adsorbed and re-released by MB, strongly affecting the MB volatile emission profile.

Ozone disrupts adsorption of Rhododendron tomentosum volatiles to neighbouring plant surfaces, but does not disturb herbivore repellency.

The perennial evergreen woody shrub, Rhododendron tomentosum, confers associational resistance against herbivory and oviposition on neighbouring plants through passive adsorption of some of its constitutively emitted volatile organic compounds (VOCs). The adsorption process is dependent on transport of VOCs in the air. In polluted atmospheres, the VOCs may be degraded and adsorption impeded. We studied the effect of elevated ozone regimes on the adsorption of R. tomentosum volatiles to white cabbage, Brassica oleracea, and the oviposition of the specialist herbivore Plutella xylostella on the exposed plants. We found evidence for adsorption and re-emission of R. tomentosum volatiles by B. oleracea plants. Ozone changed the blend of R. tomentosum volatiles and reduced the amount of R. tomentosum volatiles recovered from B. oleracea plants. However, plants exposed to R. tomentosum volatiles received fewer P. xylostella eggs than control plants exposed to filtered air irrespective of whether R. tomentosum volatiles mixed with ozone. Ozone disrupts a volatile mediated passive plant-to-plant interaction by degrading some compounds and reducing the quantity available for adsorption by neighbouring plants. The change, however, did not affect the deterrence of oviposition by P. xylostella, suggesting that aromatic companion plants of Brassica crops may confer pest-deterring properties even in ozone-polluted environments.

New Light for Phytochemicals.

Light-emitting diode (LED) lighting technology with narrow-bandwidth illumination helps to reduce energy consumption on covered crops. Here, we discuss how this new technology, which provides flexible modification of light spectra, will open new avenues for natural modulation of medicinal and crop plant metabolomes for better colour, flavour, fragrance, and antioxidant properties.