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Our objective was to study how changes in the light spectra affect growth; the concentration of carbohydrate, chlorophyll, carotenoid, terpene, alkaloid, and phenolic compounds; and the emissions of BVOC (biogenic volatile organic compound) of Norway spruce (Picea abies (L.) Karst.) seedlings. This study was conducted during the growth of the third needle generation in plant growth chambers. The plants were exposed to two light spectra with equal photon flux densities but different proportions of blue light (400–500 nm) produced by LED (light-emitting diode) lamps: (i) control (white light + 12% blue light) and (ii) increased blue light (+B) (white light + 45% blue light). The +B treatment increased the concentrations of total flavonoids and acetophenones in needles. The major changes in the phenolic profile were an accumulation of astragalin derivatives and the aglycone of picein. The +B treatment also decreased concentrations of the main alkaloid compound, epidihydropinidine, and its precursor, 2-methyl-6-propyl-1,6-piperideine; the emission rates of limonene, myrcene, and total monoterpenes; and the concentrations of a few terpenoid compounds, mainly in stems. Growth as well as the carbohydrates and pigments present in the needles were not affected. The results suggest that supplemental blue light shifts carbon allocation between secondary metabolism routes — from alkaloid and terpenoid synthesis to flavonoid and acetophenone synthesis. The changes may affect herbivory and abiotic stress tolerance of Norway spruce.

Abstract The biogenic volatile organic compounds, BVOCs have a central role in ecosystem–atmosphere interactions. High-latitude ecosystems are facing increasing temperatures and insect herbivore pressure, which may affect their BVOC emission rates, but evidence and predictions of changes remain scattered. We studied the long-term effects of + 3 °C warming and reduced insect herbivory (achieved through insecticide sprayings) on mid- and late summer BVOC emissions from field layer vegetation, supplemented with birch saplings, and the underlying soil in Subarctic mountain birch forest in Finland in 2017–2018. Reduced insect herbivory decreased leaf damage by 58–67% and total ecosystem BVOC emissions by 44–72%. Of the BVOC groups, total sesquiterpenes had 70–80% lower emissions with reduced herbivory, and in 2017 the decrease was greater in warmed plots (89% decrease) than in ambient plots (34% decrease). While non-standardized total BVOC, monoterpene, sesquiterpene and GLV emissions showed instant positive responses to increasing chamber air temperature in midsummer samplings, the long-term warming treatment effects on standardized emissions mainly appeared as changes in the compound structure of BVOC blends and varied with compounds and sampling times. Our results suggest that the effects of climate warming on the total quantity of BVOC emissions will in Subarctic ecosystems be, over and above the instant temperature effects, mediated through changes in insect herbivore pressure rather than plant growth. If insect herbivore numbers will increase as predicted under climate warming, our results forecast herbivory-induced increases in the quantity of Subarctic BVOC emissions. Graphic Abstract

Environmentally acquired chemical camouflage is a phenomenon, where a plant growing close to a strong volatile organic compound (VOC) emitter will adsorb and re-emit the VOCs produced by the neighbouring plant. The re-emitted volatile bouquet may resemble more the VOC composition of the neighbour than plant’s own typical odour, and thus act as chemical camouflage against insect detection, potentially simultaneously providing associational resistance towards herbivory. We exposed a pest-sensitive horticultural crop, Brassica oleracea var. italica (broccoli) cv. Lucky, to the volatiles emitted by Rhododendron tomentosum [RT] twigs and assessed the host selection by ovipositing females and larval instars of the major caterpillar pest Pieris brassicae between RT-exposed and control plants. Potential impact of RT exposure on herbivore natural enemies was studied using behavioural tests with a parasitoid wasp Cotesia glomerata. P. brassicae females laid significantly less eggs and egg clusters were fewer on RT-exposed plants at both night-time (6 °C) and daytime (22 °C) temperatures. Larvae preferred leaves from control plants over RT-exposed plants at both temperatures. Preceding RT-exposure did not disturb orientation of parasitoid wasp Cotesia glomerata females towards B. oleracea plants damaged by its host P. brassicae. However, host-damaged control plants were favoured by the parasitoid over RT-exposed, host-damaged plants. Our results suggest that companion plant based chemical camouflage as a mechanism of pest suppression could be developed as an additional tool for the integrated pest management toolbox in agriculture.

Volatile organic compounds (VOCs) produced by plants are essential indicators of their physiological response to environmental conditions. But evidence of natural variation in VOC emissions and their contributing factors is still limited, especially for non-cultivated species. Here we explored the natural volatile emissions of Dracophyllum subulatum Hook.f., an endemic shrub to the North Island Central Plateau of New Zealand, and determined some environmental factors driving the plant’s emissions. Volatile emissions of D. subulatum were measured on four separate occasions from December 2017 to September 2018 using the “push-pull” headspace sampling technique and analyzed using gas chromatography-mass spectrometry (GC-MS). D. subulatum was classified based on the volatiles measured on each sampling occasion using linear discriminant analysis (LDA). On each sampling occasion, we also recorded and compared ambient air temperature, herbivory damage, total soil nitrogen (N), available phosphorus (P), potassium (K), and soil moisture content. The relationship between environmental variables that differed significantly between sampling occasions and volatile emissions were estimated using generalized linear models (GLMs). Based on VOCs measured on each sampling occasion, we were able to distinguish different chemical profiles. Overall, we found that total emission and the relative proportions of all major chemical classes released by D. subulatum were significantly higher during summer. The GLMs reveal that differences in environmental factors between the four sampling occasions are highly associated with changing emissions. Higher temperatures in summer had a consistently strong positive relationship with emissions, while the impacts of soil moisture content, P and K were variable and depended on the chemical class. These results are discussed, particularly how high temperature (warming) may shape volatile emissions and plants’ ecology.

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.

Living trees are the main source of biogenic volatile organic compounds (BVOCs) in forest ecosystems, but substantial emissions originate from leaf and wood litter, the rhizosphere and from microorganisms. This review focuses on temperate and boreal forest ecosystems and the roles of BVOCs in ecosystem function, from the leaf to the forest canopy and from the forest soil to the atmosphere level. Moreover, emphasis is given to the question of how BVOCs will help forests adapt to environmental stress, particularly biotic stress related to climate change. Trees use their vascular system and emissions of BVOCs in internal communication, but emitted BVOCs have extended the communication to tree population and whole community levels and beyond. Future forestry practices should consider the importance of BVOCs in attraction and repulsion of attacking bark beetles, but also take an advantage of herbivore-induced BVOCs to improve the efficiency of natural enemies of herbivores. BVOCs are extensively involved in ecosystem services provided by forests including the positive effects on human health. BVOCs have a key role in ozone formation but also in ozone quenching. Oxidation products form secondary organic aerosols that disperse sunlight deeper into the forest canopy, and affect cloud formation and ultimately the climate. We also discuss the technical side of reliable BVOC sampling of forest trees for future interdisciplinary studies that should bridge the gaps between the forest sciences, health sciences, chemical ecology, conservation biology, tree physiology and atmospheric science.

Volatile organic compounds (VOCs) are a major currency in plant communication where they mediate above- and below-ground interactions between plants and the surrounding organisms (i.e., other plants, microorganisms, pollinators, seed dispersers, herbivores, and their natural enemies). Considering the multiple interactions mediated by VOCs and their impact on a plant’s reproductive success and survival, they can be a crucial weapon in plant-plant competition. However, this particular role of VOCs is underrepresented in the literature. Mechanisms by which volatiles can mediate plant competition can be direct or indirect. Direct mechanisms include establishing a neighbour’s identity and status to select adequate responses and affecting competitor’s seed germination or growth through VOC-mediated allelopathy. Indirect mechanisms can affect the plant's competitive ability by modifying interactions with other trophic levels, for instance, through associational resistance or chemical camouflage. These mechanisms are not mutually exclusive and can be seen as part of a continuum. In this review, we present evidence from the literature to illustrate these roles and indicate how they could influence competition in plant communities. We propose new research avenues to test if and how these mechanisms affect competitive outcomes and suggest that, in addition to morphological traits, future competition studies should also incorporate data on plant-volatile emissions and measure their effects on the surrounding plants and other trophic levels. This information would allow us to understand competition from a broader perspective by acknowledging the existence of multiple (possibly coexisting) competition strategies and the role of other trophic levels in shaping competitive outcomes.

Plants emit an extremely diverse bouquet of volatile organic compounds (VOCs) from their above-ground and below-ground parts. Emissions are constitutive or induced, e.g. by herbivores. VOCs can be classified as highly volatile, volatile and semi-volatile compounds. Sesquiterpenes (SQTs) are typical semi-volatile organic compounds (sVOCs) released by plants. Similarly, herbivore-induced homoterpenes and methyl salicylate also have relatively low volatility. SVOCs have a high boiling point (> 240 °C) and a vapour pressure below 0.005 kPa at 25 °C. Glandular trichomes on plant surfaces can store SQTs in mixtures with more volatile VOCs, which are released into the air by diffusion or after gland rupture. The sVOCs stored in glandular trichomes often have repellent effects on herbivores, while herbivore-induced sVOCs are known for their attractiveness to natural enemies of herbivores, i.e. they act in indirect chemical defence of plants. Due to their low volatility, sVOCs produced by plants may easily adhere to the surfaces of emitter and neighbouring plants during the colder temperatures that plants face, e.g. at night. On the foliage of neighbouring receiver plants, sVOCs may act in direct and indirect defence of that plant species. When the temperature rises again, sVOCs are released into the atmosphere. The semi-volatile reaction products of highly volatile plant monoterpenes and photochemical pollutants such as ozone could constitute further sVOCs on plant leaf surfaces. Here, we review recent literature of the plant surface–environment interaction of biogenic sVOCs and particularly evaluate potential crop protection strategies such as intercropping and companion planting using sVOC-emitting species. Foliage typically forms the widest surfaces on crop plants, and foliar herbivory is a major type of pest damage during the vegetative stage of crop plants. Foliage is also a major source of herbivore-induced VOC emissions. Consequently, we focus on foliage-mediated sVOCs and their potential in pest management.

Phenolic metabolites in forest trees play a key role in the defence against biotic and abiotic stressors, yet we lack information about the effects of combined abiotic factors on phenolic compounds in conifers. We studied the effects of combined abiotic factors (ozone × temperature, ozone × nitrogen, temperature × nitrogen, and ozone × temperature × nitrogen) on phenolic concentrations in stems of Scots pine (Pinus sylvestris L.) seedlings in a 3-year-long field experiment in central Finland. In current-year stems, elevated ozone increased the concentrations of total phenolics, soluble proanthocyanidins, and total proanthocyanidins, while warming reduced the concentrations of piceatannol glucoside, pinosylvin, isorhamnetin + kaempferol-3-rhamnoside, and monocoumaroyl isoquercitrin 1. Complex interaction effects on current-year stems showed that nitrogen addition increased the concentrations of some flavonoids in ambient ozone and temperature levels. In the stems from the previous years, ozone decreased the concentrations of total phenolics, total proanthocyanidins, and several flavonoids in ambient temperature, while warming increased their concentrations in combination with elevated ozone. Our results suggest that phenolic defence responses in Scots pine seedlings are affected by all three factors, but that the level of phenolics in the stems from previous years may increase under the combined exposure to elevated temperature and ozone — an expected climate trend in the Northern Hemisphere.

The New Zealand tea tree Leptospermun scoparium (mānuka) is widely known for the antimicrobial properties of its honey. Mānuka is native to New Zealand, growing in a range of environments, including the Central Volcanic Plateau of the North Island, where it is currently threatened by the spread of exotic invasive weeds such as heather (Calluna vulgaris) and Scotch broom (Cytisus scoparius). Here, we characterise for the first time the aboveground volatile organic compounds (VOCs) produced by mānuka in this area, during summer and winter seasons, in weed-invaded and non-invaded stands. We measured plant volatiles at four sites, each with a distinct combination of woody species: (1) conspecific stands of mānuka; (2) mānuka and another native species (Dracophyllum subulatum); and mānuka with one of two European invasive plants, (3) heather or (4) Scotch broom. We also quantified herbivore damage on target mānuka plants and analysed microclimatic variables (soil nutrients, air temperature and soil water content) to investigate their impact on volatile emissions. Our results reveal a strong seasonal effect on volatile emissions, but also significant differences between sites associated with biotic and abiotic changes partly driven by invasive plants. Overall, volatile emission rates from mānuka were typically lower at sites where invaders were present. We point to several factors that could contribute to the observed emission patterns and areas of interest for future research to provide a comprehensive understanding of VOC emissions in nature. Given the vital role of volatile compounds in plant communication, we also recommend future studies to be performed in multiple seasons, with larger sample sizes and more study sites to expand on these findings and explore the ecological impacts of changes in VOC emissions during plant invasion.

Hybrid aspen (Populus tremula × tremuloides Michx.) is a fast-growing tree species used for short-rotation forestry in northern latitudes. Aspen species have a rich herbivore fauna, including defoliating leaf beetles that induce emissions of volatile organic compounds (VOCs) when feeding on aspen leaves. We investigated the differential induction of VOCs by male and female Phratora laticollis leaf beetles feeding on hybrid aspen and the differences in the orientation of beetles in response to gender-specific induced VOCs. The hypotheses for the study were (1) the VOCs in the headspace of plants infested with beetles of the two genders individually and in mixed aggregates would vary subtly, and (2) foraging adult beetles would be able to detect differences in VOC blends and use them to fine-tune their orientation choices. In Y-tube bioassays, both females and males preferred VOCs from leaves damaged by one gender (females or males) over undamaged leaves. However, if leaves were damaged by a two-gender population, neither females nor males indicated a preference over volatiles of undamaged leaves. Leaves damaged by both beetle genders simultaneously had significantly increased green leaf volatile (GLV), benzenoid and homoterpene emissions compared to undamaged leaves. Emissions of these compounds possibly indicate higher herbivore pressure and a higher risk of attack by parasitoids and predators and could thus be the cause of the lack of beetle preference. Our findings provide new basic information on gender-based host plant selection by herbivores and may be helpful in the development of sustainable biogenic VOC-based herbivore-control methods for intensive short-rotation hybrid aspen production.

Carbohydrate-rich extrafloral nectar (EFN) is produced in nectaries on the leaves, stipules, and stems of plants and provides a significant energy source for ants and other plant mutualists outside of the flowering period. Our review of literature on EFN indicates that only a few forest plant species in cool boreal environments bear EFN-producing nectaries and that EFN production in many boreal and subarctic plant species is poorly studied. Boreal forest, the world’s largest land biome, is dominated by coniferous trees, which, like most gymnosperms, do not produce EFN. Notably, common deciduous tree species that can be dominant in boreal forest stands, such as Betula and Alnus species, do not produce EFN, while Prunus and Populus species are the most important EFN-producing tree species. EFN together with aphid honeydew is known to play a main role in shaping ant communities. Ants are considered to be keystone species in mixed and conifer-dominated boreal and mountain forests because they transfer a significant amount of carbon from the canopy to the soil. Our review suggests that in boreal forests aphid honeydew is a more important carbohydrate source for ants than in many warmer ecosystems and that EFN-bearing plant species might not have a competitive advantage against herbivores. However, this hypothesis needs to be tested in the future. Warming of northern ecosystems under climate change might drastically promote the invasion of many EFN-producing plants and the associated insect species that consume EFN as their major carbohydrate source. This may result in substantial changes in the diet preferences of ant communities, the preventative roles of ants against insect pest outbreaks, and the ecosystem services they provide. However, wood ants have adapted to using tree sap that leaks from bark cracks in spring, which may mitigate the effects of improved EFN availability.

Scots pine (Pinus sylvestris L.) is an important source of biogenic volatile organic compounds (BVOCs) in the boreal zone. BVOC emission rate and profile affect air quality, climate forcing, plant stress tolerance, and thus the growing conditions of forests. BVOC emission profile of shoots and forest floor, as well as emission rates from forest floor, were studied in a latitudinal provenance experiment with 19-year-old Scots pine common garden in Central Finland. The provenances studied were Saaremaa (SAA, 58°22′), Korpilahti (KOR, 62°0′), Suomussalmi (SUO, 65°10′), and Muonio (MUO, 67°56′). A chemotype with high proportion of Δ-3-carene, terpinolene, sabinene, γ-terpinene, and α-terpinene was significantly more common for the southern SAA than the northern SUO and MUO provenances. A chemotype with high proportion of α-pinene, β-pinene, limonene, and myrcene was more common in the three northernmost provenances. The main compounds emitted by forest floor were α-pinene, Δ-3-carene, and camphene. Similarly to shoot emissions, forest floor emissions from SAA had highest proportion of Δ-3-carene. Average total VOC emission rate from forest floor was 50 μg·m−2·h−1 at the end of August. Total emission rates were 65% higher in KOR than in MUO. High emission rates were explained by the high amount of decomposing needle litter and low moss coverage.

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.

Stress-induced changes to plant biochemistry and physiology can influence plant nutritional quality and subsequent interactions with herbivorous pests. However, the effects of stress combinations are unpredictable and differ to the effects of individual stressors. Here we studied the effects of exposure to the phytotoxic air-pollutant ozone (O

It is not clear how climate change in combination with increasing soil nitrogen availability and herbivory affects boreal forests, the largest terrestrial biome in the world. In this study, Scots pine ( Pinus sylvestris ) seedlings were exposed to moderate warming (ca. 1 °C), 1.5 × ambient ozone (O 3 ) concentration, fertilizer addition (120 kg N ha −1 yr −1 ) and shoot herbivory by pine sawfly ( Acantholyda posticalis ) alone and in combination. We measured fine root morphology, mycorrhizal colonization level, root fungal biomass (ergosterol), rhizosphere emission of biogenic volatile organic compounds (BVOCs), and microbial biomass (PLFAs) in the rhizosphere soil as well as seedling above- and below-ground growth. Warming and fertilization effects on fine root proportions or root fungal biomass occurred in combination with other factors, combination effects being usually negative on the studied variables, or then warming and fertilization in combination cancelled some other factors’ effects. O 3 effects on needle growth, root fungal biomass and BVOCs were more often seen after the third exposure year, and sometimes only in combination with other studied factors. Of abiotic factors, fertilizer addition had reducing effect on rhizosphere BVOCs. Though increased nitrogen availability and warming increased both shoot and root dry masses, growth allocation to above- and below-ground parts was not equally increased in the combined exposures. Thus, we conclude that climate change factors together with increased nitrogen availability and herbivory are likely to affect the below-ground compartments negatively, more often than shoots, and ultimately change in growth allocation pattern which may affect overall seedling growth and survival in later years.

Climate change may promote the frequency of insect attacks such as outbreaks of the great web-spinning pine sawfly (Acantholyda posticalis) on Scots pine (Pinus sylvestris L.). We determined the emission rates of localized biogenic volatile organic compounds (BVOCs) from A. posticalis-fed branches, and systemic BVOCs from non-fed branches of Scots pine seedlings defoliated for two growing seasons by A. posticalis larvae. Seedlings were also exposed to warming, elevated ozone and higher nitrogen availability for three years in an open-field experiment. A. posticalis feeding increased localized emissions of total non-oxygenated monoterpenes 21-fold, total oxygenated monoterpenes 9.1-fold, total sesquiterpenes 11-fold and total green leaf volatiles 9.2-fold from insect-damaged shoots on the 7th day of feeding in June. Warming reduced the effects of herbivory on the emission rates of total non-oxygenated monoterpenes by 77%. However, the effect of herbivory on total sesquiterpene emissions was enhanced by 16-fold in combination with warming and elevated ozone. The localized emission rates of total BVOCs were linearly increased when feeding damage intensity by larvae exceeded 80%. After three weeks of continuous sawfly feeding, herbivory stress increased systemic emissions of total non-oxygenated monoterpenes 5.6-fold, total sesquiterpenes 5.6-fold and total green leaf volatiles 6.5-fold from the non-damaged branch of larvae-fed seedlings, and this effect on total non-oxygenated monoterpene emission was enhanced 8.6-fold with elevated ozone. Herbivory sporadically showed post-feeding effects still by the end of 12th week from the initiation of feeding, increasing total non-oxygenated monoterpene emissions 4.4-fold at elevated nitrogen level. Our results suggest that Scots pine, at least in seedling stage, will be a stronger source of BVOC emissions in future due to expected increase of sawfly outbreaks with climate warming and by increased herbivory interactions with abiotic climate change factors.

Based on available literature, ecology and economy of LED (light emitting diode) lights in plant foods production were assessed and compared to HPS (high pressure sodium) and CFL (compact fluorescent light) lamps. The assessment summarises that LEDs are superior compared to other lamp types. LEDs are ideal in luminous efficiency, life span and electricity usage. Mercury, carbon dioxide and heat emissions are also lowest in comparison to HPS and CFL lamps. This indicates that LEDs are indeed economic and eco-friendly lighting devices. The present review indicates also that LEDs have many practical benefits compared to other lamp types. In addition, they are applicable in many purposes in plant foods production. The main focus of the review is the targeted use of LEDs in order to enrich phytochemicals in plants. This is an expedient to massive improvement in production efficiency, since it diminishes the number of plants per phytochemical unit. Consequently, any other production costs (e.g., growing space, water, nutrient and transport) may be reduced markedly. Finally, twenty four research articles published in 2013–2017 were reviewed for targeted use of LEDs in the specific, i.e. blue range (400–500 nm) of spectrum. The articles indicate that blue light is efficient in enhancing the accumulation of health beneficial phytochemicals in various species. The finding is important from the global food production.

Chemical communication is ubiquitous. The identification of conserved structural elements in visual and acoustic communication is well established, but comparable information on chemical communication displays (CCDs) is lacking. We assessed the phenotypic integration of CCDs in a meta‐analysis to characterize patterns of covariation in CCDs and identified functional or biosynthetically constrained modules. Poorly integrated plant CCDs (i.e. low covariation between scent compounds) support the notion that plants often utilize one or few key compounds to repel antagonists or to attract pollinators and enemies of herbivores. Animal CCDs (mostly insect pheromones) were usually more integrated than those of plants (i.e. stronger covariation), suggesting that animals communicate via fixed proportions among compounds. Both plant and animal CCDs were composed of modules, which are groups of strongly covarying compounds. Biosynthetic similarity of compounds revealed biosynthetic constraints in the covariation patterns of plant CCDs. We provide a novel perspective on chemical communication and a basis for future investigations on structural properties of CCDs. This will facilitate identifying modules and biosynthetic constraints that may affect the outcome of selection and thus provide a predictive framework for evolutionary trajectories of CCDs in plants and animals., The project was supported by the Graduate School ‘Evolutionary Networks: Organisms, Reactions, Molecules’ (E‐Norm) of the Heinrich‐Heine‐University, Düsseldorf, Germany, the Max Planck Society and the Deutsche Forschungsgemeinschaft (DFG JU 2856/1‐1 and DFG JU 2856/2‐2).

Red autumn colouration of trees is the result of newly synthesized anthocyanin pigments in senescing autumn leaves. As anthocyanin accumulation is costly and the trait is not present in all species, anthocyanins must have an adaptive significance in autumn leaves. According to the coevolution hypothesis of autumn colours, red autumn leaves warn herbivorous insects – especially aphids that migrate to reproduce in trees in the autumn – that the tree will not be a suitable host for their offspring in spring due to a high level of chemical defence or lack of nutrients. The signalling allows trees to avoid herbivores and herbivores to choose better host trees. In this study the coevolution hypothesis was tested with four deciduous tree species that have red autumn leaf colouration – European aspen (Populus tremula L.) (Salicaceae), rowan (Sorbus aucuparia L.) (Rosaceae), mountain birch [Betula pubescens ssp. czerepanovii (NI Orlova) Hamet‐Ahti], and dwarf birch (Betula nana L.) (Betulaceae), and with two generalist herbivores, the autumnal moth [Epirrita autumnata (Borkhausen)] and the winter moth [Operophtera brumata (L.)] (both Lepidoptera: Geometridae). Anthocyanin concentrations of autumn leaves were determined from leaf samples and the growth performance parameters of the moth larvae on the study trees were measured in the spring. Trees with higher anthocyanin concentration in the autumn were predicted to be low‐quality food for the herbivores. Our results clearly showed that anthocyanin concentration was not correlated with the growth performance of the moths in any of the studied tree species. Thus, our study does not support the coevolution hypothesis of autumn colours.

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 (O

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 alpha-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, beta-caryophyllene and alpha-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.

Plants emit biogenic volatile organic compounds (BVOCs) causing transcriptomic, metabolomic and behavioral responses in receiver organisms. Volatiles involved in such responses are often called "plant language". Arthropods having sensitive chemoreceptors can recognize language released by plants. Insect herbivores, pollinators and natural enemies respond to composition of volatiles from plants with specialized receptors responding to different types of compounds. In contrast, the mechanism of how plants "hear" volatiles has remained obscured. In a plant-plant communication, several individually emitted compounds are known to prime defense response in receiver plants with a specific manner according to the chemical structure of each volatile compound. Further, composition and ratio of volatile compounds in the plant-released plume is important in plant-insect and plant-plant interactions mediated by plant volatiles. Studies on volatile-mediated plant-plant signaling indicate that the signaling distances are rather short, usually not longer than one meter. Volatile communication from plants to insects such as pollinators could be across distances of hundreds of meters. As many of the herbivore induced VOCs have rather short atmospheric life times, we suggest that in long-distant communications with plant volatiles, reaction products in the original emitted compounds may have additional information value of the distance to emission source together with the original plant-emitted compounds.

Climate change in the boreal forests include, e.g., warming, increased tropospheric ozone concentration, higher nitrogen (N) deposition and increased risk of insect outbreaks. Climate change influences emissions of biogenic volatile organic compounds (BVOCs) affecting plant defense, communication and atmospheric feedbacks. We studied the effects of elevated temperature (ca. 1 °C), elevated ozone (ca. 1.5 × ambient), two soil N availability levels (prevailing and 120 kg N ha−1 a−1) and herbivory on BVOC emission rates, net photosynthesis and resin canals (BVOCs storage), of Scots pine (Pinus sylvestris) seedlings in an open-field exposure in central Finland. Shoot BVOCs were collected in July 2012 within a few days after feeding by larvae of pine-sawfly Acantholyda posticalis, a month later in August, and in May 2013. Elevated temperature caused twofold to fourfold increases in total emissions of non-oxygenated monoterpenes (MTs), oxygenated MTs and sesquiterpenes (SQTs) and several reactive compounds, and higher N enhanced some of these changes. Ozone and higher N together increased emissions of several MTs and total SQTs. Higher number of resin canals and higher net photosynthesis might have contributed to BVOC increases. Herbivory had the strongest effect on SQT emissions (threefold increase) shortly after feeding. In the following spring, herbivory reduced emission rates of some MTs, but also synergistically increased MTs emissions with temperature but suppressed the increase caused by ozone. Results suggest that warming and ozone, particularly in areas with increased soil N availability, can increase BVOC emissions from young boreal forests in the near future, and herbivory may modify these responses.

Stress-induced changes to plant biochemistry and physiology can influence plant nutritional quality and subsequent interactions with herbivorous pests. However, the effects of stress combinations are unpredictable and differ to the effects of individual stressors. Here we studied the effects of exposure to the phytotoxic air-pollutant ozone (O3), feeding by larvae of the large cabbage white butterfly (Pieris brassicae), and a combination of the two stresses, on the emission of volatile organic compounds (VOCs) by black mustard plants (Brassica nigra) under field and laboratory conditions. Field-grown B. nigra plants were also measured for carbon-nitrogen (C–N) content, net photosynthetic activity (Pn), stomatal conductance (gs) and biomass. The effects of O3 on interactions between plants and a herbivorous pest were addressed by monitoring the abundance of wild diamondback moth larvae (Plutella xylostella) and feeding-damage to B. nigra plants in an O3-free air concentration enrichment (O3-FACE) field site. Herbivore-feeding induced the emission of VOCs that were not emitted by undamaged plants, both under field and laboratory conditions. The combination of O3 and herbivore-feeding stresses resulted in enhanced emission rates of several VOCs from field-grown plants. Short-term O3 exposure (of 10 days) and P. brassicae-feeding did not affect C–N content, but chronic O3 exposure (of 34 and 47 days) and P. brassicae-feeding exacerbated suppression of Pn. Ozone exposure also caused visible injury and decreased the plant biomass. Field-grown B. nigra under elevated O3 were infested with fewer P. xylostella larvae and received significantly less feeding damage. Our results suggest that plants growing in a moderately polluted environment may be of reduced quality and less attractive to foraging herbivores.

Heather (Calluna vulgaris) and broom (Cytisus scoparius), originally from Europe, are the main invasive plants on New Zealand&rsquo, s North Island Central Plateau, where they threaten native flora and fauna. Given the strong link between arthropod communities and plants, we explored the impact of these invasive weeds on the diversity and composition of associated arthropod assemblages in this area. The arthropods in heather-invaded areas, broom-invaded areas, and areas dominated by the native species mānuka (Leptospermum scoparium) and Dracohyllum (Dracophyllum subulatum) were collected and identified to order. During summer and autumn, arthropods were collected using beating trays, flight intercept traps and pitfall traps. Diversity indices (Richness, Shannon&rsquo, s index and Simpson&rsquo, s index) were calculated at the order level, and permutational multivariate analysis (PERMANOVA) was used to explore differences in order-level community composition. Our results show a significant variation in community composition for all trapping methods in both seasons, whereas invasive plants did not profoundly impact arthropod order richness. The presence of broom increased arthropod abundance, while heather was linked to a reduction. Under all possible plant pairings between heather, broom, mānuka, and Dracophylum, the impact of neighbouring plant identity on arthropod community composition was further explored for the samples collected using beating trays. The results suggest that during plant invasion, arthropod communities are affected by neighbouring plant identity and that impacts vary between arthropod sampling methods and seasons.

Calluna vulgaris (heather) is an aggressive invasive weed on the Central Plateau, North Is., New Zealand (NZ), where it encounters different environmental factors compared to its native range in Europe, such as high ultraviolet radiation (UV) and a lack of specialist herbivores. The specialist herbivore Lochmaea suturalis (heather beetle) was introduced from the United Kingdom (UK) in 1996 as a biocontrol agent to manage this invasive weed. Like other plant invaders, a novel environment may be challenging for heather as it adjusts to its new conditions. This process of &ldquo, adjustment&rdquo, involves morphological and physiological changes often linked to phenotypic plasticity. The biochemical responses of exotic plants to environmental variables in their invaded range is poorly understood. The production and release of volatile organic compounds (VOCs) is essential to plant communication and highly susceptible to environmental change. This study therefore aimed to explore the VOC emissions of heather in response to different levels of UV exposure, and to feeding damage by L. suturalis. Using tunnel houses clad with UV-selective filters, we measured VOCs produced by heather under NZ ambient, 20% attenuated, and 95% attenuated solar UV treatments. We also compared VOC emissions in the field at adjacent sites where L. suturalis was present or absent. Volatiles produced by the same target heather plants were measured at four different times in the spring and summer of 2018-2019, reflecting variations in beetle&rsquo, s abundance, feeding stage and plant phenology. Heather plants under 95% attenuated UV produced significantly higher amounts of (E)-&beta, farnesene, decanal, benzaldehyde, and benzeneacetaldehyde compared to 25% attenuated and ambient UV radiation. We also found significant differences in volatiles produced by heather plants in beetle-present versus beetle-absent sites on most sampling occasions. We also recorded a lower number of generalist herbivores on heather at sites where L. suturalis was present. Interactions between invasive plants, a novel environment, and the native communities they invade, are discussed.

Biotic stresses on forest trees are caused by various pest insects and plant pathogens. Attack by these parasites is known to induce the emissions of various biogenic volatile organic compounds (BVOCs), and the profile of these emissions often differs between infested and healthy plants. This difference in emission profile can be used for the non-destructive early-stage diagnosis of the stressor organism. We studied how phloem feeding by a large pine aphid (Cinara pinea Mordvilko) on the branch bark of Scots pine (Pinus sylvestris L.) affects BVOC emissions compared to those of healthy plants in two experiments. We found that in aphid-infested plants, methyl salicylate (MeSA) emissions significantly increased, and the emission rates were dependent on aphid density on the studied branch. Aphid infestation did not significantly affect total monoterpene emission, while the emissions of total sesquiterpenes were substantially higher in aphid-infested saplings than in uninfested plants. Sesquiterpene (E, E)-&alpha, farnesene was emitted at increased rates in both experiments, and the aphid alarm pheromone sesquiterpene (E)-&beta, farnesene, only in the experiment with higher aphid pressure. We conclude that the rapid increase in MeSA emissions is the most reliable indicator of aphid infestation in pine trees together with (E, E)-&alpha, farnesene.

Invasive plants pose a threat to natural ecosystems, changing the community composition and ecological dynamics. One aspect that has received little attention is the production and emission of volatile organic compounds (VOCs) by invasive plants. Investigating VOCs is important because they are involved in vital ecological interactions such as pollination, herbivory and plant competition. Heather, Calluna vulgaris, is a major invasive weed in New Zealand, especially on the Central Plateau, where it has spread rapidly since its introduction in 1912, outcompeting native species. However, the chemical behaviour of heather in its invaded ranges is poorly understood. We aimed to explore the natural variation in volatile emissions of heather and the biotic and abiotic factors influencing them on the Central Plateau of New Zealand. To this end, foliar volatiles produced by heather at four different sites were collected and analysed using gas chromatography coupled to mass spectrometry. Soil properties, herbivory and other environmental data were also collected at each site to investigate their effects on VOC emissions using generalised linear models (GLMs). Our results reveal significant differences in VOC emissions between sites and suggest that soil nutrients are the main factor accounting for these differences. Herbivory and temperature had only a minor effect, while soil water content had no impact. Further studies are needed to investigate how these variations in the invasive plant&rsquo, s foliar volatiles influence native species.

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.

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.

Secondary organic aerosol (SOA) impact climate by scattering and absorbing radiation and contributing to cloud formation. SOA models are based on studies of simplified chemical systems that do not account for the chemical complexity in the atmosphere. This study investigated SOA formation from a mixture of real Scots pine (Pinus sylvestris) emissions including a variety of monoterpenes and sesquiterpenes. SOA generation was characterized from different combinations of volatile compounds as the plant emissions were altered with an herbivore stress treatment. During active herbivore feeding, monoterpene and sesquiterpene emissions increased, but SOA mass yields decreased after accounting for absorption effects. SOA mass yields were controlled by sesquiterpene emissions in healthy plants. In contrast, SOA mass yields from stressed plant emissions were controlled by the specific blend of monoterpene emissions. Conservative estimates using a box model approach showed a 1.5- to 2.3-fold aerosol enhancement when the terpene complexity was taken into account. This enhancement was relative to the commonly used model monoterpene, “α-pinene”. These results suggest that simplifying terpene complexity in SOA models could lead to underpredictions in aerosol mass loading.

Article, Boreal forests are a major source of climate-relevant biogenic secondary organic aerosols (SOAs) and will be greatly influenced by increasing temperature. Global warming is predicted to not only increase emissions of reactive biogenic volatile organic compounds (BVOCs) from vegetation directly but also induce large-scale insect outbreaks, which significantly increase emissions of reactive BVOCs. Thus, climate change factors could substantially accelerate the formation of biogenic SOAs in the troposphere. In this study, we have combined results from field and laboratory experiments, satellite observations and global-scale modelling in order to evaluate the effects of insect herbivory and large-scale outbreaks on SOA formation and the Earth's climate. Field measurements demonstrated 11-fold and 20-fold increases in monoterpene and sesquiterpene emissions respectively from damaged trees during a pine sawfly (Neodiprion sertifer) outbreak in eastern Finland. Laboratory chamber experiments showed that feeding by pine weevils (Hylobius abietis) increased VOC emissions from Scots pine and Norway spruce seedlings by 10–50 fold, resulting in 200–1000-fold increases in SOA masses formed via ozonolysis. The influence of insect damage on aerosol concentrations in boreal forests was studied with a global chemical transport model GLOMAP and MODIS satellite observations. Global-scale modelling was performed using a 10-fold increase in monoterpene emission rates and assuming 10 % of the boreal forest area was experiencing outbreak. Results showed a clear increase in total particulate mass (local max. 480 %) and cloud condensation nuclei concentrations (45 %). Satellite observations indicated a 2-fold increase in aerosol optical depth over western Canada's pine forests in August during a bark beetle outbreak. These results suggest that more frequent insect outbreaks in a warming climate could result in substantial increase in biogenic SOA formation in the boreal zone and, thus, affect both aerosol direct and indirect forcing of climate at regional scales. The effect of insect outbreaks on VOC emissions and SOA formation should be considered in future climate predictions., published version, http://purl.org/eprint/status/PeerReviewed

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.

Some insectivorous birds orient towards insect-defoliated trees even when they do not see the foliar damage or the herbivores. There are, however, only a few studies that have examined the mechanisms behind this foraging behaviour. Previous studies suggest that birds can use olfactory foraging cues (e.g. volatile organic compounds (VOCs) emitted by defoliated plants), indirect visual cues or a combination of the two sensory cues. VOCs from insect-defoliated plants are known to attract natural enemies of herbivores, and researchers have hypothesized that VOCs could also act as olfactory foraging cues for birds. We conducted three experiments across a range of spatial scales to test this hypothesis. In each experiment, birds were presented with olfactory cues and their behavioural responses or foraging outcomes were observed. In the first experiment, two different VOC blends, designed to simulate the volatile emissions of mountain birch (Betula pubescens ssp. czerepanovii) after defoliation by autumnal moth (Epirrita autumnata) larvae, were used in behavioural experiments in aviaries with pied flycatchers (Ficedula hypoleuca). The second experiment was a field-based trial of bird foraging efficiency; the same VOC blends were applied to mountain birches, silver birches (B. pendula) and European white birches (B. pubescens) with plasticine larvae attached to the trees to serve as artificial prey for birds and provide a means to monitor predation rate. In the third experiment, the attractiveness of silver birch saplings defoliated by autumnal moth larvae versus intact controls was tested with great tits (Parus major) and blue tits (Cyanistes caeruleus) in an aviary. Birds did not orient towards either artificial or real trees with VOC supplements or towards herbivore-damaged saplings when these saplings and undamaged alternatives were hidden from view. These findings do not support the hypothesis that olfactory foraging cues are necessary in the attraction of birds to herbivore-damaged trees.

Plant volatiles mediate a range of interactions across and within trophic levels, including plant–plant interactions. Volatiles emitted by a plant may trigger physiological responses in neighboring plants or adhere to their surfaces, which, in turn, may affect the responses of the neighboring plant to herbivory. These volatiles are subject to chemical reactions during transport in air currents, especially in a polluted atmosphere. We conducted a field experiment to test for the adsorption of dispenser-released myrcene on the surfaces of cabbage plants and the effects of distance from the dispenser and elevated ozone levels (1.4× ambient) on the process. We also tested the effects of the same treatments on oviposition on cabbage plants by naturally occurring Plutella xylostella. Under low ambient ozone conditions of central Finland, there was evidence for the adsorption and re-release of myrcene by cabbage plants growing at a distance of 50 cm from myrcene dispensers. This effect was absent at elevated ozone levels. The number of eggs deposited by P. xylostella was generally lower in plots under elevated ozone compared to ambient control plots. Our results indicate that passive adsorption and re-release of a volatile monoterpene can occur in nature; however, this process is dependent upon the distance between emitter source and receiver plants as well as the concentration of atmospheric pollutants in the air. We conclude that, in the development of field-scale use of plant volatiles in modern pest control, the effects of distances and air pollution should be considered., final draft, peerReviewed

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.

Acclimation of conifer needle anatomy to climate change is poorly understood. We studied needle anatomy, shoot gas exchange, current-year shoot length, and stem diameter growth in Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) seedlings exposed to elevated ozone (1.35× to 1.5× ambient concentration) and elevated temperature (0.9–1.3 °C + ambient temperature) alone and in combination for two exposure seasons in two separate open-field experiments in central Finland. Pines grew also at two soil nitrogen levels. In spruce, warming increased mesophyll intercellular space and reduced gas exchange and shoot growth and made needles narrower and the epidermis and hypodermis thinner. In pine, warming made needles bigger, increased shoot and stem growth, stomatal row number, and proportions of vascular cylinder, phloem, and xylem and reduced the proportion of mesophyll. These responses indicate that pine benefited and spruce suffered from moderate warming. Ozone caused a thickening of epi- and hypo-dermis and a lower stomatal conductance in both species, reduced stomatal density in spruce, and increased proportions of phloem, xylem, and sclerenchyma and reduced growth in pine. Ozone responses suggest increased oxidative stress defense. Stomatal responses were affected by interactions of elevated temperature and ozone in both species. Nitrogen availability modified ozone and temperature responses, particularly in the vascular tissues in pine., final draft, peerReviewed