Tritrophic Interactions among Arthropod Natural Enemies, Herbivores and Plants Considering Volatile Blends at Different Scale Levels.

Herbivore-induced plant volatiles (HIPVs) are released by plants upon damaged or disturbance by phytophagous insects. Plants emit HIPV signals not merely in reaction to tissue damage, but also in response to herbivore salivary secretions, oviposition, and excrement. Although certain volatile chemicals are retained in plant tissues and released rapidly upon damaged, others are synthesized de novo in response to herbivore feeding and emitted not only from damaged tissue but also from nearby by undamaged leaves. HIPVs can be used by predators and parasitoids to locate herbivores at different spatial scales. The HIPV-emitting spatial pattern is dynamic and heterogeneous in nature and influenced by the concentration, chemical makeup, breakdown of the emitted mixes and environmental elements (e.g., turbulence, wind and vegetation) which affect the foraging of biocontrol agents. In addition, sensory capability to detect volatiles and the physical ability to move towards the source were also different between natural enemy individuals. The impacts of HIPVs on arthropod natural enemies have been partially studied at spatial scales, that is why the functions of HIPVs is still subject under much debate. In this review, we summarized the current knowledge and loopholes regarding the role of HIPVs in tritrophic interactions at multiple scale levels. Therefore, we contend that closing these loopholes will make it much easier to use HIPVs for sustainable pest management in agriculture.

Alteration in Light Spectra Causes Opposite Responses in Volatile Phenylpropanoids and Terpenoids Compared with Phenolic Acids in Sweet Basil (Ocimum basilicum) Leaves.

Basil (Ocimum basilicum, cv. Dolly) grew under three different light spectra (A, B, and C) created by light-emitting diode lamps. The proportions of UV-A, blue, and green-yellow wavelengths decreased linearly from A to C, and the proportions of red and far-red wavelengths increased from A to C. Photosynthetic photon flux density was 300 µmol m-2 s-1 in all spectra. The spectrum C plants had highest concentrations of phenolic acids (main compounds: rosmarinic acid and cichoric acid), lowest concentrations and emissions of phenylpropanoid eugenol and terpenoids (main compounds: linalool and 1,8-cineole), highest dry weight, and lowest water content. Conversely, spectra A and B caused higher terpenoid and eugenol concentrations and emissions and lower concentrations of phenolic acids. High density of peltate glandular trichomes explained high terpenoid and eugenol concentrations and emissions. Basil growth and secondary compounds affecting aroma and taste can be modified by altering light spectra; however, increasing terpenoids and phenylpropanoids decreases phenolic acids and growth and vice versa.

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.

Seasonal Volatile Emission Patterns of the Endemic New Zealand Shrub Dracophyllum subulatum on the North Island Central Plateau.

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.

The phytotoxic air-pollutant O3 enhances the emission of herbivore-induced volatile organic compounds (VOCs) and affects the susceptibility of black mustard plants to pest attack.

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.

Effects of Two Invasive Weeds on Arthropod Community Structure on the Central Plateau of New Zealand.

Heather (Calluna vulgaris) and broom (Cytisus scoparius), originally from Europe, are the main invasive plants on New Zealand'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 manuka (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's index and Simpson'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, manuka, 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.

Seasonal and environmental variation in volatile emissions of the New Zealand native plant Leptospermum scoparium in weed-invaded and non-invaded sites.

The New Zealand tea tree Leptospermun scoparium (manuka) is widely known for the antimicrobial properties of its honey. Manuka 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 manuka 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 manuka; (2) manuka and another native species (Dracophyllum subulatum); and manuka with one of two European invasive plants, (3) heather or (4) Scotch broom. We also quantified herbivore damage on target manuka 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 manuka 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.

Herbivory and Attenuated UV Radiation Affect Volatile Emissions of the Invasive Weed Calluna vulgaris.

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 "adjustment" 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's abundance, feeding stage and plant phenology. Heather plants under 95% attenuated UV produced significantly higher amounts of (E)-ß-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.

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.

Natural Variation in Volatile Emissions of the Invasive Weed Calluna vulgaris in New Zealand.

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's foliar volatiles influence native species.

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.

Covariation and phenotypic integration in chemical communication displays: biosynthetic constraints and eco-evolutionary implications.

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.

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.