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.

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

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

Pre-exposure to nitric oxide modulates the effect of ozone on oxidative defenses and volatile emissions in lima bean.

The roles that ozone and nitric oxide (NO), the chief O3 precursor, play in the antioxidative balance and inducible volatile emissions of lima bean were assessed. Exposure to O3 inhibited APX, CAT, and GR, decreased GSH content and induced emissions of (E)-ß-ocimene, limonene, 1,8-cineole, linalool, (E)-4,8-dimethyl-1,3,7-nonatriene (E)-DMNT, 2-butanone and nonanal. O3 did not induce emissions of (E)-ß-caryophyllene and appeared to reduce the antioxidative capacity of plants to a greater extent than NO and NO followed by O3 (NO/O3) treatments. There were significant differences in emissions of (E)-ß-ocimene and linalool between NO/O3 treated plants and controls, but no differences in antioxidant concentrations. A model to explain the relationships between the ascorbate-glutathione cycle and O3 and NO inducible volatiles was proposed. Our findings suggest that prior exposure to NO modulates the oxidative effect of ozone by the process of cross-tolerance, which might regulate the antioxidative system and induction of volatile organic compounds.

Molecular plant volatile communication.

Plants produce a wide array of volatile organic compounds (VOCs) which have multiple functions as internal plant hormones (e.g., ethylene, methyl jasmonate and methyl salicylate), in communication with conspecific and heterospecific plants and in communication with organisms of second (herbivores and pollinators) and third (enemies of herbivores) trophic levels. Species specific VOCs normally repel polyphagous herbivores and those specialised on other plant species, but may attract specialist herbivores and their natural enemies, which use VOCs as host location cues. Attraction of predators and parasitoids by VOCs is considered an evolved indirect defence, whereby plants are able to indirectly reduce biotic stress caused by damaging herbivores. In this chapter we review these interactions where VOCs are known to play a crucial role. We then discuss the importance of volatile communication in self and nonself detection. VOCs are suggested to appear in soil ecosystems where distinction of own roots from neighbours roots is essential to optimise root growth, but limited evidence of above-ground plant self-recognition is available.

Birch (Betula spp.) leaves adsorb and re-release volatiles specific to neighbouring plants--a mechanism for associational herbivore resistance?

Plant-emitted semi-volatile compounds have low vaporization rates at 20-25 degrees C and may therefore persist on surfaces such as plant foliage. The passive adsorption of arthropod-repellent semi-volatiles to neighbouring foliage could convey associational resistance, whereby a plant's neighbours reduce damage caused by herbivores. We found that birch (Betula spp.) leaves adsorb and re-release the specific arthropod-repelling C(15) semi-volatiles ledene, ledol and palustrol produced by Rhododendron tomentosum when grown in mixed association in a field setup. In a natural habitat, a higher concentration of ledene was released from birches neighbouring R. tomentosum than from birches situated > 5 m from R. tomentosum. Emission of alpha-humulene, a sesquiterpene synthesized by both Betula pendula and R. tomentosum, was also increased in R. tomentosum-neighbouring B. pendula. In assessments for associational resistance, we found that the polyphagous green leaf weevils (Polydrusus flavipes) and autumnal moth (Epirrita autumnata) larvae both preferred B. pendula to R. tomentosum. P. flavipes also preferred birch leaves not exposed to R. tomentosum to leaves from mixed associations. In the field, a reduction in Euceraphis betulae aphid density occurred in mixed associations. Our results suggest that plant/tree species may be protected by semi-volatile compounds emitted by a more herbivore-resistant heterospecific neighbour.

Pine weevil feeding on Norway spruce bark has a stronger impact on needle VOC emissions than enhanced ultraviolet-B radiation.

Plants can respond physiologically to damaging ultraviolet-B radiation by altering leaf chemistry, especially UV absorbing phenolic compounds. However, the effects on terpene emissions have received little attention. We conducted two field trials in plots with supplemented UV-B radiation and assessed the influence of feeding by pine weevils, Hylobius abietis L., on volatile emissions from 3-year old Norway spruce trees (Picea abies L. Karst.). We collected emissions from branch tips distal to the feeding weevils, and from whole branches including the damage sites. Weevil feeding clearly induced the emission of monoterpenes and sesquiterpenes, particularly linalool and (E)-beta-farnesene, from branch tips, and the sums of monoterpenes and sesquiterpenes emitted by whole branches were substantially increased. We discovered little effect of UV-B radiation up to 30% above the ambient level on volatile emissions from branch tips distal to damage sites, but there was a possible effect on bark emissions from damage sites.

Isoprene emission from a subarctic peatland under enhanced UV-B radiation.

Isoprene is a reactive hydrocarbon with an important role in atmospheric chemistry, and emissions from vegetation contribute to atmospheric carbon fluxes. The magnitude of isoprene emissions from arctic peatlands is not known, and it may be altered by increasing UV-B radiation. Isoprene emission was measured with the dynamic chamber method from a subarctic peatland under long-term enhancement of UV-B radiation targeted to correspond to a 20% loss in the stratospheric ozone layer. The site type of the peatland was a flark fen dominated by the moss Warnstorfia exannulata and sedges Eriophorum russeolum and Carex limosa. The relationship between species densities and the emission was also assessed. Isoprene emissions were significantly increased by enhanced UV-B radiation during the second (2004) and the fourth (2006) growing seasons under the UV-B exposure. Emissions were related to the density of E. russeolum. The dominant moss, W. exannulata, proved to emit small amounts of isoprene in a laboratory trial. Subarctic fens, even without Sphagnum moss, are a significant source of isoprene to the atmosphere, especially under periods of warm weather. Warming of the Arctic together with enhanced UV-B radiation may substantially increase the emissions.

Long-term exposure to enhanced UV-B radiation has no significant effects on growth or secondary compounds of outdoor-grown Scots pine and Norway spruce seedlings.

The effects of long-term enhanced UV-B radiation on growth and secondary compounds of two conifer species were studied in an outdoor experiment. Scots pine (Pinus sylvestris) seedlings were exposed for two growing seasons and Norway spruce (Picea abies) seedlings for three growing seasons to supplemental UV-B radiation, corresponding to a 30% increase in ambient UV-B radiation. The experiment also included appropriate controls for ambient and increased UV-A radiation. Enhanced UV-B did not affect the growth of the conifer seedlings. In addition, neither the concentrations of terpenes and phenolics in the needles nor the concentrations of terpenes in the wood were affected. However, in the UV-A control treatment the concentrations of diterpenes in the wood of Scots pine decreased significantly compared to the ambient control. Apparently, a small increase in UV-B radiation has no significant effects on the secondary compounds and growth of Scots pine and Norway spruce seedlings.

Influence of carrot psyllid (Trioza apicalis) feeding or exogenous limonene or methyl jasmonate treatment on composition of carrot (Daucus carota) leaf essential oil and headspace volatiles.

The effect of carrot psyllid (Trioza apicalis Förster) feeding and limonene and methyl jasmonate (MeJA) treatments on the essential oil composition and headspace volatiles of carrot (Daucus carota ssp. sativus), cvs. Parano and Splendid, leaves was studied. Carrot psyllid feeding significantly increased the concentrations of sabinene, beta-pinene, and limonene, whereas limonene treatment increased the concentration of (Z)-beta-ocimene in the leaves of both cultivars. The limonene treatment significantly increased the concentration of total phenolics in the leaves of both cultivars, and MeJA treatment increased phenolic concentration in the leaves of Parano. Exogenous limonene spray did not decrease the number of carrot psyllid eggs laid either 2 or 24 h after treatment. The results suggest that carrot psyllid feeding induces changes in the endogenous monoterpene pool in the carrot leaves. Limonene and MeJA treatments affect some induced defenses of the carrot, but the exogenous limonene spray is not an effective oviposition deterrent against carrot psyllid.

Resource partitioning to growth, storage and defence in nitrogen-fertilized Scots pine and susceptibility of the seedlings to the tarnished plant bug Lygus rugulipennis.

We tested how variable nitrogen availability affects the above-and below-ground growth of first-year Scots pine (Pinus sylvestris L.) seedlings and carbon allocation to defensive allelochemicals and storage. Concentrations of free ammo acids were considered as indicators of nutritive quality. Suitability of seedlings for polyphagous Lygus rugulipennis Popp. (Heteroptera: Miridae) was tested with oviposition preference and nymphal growth experiments. At the end of the growing season, needle length increased while root hiomass decreased with elevated N fertilization, but shoot length was not affected. Concentration of starch in needles and roots, representing carbon storage, was not significantly affected by N fertilization, although there was a decrease in the starch concentration of needles when nitrogen input increased. Nitrogen fertilization significantly increased the pool of total and non-essential ammo acids in the shoots. Arginine, proline and glutamine were the individual ammo acids most affected by elevated N. Of the carbon-based defence compounds, total resin acid concentrations in shoots were significantly reduced with elevated nitrogen in 9-wk-old seedlings. Palustric acid and neoabietic acid were the most affected individual) resin acids, where as foliar monoterpenes were not influenced by N availability. Total phenolics in shoot and root showed variable response. The increasing effect of nitrogen on the oviposition rate of Lygus females was almost linear. Mean relative growth rate of the nymphs was significantly affected by the level of N fertilization. but the mortality of nymphs was high in all treatments. The results suggest that in nitrogen-rich environments the needle growth of small Scots pine seedlings is improved, but their Susceptibility to insect attack is increased and they remain less defended as predicted by the carbon/nutrient balance hypothesis. Total phenolics and resin acids, representing phenylalanine and mevalonic acid pathways, respectively, were both reduced by increased nitrogen availability. Together with the simultaneous increase of foliar free nitrogen in the form amino acids, the nutritive value of seedlings is ameliorated and this might explain susceptibility of nursery-grown. N fertilized seedlings to polyphagous Lygus bugs.