Characterisation of New Zealand Propolis from Different Regions Based on Its Volatile Organic Compounds.

Propolis is a bee product mainly consisting of plant resins and is used by bees to maintain the structural integrity of the colony. Propolis is known to contribute to bee health via its antimicrobial activity and is a valued product for human use owing to its nutritional and medicinal properties. Propolis is often characterised into seven categories depending on the resin source. New Zealand propolis is typically assumed as being poplar-type propolis, but few studies have chemically characterised New Zealand propolis to confirm or reject this assumption. Here, for the first time, we characterise propolis originating from different regions in New Zealand based on its volatile organic compounds, using gas chromatography coupled with mass spectrometry (GC-MS). To support this characterisation, we also collected and analysed resin samples from a variety of resin-producing plants (both native to New Zealand and introduced). Our findings suggest that bees mainly use poplar as a resin source, but also utilize native plant species to produce propolis. While regional variation did not allow for clear separation between samples, some patterns emerged, with samples from some regions having more chemical complexity and a higher contribution from native species (as suggested by a higher number of compounds unique to native species resin). Further studies are needed to accurately identify the botanical sources contributing to these samples. It may be also of interest to explore the biological activity of regional propolis samples and their potential nutritional or medicinal benefits.

Environmental Factors Affecting Monoterpene Emissions from Terrestrial Vegetation.

Monoterpenes are volatile organic compounds that play important roles in atmospheric chemistry, plant physiology, communication, and defense. This review compiles the monoterpene emission flux data reported for different regions and plant species and highlights the role of abiotic environmental factors in controlling the emissions of biogenic monoterpenes and their emission fluxes for terrestrial plant species (including seasonal variations). Previous studies have demonstrated the role and importance of ambient air temperature and light in controlling monoterpene emissions, likely contributing to higher monoterpene emissions during the summer season in temperate regions. In addition to light and temperature dependence, other important environmental variables such as carbon dioxide (CO2), ozone (O3), soil moisture, and nutrient availability are also known to influence monoterpene emissions rates, but the information available is still limited. Throughout the paper, we identify knowledge gaps and provide recommendations for future studies.

A mini-review on the impact of common gorse in its introduced ranges.

It is indisputable that invasive plant species strongly impact the ecosystems they invade. Many of such impacts can be negative and threaten the local species through competition, environmental change, or habitat loss. However, introduced plants may also have positive roles in the ecosystems they invade. This review extracted information from reports on common gorse (Ulex europaeus), one of the top 100 invasive plants on the earth, including its detrimental effects and potential beneficial roles in invaded ecosystems. The reduction of native fauna and flora are the main harmful effects of common gorse identified by the literature review. Soil impoverishment and fire hazards are other negative impacts reported for common gorse that could affect agricultural systems and local economies. Despite the negative impacts, reports of positive ecological services provided by common gorse also exist, e.g., as a nursery plant or habitat for endangered native animals. We also reviewed the known human uses of this plant that could support management strategies through harvest and benefit the local communities, including its use as biofuel, raw matter for xylan extraction, medicine, and food. Finally, our review identified the gaps in the literature regarding the understanding of the beneficial role of common gorse on native ecosystems and potential human uses, especially in the tropics.

Exploring plant volatile-mediated interactions between native and introduced plants and insects.

In invasion scenarios, native and introduced species co-occur creating new interactions and modifying existing ones. Many plant-plant and plant-insect interactions are mediated by volatile organic compounds (VOCs), however, these have seldom been studied in an invasion context. To fill this knowledge gap, we explored some interactions mediated by VOCs between native and introduced plants and insects in a New Zealand system. We investigated whether a native plant, Leptospermum scoparium (manuka), changes its volatile profile when grown adjacent to two European introduced plants, Calluna vulgaris (heather) and Cytisus scoparius (Scotch broom), in a semi-field trial using potted plants without above- or below-ground physical contact. We also investigated the influence of plant cues on the host-searching behaviour of two beetles, the native Pyronota festiva (manuka beetle), and the introduced biocontrol agent Lochmaea suturalis (heather beetle), by offering them their host-plant and non-host volatiles versus clean air, and their combination in a Y-tube olfactometer. As a follow-up, we performed preference/feeding tests in Petri dishes with fresh plant material. Results of the semi-field experiment show a significant reduction in green leaf volatiles, sesquiterpenes and total volatile emissions by manuka plants neighbouring heather. In the Y-tube assays, the native beetle P. festiva performed poorly in discriminating between host and non-host plants based on plant volatile cues only. However, it performed relatively well in the Petri dish tests, where other cues (i.e., visual, gustatory or tactile) were present. In contrast, the introduced beetle L. suturalis showed high host-specificity in both Y-tube and Petri dish assays. This study illustrates the importance of VOCs in mediating interactions between introduced and native species, suggesting that invasive plants can disrupt native plants' communication and affect the host-searching behaviour of native insects. It also reinforces the relevance of regular host testing on introduced weed biocontrol agents to avoid unwanted host shifts or host-range expansion.

Exploring the Chemical Properties and Biological Activity of Four New Zealand Monofloral Honeys to Support the Maori Vision and Aspirations.

Honey production and export are significant contributors to the Aotearoa New Zealand economy, generating over 400 million dollars in revenue. Its main export is manuka (Leptospermum scoparium) honey, which has a high commercial value due to its medicinal properties that are linked to its unique chemical composition. The compound methylglyoxal (MGO) has been identified as the main floral marker and is used as a quality indicator, often labelled as unique manuka factor (UMF). However, the high demand for manuka honey creates pressure on beekeepers and may have negative ecological consequences by favouring extensive manuka monocultures to the detriment of other native species. There are other honeys native to New Zealand, such as kamahi (Weinmannia racemosa), kanuka (Kunzea ericoides), rata (Metrosideros robusta) and rewarewa (Knightia excelsa), that also have medicinal properties; however, they are less well known in the local and global market. Indigenous Maori communities envision the production and commercialization (locally and internationally) of these honeys as an opportunity to generate income and secure a sustainable future in alignment with their worldview (Te Ao Maori) and values (tikanga Maori). Diversifying the market could lead to a more sustainable income for beekeepers and reduce pressure on Maori and the conservation land, while supporting indigenous communities to realize their vision and aspirations. This manuscript provides an extensive review of the scientific literature, technical literature and traditional knowledge databases describing the plants of interest and their traditional medicinal uses (rongoa) and the chemical properties of each honey, potential floral markers and their biological activity. For each honey type, we also identify knowledge gaps and potential research avenues. This information will assist Maori beekeepers, researchers, consumers and other stakeholders in making informed decisions regarding future research and the production, marketing and consumption of these native monofloral honeys.

Chemical Ecology and Olfaction in Short-Horned Grasshoppers (Orthoptera: Acrididae).

Chemoreception plays a crucial role in the reproduction and survival of insects, which often rely on their sense of smell and taste to find partners, suitable habitats, and food sources, and to avoid predators and noxious substances. There is a substantial body of work investigating the chemoreception and chemical ecology of Diptera (flies) and Lepidoptera (moths and butterflies); but less is known about the Orthoptera (grasshoppers, locusts, crickets, and weta). Within the Orthoptera, the family Acrididae contains about 6700 species of short-horned grasshoppers. Grasshoppers are fascinating organisms to study due to their significant taxonomic and ecological divergence, however, most chemoreception and chemical ecology studies have focused on locusts because they are agricultural pests (e.g., Schistocerca gregaria and Locusta migratoria). Here we review studies of chemosensory systems and chemical ecology of all short-horned grasshoppers. Applications of genome editing tools and entomopathogenic microorganism to control locusts in association with their chemical ecology are also discussed. Finally, we identify gaps in the current knowledge and suggest topics of interest for future studies.

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.

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.

Honeydew Deposition by the Giant Willow Aphid (Tuberolachnus salignus) Affects Soil Biota and Soil Biochemical Properties.

Infestation of willow plants by the giant willow aphid Tuberolachnus salignus (Hemiptera: Aphididae) is associated with copious deposition of sugar-rich honeydew under the plant canopy. We explored the effect of aphid honeydew on the soil biota and biochemical indicators in a two-year field trial. Soil samples from under aphid-infested and control willow trees, as well as samples from black sooty mould spots under the aphid-infested willows were compared; soil samples before aphid inoculation were used as a baseline. The honeydew deposition had a positive effect on the total soil carbon (C), but not on the total soil nitrogen content or soil pH. Microbial biomass C, basal respiration, number of yeast colony forming units, and the geometric mean of activities for six enzymes were significantly higher in honeydew-affected soils than in the control treatment on both years. The honeydew deposition also increased soil meso-fauna abundance, especially in the black sooty mould spots. The soil biochemical properties, which differed before and after aphid infestation, showed considerable overlap between the first and second year post-infestation. The results highlight the cascading effects of T. salignus on soil biological activity and the importance of using a multitrophic approach to explore similar scenarios.

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.

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.

Comparative Seed Morphology of Tropical and Temperate Orchid Species with Different Growth Habits.

Seed morphology underpins many critical biological and ecological processes, such as seed dormancy and germination, dispersal, and persistence. It is also a valuable taxonomic trait that can provide information about plant evolution and adaptations to different ecological niches. This study characterised and compared various seed morphological traits, i.e., seed and pod shape, seed colour and size, embryo size, and air volume for six orchid species; and explored whether taxonomy, biogeographical origin, or growth habit are important determinants of seed morphology. We investigated this on two tropical epiphytic orchid species from Indonesia (Dendrobium strebloceras and D. lineale), and four temperate species from New Zealand, terrestrial Gastrodia cunnninghamii, Pterostylis banksii and Thelymitra nervosa, and epiphytic D. cunninghamii. Our results show some similarities among related species in their pod shape and colour, and seed colouration. All the species studied have scobiform or fusiform seeds and prolate-spheroid embryos. Specifically, D. strebloceras, G. cunninghamii, and P. banksii have an elongated seed shape, while T. nervosa has truncated seeds. Interestingly, we observed high variability in the micro-morphological seed characteristics of these orchid species, unrelated to their taxonomy, biogeographical origin, or growth habit, suggesting different ecological adaptations possibly reflecting their modes of dispersal.

Quality Improvement Basics: A Crash Course for Pediatric Cardiac Care.

BACKGROUND: Lack of knowledge of quality improvement (QI) methodology and change management principles can explain many of the difficulties encountered when trying to develop effective QI initiatives in health care. METHODS: An interactive QI workshop at the 14th Annual Meeting of the Pediatric Cardiac Intensive Care Society provided an overview of the role of QI in health care, basic QI frameworks and tools, and leadership and organizational culture pitfalls. The top five QI projects submitted to the meeting were later presented to an expert QI panel in a separate session to illustrate examples of QI principles. RESULTS: Workshop presenters introduced two major QI methodologies used to design QI projects. Important first steps include identifying a problem, forming a multidisciplinary team, and developing an aim statement. Key driver diagrams were highlighted as an important tool to develop a project's framework. Several diagnostic tools used to understand the problem were discussed, including the "5 Why's," cause-and-effect charts, and process flowcharts. The importance of outcome, process, and balancing measures was emphasized. Identification of interventions, the value of plan-do-study-act cycles to fuel continuous QI, and use of statistical process control, including run charts or control charts, were reviewed. The importance of stakeholder engagement, transparency, and sustainability was discussed. Later, the top five QI projects presented highlighted multiple "QI done well" practices discussed during the preconference QI workshop. CONCLUSIONS: Understanding QI methodology and appropriately applying basic QI tools are pivotal steps to realizing meaningful and sustained improvement.

Parthenogenetic Females of the Stick Insect Clitarchus hookeri Maintain Sexual Traits.

The New Zealand stick insect Clitarchus hookeri has both sexual and parthenogenetic (all-female) populations. Sexual populations exhibit a scramble competition mating system with distinctive sex roles, where females are signalers and males are searchers, which may lead to differences in the chemical and morphological traits between sexes. Evidence from a range of insect species has shown a decay of sexual traits is common in parthenogenetic lineages, especially those traits related to mate attraction and location, presumably due to their high cost. However, in some cases, sexual traits remain functional, either due to the recent evolution of the parthenogenetic lineage, low cost of maintenance, or because there might be an advantage in maintaining them. We measured morphological and chemical traits of C. hookeri to identify differences between males and females and between females from sexual and parthenogenetic populations. We also tested the ability of males to discriminate between sexual and parthenogenetic females in a laboratory bioassay. Our results show that male C. hookeri has morphological traits that facilitate mobility (smaller body with disproportionately longer legs) and mate detection (disproportionately longer antennae), and adult females release significantly higher amounts of volatile organic compounds than males when this species is sexually active, in accordance with their distinctive sex roles. Although some differences were detected between sexual and parthenogenetic females, the latter appear to maintain copulatory behaviors and chemical signaling. Males were unable to distinguish between sexual and parthenogenetic females, suggesting that there has been little decay in the sexual traits in the parthenogenetic lineage of C. hookeri.

Divergent behavioural responses of gypsy moth (Lymantria dispar) caterpillars from three different subspecies to potential host trees.

Almost all previous work on host-plant selection by insect herbivores has focused on adult behaviour; however, immature life stages can also play an active role in host discrimination. The important forest pest Lymantria dispar (gypsy moth) has three recognised subspecies: the European, Asian, and Japanese gypsy moth. Unlike the other two subspecies, the European subspecies is characterised by a loss of female flight ability, which might impose a selective pressure on larvae to actively engage in host-plant selection. We therefore explored the interactions of early-instar larvae from laboratory colonies of each subspecies with four potential hosts of differing quality: oak, beech, maple, and pine-measuring larval survival and performance, feeding preferences, responses to host-derived odour cues, and the propensity to disperse from hosts via ballooning. Compared to larvae from the Asian and Japanese subspecies, larvae from the (American-originated) European gypsy moth colony exhibited (i) significantly lower survival on the poorest quality host (pine), (ii) an ability to discriminate among hosts via olfactory cues; and (iii) higher propensity to disperse from sub-optimal hosts. These results are consistent with the hypothesis that larvae from flightless female European Gypsy moth subspecies play a more active role in host-plant selection.

Herbivore-induced volatile emission from old-growth black poplar trees under field conditions.

Herbivory is well known to trigger increased emission of volatile organic compounds (VOCs) from plants, but we know little about the responses of mature trees. We measured the volatiles emitted by leaves of old-growth black poplar (Populus nigra) trees after experimental damage by gypsy moth (Lymantria dispar) caterpillars in a floodplain forest, and studied the effect of herbivory on the transcript abundance of two genes involved in the biosynthesis of VOCs, and the accumulation of defence phytohormones. Herbivory significantly increased volatile emission from the experimentally damaged foliage, but not from adjacent undamaged leaves in the damaged branches (i.e., no systemic response). Methylbutyraldoximes, 4,8-dimethyl-1,3,7-nonatriene (DMNT), (Z)-3-hexenol and (E)-ß-ocimene, amongst other compounds, were found to be important in distinguishing the blend of herbivore-damaged vs. undamaged leaves. Herbivory also increased expression of PnTPS3 (described here for the first time) and PnCYP79D6-v4 genes at the damaged sites, these genes encode for an (E)-ß-ocimene synthase and a P450 enzyme involved in aldoxime formation, respectively, demonstrating de novo biosynthesis of the volatiles produced. Herbivore-damaged leaves had significantly higher levels of jasmonic acid and its conjugate (-)-jasmonic acid-isoleucine. This study shows that mature trees in the field have a robust response to herbivory, producing induced volatiles at the damaged sites even after previous natural herbivory and under changing environmental conditions, however, further studies are needed to establish whether the observed absence of systemic responses is typical of mature poplar trees or if specific conditions are required for their induction.

Exploring the Effects of Plant Odors, from Tree Species of Differing Host Quality, on the Response of Lymantria dispar Males to Female Sex Pheromones.

A widely accepted hypothesis for host-plant selection in herbivorous insects is that ovipositing females select host-plants that maximize the survival and performance of their offspring. However, numerous studies indicate that this is not always the case for polyphagous species. Lymantria dispar is a highly polyphagous forest defoliator and has flightless females in some subspecies, resulting in a limited capacity to make host-choices. Males of other Lepidopteran species utilize a combination of sexual pheromones and plant volatiles in their mating choices and exhibit preferences among plant species. We explored the behavior of L. dispar males towards sexual pheromone in the presence and absence of plant volatiles and their ability to discriminate between two plant species with different degrees of suitability for their offspring: a suboptimal host (Pinus sylvestris), and an optimal host (Quercus robur). In no-choice wind tunnel assays, we found that rates of male success in locating a pheromone source were not altered by the presence of plant odors; however, the time spent by males searching for the pheromone source after reaching the full length of the tunnel was reduced by more than 50% in the presence of plant volatiles. In dual choice assays, males exhibited a clear preference for a combination of pheromones and plant volatiles over the pheromone alone. However, we did not find evidence of an innate ability to discriminate between the odors of optimal and suboptimal host plants. We discuss possible ecological and evolutionary explanations for these observations.

Can plant-natural enemy communication withstand disruption by biotic and abiotic factors?

The attraction of natural enemies towards herbivore-induced plant volatiles is a well-documented phenomenon. However, the majority of published studies are carried under optimal water and nutrient regimes and with just one herbivore. But what happens when additional levels of ecological complexity are added? Does the presence of a second herbivore, microorganisms, and abiotic stress interfere with plant-natural enemy communication? or is communication stable enough to withstand disruption by additional biotic and abiotic factors?Investigating the effects of these additional levels of ecological complexity is key to understanding the stability of tritrophic interactions in natural ecosystems and may aid to forecast the impact of environmental disturbances on these, especially in climate change scenarios, which are often associated with modifications in plant and arthropod species distribution and increased levels of abiotic stress.This review explores the literature on natural enemy attraction to herbivore-induced volatiles when, besides herbivory, plants are challenged by additional biotic and abiotic factors.The aim of this review was to establish the impact of different biotic and abiotic factors on plant-natural enemy communication and to highlight critical aspects to guide future research efforts.

Feeding Experience Affects the Behavioral Response of Polyphagous Gypsy Moth Caterpillars to Herbivore-induced Poplar Volatiles.

Plant volatiles influence host selection of herbivorous insects. Since volatiles often vary in space and time, herbivores (especially polyphagous ones) may be able to use these compounds as cues to track variation in host plant quality based on their innate abilities and previous experience. We investigated the behavioral response of naïve (fed on artificial diet) and experienced (fed on poplar) gypsy moth (Lymantria dispar) caterpillars, a polyphagous species, towards constitutive and herbivore-induced black poplar (Populus nigra) volatiles at different stages of herbivore attack. In Y-tube olfactometer assays, both naïve and experienced caterpillars were attracted to constitutive volatiles and volatiles released after short-term herbivory (up to 6 hr). Naïve caterpillars also were attracted to volatiles released after longer-term herbivory (24-30 hr), but experienced caterpillars preferred the odor of undamaged foliage. A multivariate statistical analysis comparing the volatile emission of undamaged plants vs. plants after short and longer-term herbivory, suggested various compounds as being responsible for distinguishing between the odors of these plants. Ten compounds were selected for individual testing of caterpillar behavioral responses in a four-arm olfactometer. Naïve caterpillars spent more time in arms containing (Z)-3-hexenol and (Z)-3-hexenyl acetate than in solvent permeated arms, while avoiding benzyl cyanide and salicyl aldehyde. Experienced caterpillars avoided benzyl cyanide and preferred (Z)-3-hexenyl acetate and the homoterpene (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT) over solvent. Only responses to DMNT were significantly different when comparing experienced and naïve caterpillars. The results show that gypsy moth caterpillars display an innate behavioral response towards constitutive and herbivore-induced plant volatiles, but also that larval behavior is plastic and can be modulated by previous feeding experience.