Plant Volatile Organic Compounds Emissions Research Articles

Individual and Combined Effects of Predatory Bug Engytatus nicotianae and Trichoderma atroviride in Suppressing the Tomato Potato Psyllid Bactericera cockerelli in Greenhouse Grown Tomatoes

The tomato potato psyllid (TPP) Bactericera cockerelli is a serious pest of the Solanaceae family. The management of this pest using synthetic pesticides is problematic because of the development of pesticide resistance and environmental concerns including impacts on non-target organisms. The predatory bug Engytatus nicotianae has recently been identified as a useful biocontrol agent for TPP in greenhouses. The soil fungus Trichoderma Pers. is commonly used as a plant growth enhancer and biocontrol agent against phytopathogenic fungi. Therefore, there could be advantages associated with the combined use of these biocontrol agents. Some reports in other systems suggest that Trichoderma inoculation may alter the behaviour of pests and their natural enemies by modifying plant defence metabolites such as volatile organic compounds (VOCs). For this reason, this study aimed to investigate the individual and combined efficacy of these biocontrol agents (i.e., Trichoderma atroviride and E. nicotianae) against TPP in greenhouse grown tomatoes (Solanum lycopersicum cv. Merlice). To this end, we compared the effect of each biocontrol agent and their combination on TPP abundance across different developmental stages (egg, nymphs, adults) and the number of infested leaves. We also investigated plant VOC emissions under the different treatments. Across all measured TPP stages, the treatments tested (E. nicotianae alone, T. atrovirdae alone, and T. atrovirdae + E. nicotianae) significantly reduced mean TPP counts relative to the control, and no significant differences were observed in VOC emissions among treatments. Overall, T. atrovirdae alone was less effective than E. nicotianae alone and its combination with T. atrovirdae in suppressing TPP populations. However, the combined use of Trichoderma + E. nicotianae did not show significant advantages over the use of E. nicotianae alone in controlling TPP. Therefore, their combined use needs to be further assessed in light of other advantages of Trichoderma to the crop (e.g., growth promotion or pathogen defence).

Volatile responses of dwarf birch to mimicked insect herbivory and experimental warming at two elevations in Greenlandic tundra.

Plants release a complex blend of volatile organic compounds (VOCs) in response to stressors. VOC emissions vary between contrasting environments and increase with insect herbivory and rising temperatures. However, the joint effects of herbivory and warming on plant VOC emissions are understudied, particularly in high latitudes, which are warming fast and facing increasing herbivore pressure. We assessed the individual and combined effects of chemically mimicked insect herbivory, warming, and elevation on dwarf birch (Betula glandulosa) VOC emissions in high-latitude tundra ecosystems in Narsarsuaq, South Greenland. We hypothesized that VOC emissions and compositions would respond synergistically to warming and herbivory, with the magnitude differing between elevations. Warming increased emissions of green leaf volatiles (GLVs) and isoprene. Herbivory increased the homoterpene, (E)-4,8-dimethyl-1,3,7-nonatriene, emissions, and the response was stronger at high elevation. Warming and herbivory had synergistic effects on GLV emissions. Dwarf birch emitted VOCs at similar rates at both elevations, but the VOC blends differed between elevations. Several herbivory-associated VOC groups did not respond to herbivory. Harsher abiotic conditions at high elevations might not limit VOC emissions from dwarf birch, and high-elevation plants might be better at herbivory defense than assumed. The complexity of VOC responses to experimental warming, elevation, and herbivory are challenging our understanding and predictions of future VOC emissions from dwarf birch-dominated ecosystems.

Unearthing the alteration in plant volatiles induced by mycorrhizal fungi: A shield against plant pathogens

Plants produce a large range of structurally varied low molecular weight secondary metabolites, which evaporate, known as volatile organic compounds (VOCs). Several of them are emitted in response to biotic stress as a defensive measure against pathogen attacks. Arbuscular Mycorrhizal Fungi (AMFs) can change the VOC pattern in parts of the plant and may promote plant defense via direct or indirect mechanisms. Mycorrhization of plants positively affects plant immunization along with growth and yield. The presence of AMF may raise the concentration of phenolic compounds and the activity of critical defense-related enzymes. AMF-induced changes in plant chemistry and associated volatile emissions lead to stronger immunity against pathogenic microorganisms. Despite substantial research into the origins of diversity in VOC-mediated plant communication, very little is known about the mechanism of influence of several AMFs on plant VOC emissions and modulation of plant immunization. Moreover, the molecular mechanism for VOC sensing in plants and mycorrhizal association is still unclear. In the present review, we have presented an up-to-date understanding of the cross-talk of AMF and VOC patterns in plants and the subsequent modulation of resistance against microbial pathogens.

Comparative Analysis of Volatiles Emitted from Tomato and Pepper Plants in Response to Infection by Two Whitefly-Transmitted Persistent Viruses.

Simple SummaryViruses transmitted by the whitefly Bemisia tabaci, a cosmopolitan pest that inflicts damage on many agricultural crops, are among the most important plant viruses and cause devastating damage to agriculture worldwide. Begomoviruses and a new polerovirus transmitted by the whitefly cause serious damage to tomato and pepper crops in Israel, respectively. Here, both viruses were studied with regard to their ability to induce the production of volatile organic compounds (VOCs) upon plant infection, which are thought to mediate communication with the environment. The results showed that infection with each virus alone, or infestation with whiteflies, without viruses, induced shared and unique VOC accumulation. The VOCs identified in these unique responses suggest that plants can respond specifically to virus infection or insect infestation, and those VOCs can be used individually or as blends for monitoring and disrupting pest populations.The whitefly Bemisia tabaci is one of the most important agricultural pests due to its extreme invasiveness, insecticide resistance, and ability to transmit hundreds of plant viruses. Among these, Begomoviruses and recombinant whitefly-borne Poleroviruses are transmitted persistently. Several studies have shown that upon infection, plant viruses manipulate plant-emitted volatile organic compounds (VOCs), which have important roles in communication with insects. In this study, we profiled and compared the VOCs emitted by tomato and pepper plant leaves after infection with the Tomato yellow leaf curl virus (TYLCV) (Bogomoviruses) and the newly discovered Pepper whitefly-borne vein yellows virus (PeWBVYV) (Poleroviruses), respectively. The results identified shared emitted VOCs but also uncovered unique VOC signatures for each virus and for whitefly infestation (i.e., without virus infection) independently. The results suggest that plants have general defense responses; however, they are also able to respond individually to infection with specific viruses or infestation with an insect pest. The results are important to enhance our understanding of virus- and insect vector-induced alteration in the emission of plant VOCs. These volatiles can eventually be used for the management of virus diseases/insect vectors by either monitoring or disrupting insect–plant interactions.

Cropping systems alter plant volatile emissions in the field through soil legacy effects

AbstractCrops emit a variety of volatile organic compounds (VOCs) that serve as attractants or repellents for pests and their natural enemies. Crop rotations, off-farm chemical inputs, and mechanical and cultural tactics – collectively called cropping systems – alter soil nutrients, moisture content, and microbial communities, all of which have the potential to alter crop VOC emissions. Soil legacy effects of diversified cropping systems have been shown to enhance crop VOC emissions in greenhouse studies, but how they influence emissions under field conditions remains virtually unknown. To determine the effect of cropping systems on plant VOC emissions in the field, air samples were collected from the headspace of wheat (Triticum aestivum L. Judee) grown in simplified wheat-fallow rotations or diversified wheat-cover crop rotations where cover crops were terminated by grazing cattle. Across two growing seasons, wheat grown in rotation with fallow emitted greater amounts of Z-3-hexenyl acetate and β-ocimene, key attractants for wheat stem sawfly (Cephus cinctus Norton), a major pest of wheat. While overall VOC blends were relatively similar among cropping system during the first growing season, emissions varied substantially in the second year of this study where wheat grown in rotation with cover crops emitted substantially greater quantities of volatile compounds characteristic of abiotic stress. Below-average precipitation in the second growing season, in addition to reduced soil water content in cover crop rotations, suggests that cropping system effects on wheat VOCs may have been driven primarily by water availability, a major factor limiting crop growth in dryland agriculture. While the specific mechanisms driving changes in VOC emissions were not explicitly tested, this work shows that agricultural practices applied in one growing season can differentially influence crop VOC emissions in the next through soil legacy effects, illustrating additional avenues through which cropping systems may be leveraged to enhance pest management.

Herbivory Protection via Volatile Organic Compounds Is Influenced by Maize Genotype, Not Bacillus altitudinis-Enriched Bacterial Communities.

Belowground, plants interact with beneficial soil microbes such as plant growth-promoting rhizobacteria (PGPR). PGPR are rhizosphere bacteria that colonize roots and elicit beneficial effects in plants such as improved plant growth, pathogen resistance, abiotic stress tolerance, and herbivore protection. Treatment of plants with PGPR has been shown to trigger the emission of volatile organic compounds (VOCs). Volatile emissions can also be triggered by herbivory, termed herbivore-induced plant volatiles (HIPV), with important ramifications for chemical-mediated plant and insect interactions. Much of our current understanding of PGPR and herbivore-induced volatiles is based on studies using one plant genotype, yet domestication and modern breeding has led to the development of diverse germplasm with altered phenotypes and chemistry. In this study, we investigated if volatile emissions triggered by PGPR colonization and herbivory varies by maize genotype and microbial community assemblages. Six maize genotypes representing three decades of crop breeding and two heterotic groups were used, with four microbiome treatments: live or sterilized soil, with or without a Bacillus inoculant. Soil sterilization was used to delay microbiome establishment, resulting in low-diversity treatments. At planting, maize seeds were inoculated with PGPR Bacillus altitudinis AP-283 and grown under greenhouse conditions. Four weeks post planting, plants were subjected to feeding by third instar Helicoverpa zea (Lepidoptera: Noctuidae) larvae. Volatiles were collected using solid phase microextraction and analyzed with gas chromatography-mass spectrometry. Illumina NovaSeq 16S rRNA amplicon sequencing was carried out to characterize the rhizosphere microbiome. Maize genotype significantly influenced total volatile emissions, and relative abundance of volatile classes. We did not document a strong influence of microbe treatment on plant VOC emissions. However, inoculating plants with PGPR improved plant growth under sterile conditions. Taken together, our results suggest that genotypic variation is the dominant driver in HIPV composition and individual HIPV abundances, and any bacterial-mediated benefit is genotype and HIPV-specific. Therefore, understanding the interplay of these factors is necessary to fully harness microbially-mediated benefits and improve agricultural sustainability.

Elevated CO2‐mediated plant VOCs change aggravates invasive thrips occurrence by altering their host‐selection behaviour

AbstractGlobally rising atmosphere CO2 has been predicted to affect the emission of plant volatile organic compounds (VOCs) and the interaction with insect herbivores. CO2‐mediated plant VOCs change might contribute to develop effective management strategies for insect pests by using VOCs related biological control methods. In this study, we analysed the effect of elevated CO2 on the host‐selection behaviour of western flower thrips (Frankliniella occidentalis) and studied how they were affected by the release of VOCs from kidney bean plants (Phaseolus vulgaris). A four‐chamber olfactometer was used to quantify the host‐selection of F. occidentalis for P. vulgaris. Elevated CO2 increased F. occidentalis' selection for P. vulgaris wounded by mechanical damage (MD) and thrips infestation (TI) that might via regulating the gene expression of CSP1, CSP1‐q, OBP1 and OBP1‐q in F. occidentalis. Besides, we cultivated kidney beans at ambient CO2 (400 ppm) and elevated CO2 (800 ppm), and quantified the emission of plant VOCs by using GC‐MS. Thirty VOCs belonging to ten chemical groups were identified from P. vulgaris, including aromatic hydrocarbons, ethers, alkanes, cycloalkanes, alcohols, alkenes, aromatic derivatives, phenols, ketones and esters. Furthermore, six VOCs from P. vulgaris were significantly affected by CO2 level, wounding way and the interaction between them, including ethyl benzene, 1,3‐dimethyl benzene, 1,3‐dimethyl‐4‐ethyl benzene, (E,E,E,E)‐squalene, 2,6‐ditert‐butyl‐4‐methyl phenol and dioctyl phthalate. Our study indicates that elevated CO2 might increase the host‐selection of F. occidentalis for wounded P. vulgaris due to the changed plant VOCs.

Mycorrhizae Alter Constitutive and Herbivore-Induced Volatile Emissions by Milkweeds.

Plants use volatile organic compounds (VOCs) to cue natural enemies to their herbivore prey on plants. Simultaneously, herbivores utilize volatile cues to identify appropriate hosts. Despite extensive efforts to understand sources of variation in plant communication by VOCs, we lack an understanding of how ubiquitous belowground mutualists, such as arbuscular mycorrhizal fungi (AMF), influence plant VOC emissions. In a full factorial experiment, we subjected plants of two milkweed (Asclepias) species under three levels of AMF availability to damage by aphids (Aphis nerii). We then measured plant headspace volatiles and chemical defenses (cardenolides) and compared these to VOCs emitted and cardenolides produced by plants without herbivores. We found that AMF have plant species-specific effects on constitutive and aphid-induced VOC emissions. High AMF availability increased emissions of total VOCs, two green leaf volatiles (3-hexenyl acetate and hexyl acetate), and methyl salicylate in A. curassavica, but did not affect emissions in A. incarnata. In contrast, aphids consistently increased emissions of 6-methyl-5-hepten-2-one and benzeneacetaldehyde in both species, independent of AMF availability. Both high AMF availability and aphids alone suppressed emissions of individual terpenes. However, aphid damage on plants under high AMF availability increased, or did not affect, emissions of those terpenes. Lastly, aphid feeding suppressed cardenolide concentrations only in A. curassavica, and AMF did not affect cardenolides in either plant species. Our findings suggest that by altering milkweed VOC profiles, AMF may affect both herbivore performance and natural enemy attraction.

Development of a HS-SPME-GC/MS protocol assisted by chemometric tools to study herbivore-induced volatiles in Myrcia splendens

A headspace solid phase microextraction (HS-SPME) method combined with gas chromatography-mass spectrometry (GC/MS) was developed and optimized for extraction and analysis of volatile organic compounds (VOC) of leaves and galls of Myrcia splendens. Through a process of optimization of main factors affecting HS-SPME efficiency, the coating divivnilbenzene-carboxen-polydimethylsiloxane (DVB/Car/PDMS) was chosen as the optimum extraction phase, not only in terms of extraction efficiency, but also for its broader analyte coverage. Optimum extraction temperature was 30°C, while an extraction time of 15min provided the best compromise between extraction efficiencies of lower and higher molecular weight compounds. The optimized protocol was demonstrated to be capable of sampling plant material with high reproducibility, considering that most classes of analytes met the 20% RSD FDA criterion. The optimized method was employed for the analysis of three classes of M. splendens samples, generating a final list of 65 tentatively identified VOC, including alcohols, aldehydes, esters, ketones, phenol derivatives, as well as mono and sesquiterpenes. Significant differences were evident amongst the volatile profiles obtained from non-galled leaves (NGL) and leaf-folding galls (LFG) of M. splendens. Several differences pertaining to amounts of alcohols and aldehydes were detected between samples, particularly regarding quantities of green leaf volatiles (GLV). Alcohols represented about 14% of compounds detected in gall samples, whereas in non-galled samples, alcohol content was below 5%. Phenolic derived compounds were virtually absent in reference samples, while in non-galled leaves and galls their content ranged around 0.2% and 0.4%, respectively. Likewise, methyl salicylate, a well-known signal of plant distress, amounted for 1.2% of the sample content of galls, whereas it was only present in trace levels in reference samples. Chemometric analysis based on Heatmap associated with Hierarchical Cluster Analysis (HCA) and Principal Component Analysis (PCA) provided a suitable tool to differentiate VOC profiles in vegetal material, and could open new perspectives and opportunities in agricultural and ecological studies for the detection and identification of herbivore-induced plant VOC emissions.

Bidirectional Interaction between Phyllospheric Microbiotas and Plant Volatile Emissions.

Due to their antimicrobial effects and their potential role as carbon sources, plant volatile organic compound (VOC) emissions play significant roles in determining the characteristics of the microbial communities that can establish on plant surfaces. Furthermore, epiphytic microorganisms, including bacteria and fungi, can affect plant VOC emissions in different ways: by producing and emitting their own VOCs, which are added to and mixed with the plant VOC blend; by affecting plant physiology and modifying the production and emission of VOCs; and by metabolizing the VOCs emitted by the plant. The study of the interactions between plant VOC emissions and phyllospheric microbiotas is thus of great interest and deserves more attention.

Secretions from the ventral eversible gland of Spodoptera exigua caterpillars activate defense-related genes and induce emission of volatile organic compounds in tomato, Solanum lycopersicum.

BackgroundPlant induced defense against herbivory are generally associated with metabolic costs that result in the allocation of photosynthates from growth and reproduction to the synthesis of defense compounds. Therefore, it is essential that plants are capable of sensing and differentiating mechanical injury from herbivore injury. Studies have shown that oral secretions (OS) from caterpillars contain elicitors of induced plant responses. However, studies that shows whether these elicitors originated from salivary glands or from other organs associated with feeding, such as the ventral eversible gland (VEG) are limited. Here, we tested the hypothesis that the secretions from the VEG gland of Spodoptera exigua caterpillars contain elicitors that induce plant defenses by regulating the expression of genes involved in the biosynthesis of volatile organic compounds (VOCs) and other defense-related genes. To test this hypothesis, we quantified and compared the activity of defense-related enzymes, transcript levels of defense-related genes and VOC emission in tomato plants damaged by S. exigua caterpillars with the VEG intact (VEGI) versus plants damaged by caterpillars with the VEG ablated (VEGA).ResultsThe quantified defense-related enzymes (i.e. peroxidase, polyphenol oxidase, and lipoxigenase) were expressed in significantly higher amounts in plants damaged by VEGI caterpillars than in plants damaged by VEGA caterpillars. Similarly, the genes that encode for the key enzymes involved in the biosynthesis of jasmonic acid and terpene synthase genes that regulate production of terpene VOCs, were up-regulated in plants damaged by VEGI caterpillars. Moreover, the OS of VEGA caterpillars were less active in inducing the expression of defense genes in tomato plants. Increased emissions of VOCs were detected in the headspace of plants damaged by VEGI caterpillars compared to plants damaged by VEGA caterpillars.ConclusionThese results suggest that the VEG of S. exigua caterpillars contains elicitors of late plant defense signaling in tomato which trigger defense-related enzymatic activity, regulate expression of defense-related genes, and induce emission of plant VOCs. These signaling cascades may have important ramifications for plant-insect and tritrophic interactions.

Different Reactions of Potato Varieties to Infection by Potato Leafroll Virus, and Associated Responses by its Vector, Myzus persicae (Sulzer)

Vector-dependent plant pathogens can alter their hosts such that vector behavior and pathogen spread are affected. For example, Potato leafroll virus (PLRV)-induces changes in volatiles emitted by potato plants (Solanum tuberosum L.) that influence settling behavior by Myzus persicae, a principal vector of the virus. Prior work in this pathosystem has utilized a single potato variety, Russet Burbank, but as is true for other plant responses to biotic stresses, responses may differ among plant genotypes. To examine this, PLRV-induced changes in headspace volatile organic compounds (VOCs) and associated aphid responses to these VOCs were compared among four potato varieties (Chipeta, Desiree, IdaRose, and Russet Burbank). Total headspace VOCs differed among the varieties and were differentially induced by PLRV infection such that the effect of variety, infection, and their interaction was significant; two of the varieties had increased concentrations of headspace VOCs, and two did not. MANOVA for the effect of infection and variety on total VOCs and major VOC classes also was significant. A principal component analysis (PCA) partially separated the VOC profiles from the four varieties. Aphid arrestment differed in response to the VOCs of the four varieties, and was greater on those that were more readily infected by PLRV (Desiree and Russet Burbank) as compared with those that were less readily infected (Chipeta and IdaRose). Aphid responses were not clearly related to specific characteristics of blends, such that total VOCs and composition appear to contribute. The four varieties used in this study have distinct pedigrees representative of different cultivated forms of S. tuberosum. Although cultivated potato varieties have been subjected to genetic manipulation by humans, the differences in PLRV induced changes in VOCs nonetheless indicate the potential for complex effects of PLRV infection on host plant VOC emissions, and vector responses in managed, and natural systems.

Floral scent in a whole‐plant context: moving beyond pollinator attraction

The natural world is awash with odours – blends of volatile organic compounds (VOCs) – whose chemical structures and composition convey information to organisms endowed with the sensory capacity to perceive them. Pheromones and other chemical signals provide specific information that facilitates kin recognition, mate acquisition, social interactions and defence across the tree of life (Meinwald & Eisner 2008). Even more general volatile cues, such as CO2, convey context-dependent information about habitat and host or food quality, illness and microbial decay (Guerenstein & Hildebrand 2007). The terrestrial fossil record teems with evidence of chemical communication and warfare, in the form of animal olfactory appendages (Strausfeld & Hildebrand 1999; Labandeira 2002) and plant essential oil glands (Fahn 2002; Krings, Taylor & Kellogg 2002). This evidence, combined with our knowledge of cycads, water lilies and other ‘living fossils’ (Thien, Azuma & Kawano 2000; Terry et al. 2007), attests to the antiquity of volatile communication between plants, their mutualists and antagonists.

Treatment of wet process hardboard plant VOC emissions by a pilot scale biological system

A pilot scale biological treatment system for air emissions was installed and tested at a forest products plant in western Oregon, USA, which collected and treated gaseous emissions from the hardboard steam press vents on the top of the plant building. This system was installed mainly to demonstrate the effectiveness of biological treatment technologies in removing volatile organic compounds (VOC) and hazardous air pollutants (HAP) from the wet-process hardboard press emissions, and to test the efficiency of the system on fine particles and condensable organics with the presence of a pre-treatment wet dust collector. The bio-oxidation system was comprised of a particle pre-treatment unit Type W Rotoclone (wet hydrocyclone), a biotrickling filter and a biofilter with airflow capacity of up to 4.72 m 3/s. This unit operated at approximately 0.71 m 3/s, which is the optimal flow required for the Rotoclone's throughput, and provided an EBCT (empty bed contact time) of 45 s. Analysis of total VOC measurements from the system indicated removals down to less than 5 ppm in the effluent emissions. Evaluations of opacity reductions also met project objectives with routine outlet measurements of 0–5%, which are in compliance with state regulatory guidelines. Emissions air samples were collected at different locations in the biological system for GC–MS analysis and characterization to identify specific VOCs and their removals.

Dynamics of the enhanced emissions of monoterpenes and methyl salicylate, and decreased uptake of formaldehyde, by Quercus ilex leaves after application of jasmonic acid

Jasmonic acid (JA) is a signalling compound with a key role in both stress and development in plants, and is reported to elicit the emission of volatile organic compounds (VOCs). Here we studied the dynamics of such emissions and the linkage with photosynthetic rates and stomatal conductance. We sprayed JA on leaves of the Mediterranean tree species Quercus ilex and measured the photosynthetic rates, stomatal conductances, and emissions and uptake of VOCs using proton transfer reaction mass spectrometry and gas chromatography after a dark-light transition. Jasmonic acid treatment delayed the induction of photosynthesis and stomatal conductance by approx. 20 min, and decreased them 24 h after spraying. Indications were found of both stomatal and nonstomatal limitations of photosynthesis. Monoterpene emissions were enhanced (20-30%) after JA spraying. Jasmonic acid also increased methyl salicylate (MeSa) emissions (more than twofold) 1 h after treatment, although after 24 h this effect had disappeared. Formaldehyde foliar uptake decreased significantly 24 h after JA treatment. Both biotic and abiotic stresses can thus affect plant VOC emissions through their strong impact on JA levels. Jasmonic acid-mediated increases in monoterpene and MeSa emissions might have a protective role when confronting biotic and abiotic stresses.

Headspace solid-phase microextraction--comprehensive two-dimensional gas chromatography of wound induced plant volatile organic compound emissions.

Plant emissions of volatile organic compounds from mechanically wounded Agrostis stolonifera, Pennisetum clandestinum, Eucalyptus leucoxylon and Trifolium repens have been sampled by headspace-solid phase microextraction (HS-SPME) and analysed by using comprehensive two-dimensional gas chromatography (GCxGC) for measurement of the plant emissions. GCxGC produces a fingerprint of the volatile organic compounds in a 2D separation space that may be approximately interpreted as a boiling point-polarity space, and may then be presented as a two-dimensional contour plot. This allows identification of sample-dependent variations in component distributions in the 2D plot, which will contain information about plant differences and should therefore facilitate recognition of different plant materials and displays the gross differences in volatiles between each plant species.