Volatile Organic Compounds Mediate Host Selection of Wheat Midge, Sitodiplosis Mosellana (Géhin) (Diptera: Cecidomyiidae) between Preanthesis and Postanthesis Stages of Wheat.

The orange wheat blossom midge, Sitodiplosis mosellana (Géhin) (Diptera: Cecidomyiidae), is a significant wheat pest in the Prairie Provinces of Canada and northern regions of the USA. Wheat phenology plays a critical role in wheat midge oviposition. We hypothesized that S. mosellana oviposition behaviour is influenced by volatile organic compounds (VOCs) emitted by wheat at two adjacent wheat growth stages: preanthesis and postanthesis. A higher number of S. mosellana eggs laid on preanthesis than postanthesis spikes in an oviposition choice experiment using the susceptible spring wheat cultivar 'Roblin'. In preanthesis, wheat emitted higher amounts of Z-3-hexenyl acetate (Z3-06:OAc) than at the postanthesis stage. Higher amounts of methyl ketones such as 2-tridecanone, 2-pentadecanone, and 2-undecanone were emitted by wheat in the postanthesis stage and these VOCs were sensitive to S. mosellana antennae used in the Gas Chromatography-Electroantennographic Detection. Females were attracted to synthetic Z3-06:OAc but were deterred by 2-tridecanone relative to the solvent control in the vertical Y-tube olfactometer. 2-Undecanone and 2-pentadecanone did not show any attractiveness or deterrence. In a no-choice oviposition experiment, fewer eggs were laid in preanthesis wheat exposed to a synthetic VOC blend of Z3-06:OAc, 2-undecanone, 2-tridecanone, and 2-pentadecanone at the concentrations released by postanthesis spikes. This study shows that the reduction of Z3-06:OAc, in the VOC mix, and possibly the increase in 2-tridecanone, are likely responsible for the reduction in oviposition on postanthesis wheat. These results elucidate for the first time the role of specific VOCs mediating S. mosellana oviposition in preanthesis and postanthesis wheat.

Pheromone and Host Plant Odor Detection in Eastern Spruce Budworm, Choristoneura fumiferana Clemens (Lepidoptera: Tortricidae).

Spruce budworm, Choristoneura fumiferana Clemens, is an ecologically significant defoliator of spruce and balsam fir in North America. Optimization of semiochemical-mediated control is needed to improve the existing integrated pest management systems such as mating disruption and population estimation. This study used single sensillum recordings (SSR) to identify the responses of olfactory receptor neurons (ORNs) in the antennal sensilla of adult male and female C. fumiferana to host plant volatiles, and female sex pheromones. There have been few SSR studies done on spruce budworm, and to our knowledge, the present study represents the first attempt to examine the responses of ORNs from antennal sensilla in response to a range of host and conspecific stimuli. A total of 86 sensilla were characterized and sorted into 15 possible sensillum categories based on odor responses. We observed that specialist sensilla responding to few ligands were more abundant in both male and female than sensilla exhibiting more generalized odorant responses. (E/Z)-11-tetradecenal elicited responses from ORNs from any sensilla which were sensitive to pheromones in both males and females. Female C. fumiferana ORNs were able to detect and physiologically respond to female-produced sex pheromones with the same degree of sensitivity as their male counterparts. Together, these data improve our knowledge of mechanisms by which adult budworms respond to pheromone and host plant volatiles and provide insights that may be complementary to existing integrated pest management (IPM) strategies based on the chemical ecology of spruce budworm.

Analysis of Streptomyces Volatilomes Using Global Molecular Networking Reveals the Presence of Metabolites with Diverse Biological Activities.

Streptomyces species produce a wide variety of specialized metabolites, some of which are used for communication or competition for resources in their natural environments. In addition, many natural products used in medicine and industry are derived from Streptomyces, and there has been interest in their capacity to produce volatile organic compounds (VOCs) for different industrial and agricultural applications. Recently, a machine-learning workflow called MSHub/GNPS was developed, which enables auto-deconvolution of gas chromatography-mass spectrometry (GC-MS) data, molecular networking, and library search capabilities, but it has not been applied to Streptomyces volatilomes. In this study, 131 Streptomyces isolates from the island of Newfoundland were phylogenetically typed, and 37 were selected based on their phylogeny and growth characteristics for VOC analysis using both a user-guided (conventional) and an MSHub/GNPS-based approach. More VOCs were annotated by MSHub/GNPS than by the conventional method. The number of unknown VOCs detected by the two methods was higher than those annotated, suggesting that many novel compounds remain to be identified. The molecular network generated by GNPS can be used to guide the annotation of such unknown VOCs in future studies. However, the number of overlapping VOCs annotated by the two methods is relatively small, suggesting that a combination of analysis methods might be required for robust volatilome analysis. More than half of the VOCs annotated with high confidence by the two approaches are plant-associated, many with reported bioactivities such as insect behavior modulation. Details regarding the properties and reported functions of such VOCs are described. IMPORTANCE This study represents the first detailed analysis of Streptomyces volatilomes using MSHub/GNPS, which in combination with a routinely used conventional method led to many annotations. More VOCs could be annotated using MSHub/GNPS as compared to the conventional method, many of which have known antimicrobial, anticancer, and insect behavior-modulating activities. The identification of numerous plant-associated VOCs by both approaches in the current study suggests that their production could be a more widespread phenomenon by members of the genus, highlighting opportunities for their large-scale production using Streptomyces. Plant-associated VOCs with antimicrobial activities, such as 1-octen-3-ol, octanol, and phenylethyl alcohol, have potential applications as fumigants. Furthermore, many of the annotated VOCs are reported to influence insect behavior, alluding to a possible explanation for their production based on the functions of other recently described Streptomyces VOCs in dispersal and nutrient acquisition.

Methylmercury in caddisflies and mayflies: Influences of water and sediment chemistry.

Quantifying methylmercury (MeHg) concentrations and uptake at the base of the food web is useful for assessing mercury exposure risk to higher trophic level organisms. Higher MeHg concentrations near the base of the food web may result in more MeHg exposure and accumulation in higher trophic organisms. Here, we analyze MeHg in caddisflies, mayflies, lake water, and sediment collected from two temperate lakes and one brook in Kejimkujik National Park, Nova Scotia, Canada. Overall, caddisfly larvae MeHg (15.38-276.96 ng/g; n = 29) was not significantly correlated with water chemistry. Whereas mayfly naiads MeHg (14.28-166.82 ng/g; n = 31) was positively correlated with water MeHg (rs = 0.43), negatively correlated with pH (rs = -0.49), and positively correlated with dissolved organic carbon (DOC; rs = 0.48). Of the mercury in insect tissues, the %MeHg ranged from 56 to 75 % in caddisfly larvae and 38-47 % in mayfly naiads. MeHg bioaccumulation factors (BAF) varied greatly (water to tissue BAFs = 0.145 × 106-1.054 × 106; sediment to tissue BAFs = 0.017 × 106-0.541 × 106). This study highlights the importance of quantifying variations in MeHg bioaccumulation and BAFs of common aquatic insect bioindicators at the base of complex food webs.

Assessment of Attractant Lures and Monitoring Traps for Drosophila suzukii (Diptera: Drosophidae) Using Electrophysiology, Laboratory Choice Assays, and Field Trials.

Monitoring is critical to control efforts for Drosophila suzukii Matsumura, an invasive polyphagous fly that has the potential to cause significant losses in commercial soft fruit and berry production worldwide. We used an iterative process to identify trap colors, trap designs, and volatile mixtures to improve monitoring efforts in commercial blueberry, raspberry, and blackberry crops. Our results suggest that the selection of trap color and design and attractant lures should be customized to the crop in which they are deployed. In raspberries grown in high tunnel systems, DrosaLure paired with Drosal traps painted green and purple were highly specific to D. suzukii although actual capture counts were low. However, in field grown raspberries, BioLure and Multilure traps were most effective, but with significant nontarget bycatch. In blueberries, we had greatest success with a 5 µg:50 ng mixture of ethyl acetate-acetoin in a green/purple-colored jar-style trap with large (5 cm) mesh covered openings.

Preliminary Evaluation of a Granite Rock Dust Product for Pest Herbivore Management in Field Conditions.

The effects of granite rock dust in dry and aqueous formulations were evaluated under field conditions for control of insect pests in different crop systems and ornamental plants. We tested efficacy of crop protection following foliar applications on lily, squash, and cabbage plants by evaluating subsequent pest damage, overall plant health, and quantity of crops produced over one season. Lily plants treated with dry and aqueous formulations of rock dust were subject to lower herbivore damage (>1% and 11% herbivory damage, respectively) when compared to the controls (30% herbivory damage). Treatment on cabbage was less effective to protect plants against herbivory damage, and no statistically significant differences were reported within treatments. The foliar applications (dry and aqueous formulations) had positive impacts on growth of squash fruit resulting in a 2.5-fold increase in size relative to the control squash fruit. These results support the potential field application of granite dust to protect ornamental plants against herbivory attack, and reveal an alternative positive effect of the silica-based product on plant growth and development.

Production of Plant-Associated Volatiles by Select Model and Industrially Important Streptomyces spp.

The Streptomyces produce a great diversity of specialized metabolites, including highly volatile compounds with potential biological activities. Volatile organic compounds (VOCs) produced by nine Streptomyces spp., some of which are of industrial importance, were collected and identified using gas chromatography-mass spectrometry (GC-MS). Biosynthetic gene clusters (BGCs) present in the genomes of the respective Streptomyces spp. were also predicted to match them with the VOCs detected. Overall, 33 specific VOCs were identified, of which the production of 16 has not been previously reported in the Streptomyces. Among chemical classes, the most abundant VOCs were terpenes, which is consistent with predicted biosynthetic capabilities. In addition, 27 of the identified VOCs were plant-associated, demonstrating that some Streptomyces spp. can also produce such molecules. It is possible that some of the VOCs detected in the current study have roles in the interaction of Streptomyces with plants and other higher organisms, which might provide opportunities for their application in agriculture or industry.

Detection of heliothine sex pheromone components in the Australian budworm moth, Helicoverpa punctigera: electrophysiology, neuroanatomy, and behavior.

This study examined electrophysiological responses of the Australian budworm moth Helicoverpa punctigera, to heliothine sex pheromone components, via single sensillum recordings (SSR), and examined male neuroanatomy using confocal microscopy and 3D imaging tools. We found that male H. punctigera have three distinct regions of the macroglomerular complex (MGC) in the antennal lobe. Male antennae have only two functional types of sensilla trichoidea (A and C) and type A sensilla contain an olfactory sensory neuron (OSN) that responds to the major sex pheromone component (Z)-11-hexadecenal (Z11-16:Ald) with axons projecting to the cumulus of the macroglomerular complex (MGC) in the antennal lobe. Type C sensilla contained large-spiking receptor neurons which responded primarily to (Z)-9-tetradecenal (Z9-14:Ald) and to a lesser degree to (Z)-11-hexadecenol (Z11-16:OH). These were co-compartmentalized with small-spiking receptor neurons in type C sensilla which responded strongly to Z9-14:Ald and (Z)-9-hexadecenal (Z9-16:Ald), and to a lesser degree to (Z)-11-hexadecenyl acetate (Z11-16:OAc) and Z11-16:OH. Axons from the two co-localized neurons in Type C sensilla projected to the two small MGC units, the dorsomedial anterior and dorsomedial posterior, respectively. In wind tunnel assays, the addition of Z9-16:Ald to an otherwise attractive blend completely shut down male H. punctigera upwind flight.

Electrophysiological Responses of the Beech Leaf-Mining Weevil, Orchestes fagi, to Seasonally-Variant Volatile Organic Compounds Emitted by American Beech, Fagus grandifolia.

The beech leaf-mining weevil, Orchestes fagi, is a common pest of European beech, Fagus sylvatica, and has recently become established in Nova Scotia, Canada where it similarly infests American beech, F. grandifolia. We collected volatile organic compounds (VOCs) emitted by F. grandifolia leaves at five developmental stages over one growing season and simultaneously analyzed them for volatile emissions and O. fagi antennal response using gas chromatography-electroantennographic detection (GC-EAD). Volatile profiles changed significantly throughout the growing season, shifting from primarily ß-caryophyllene, methyl jasmonate, and simple monoterpene emissions to dominance of the bicyclic monoterpene sabinene during maturity. Two VOCs dominant during bud burst, (R)-(+)-limonene and geranyl-p-cymene, may be of biological relevance due to the highly specific oviposition period of O. fagi at this stage though antennal responses were inconclusive. Senescence showed a decrease in blend complexity with an increase in (Z)-3-hexenyl acetate and (Z)-3-hexen-1-ol as well as a resurgence of α-terpinene and geranyl-p-cymene. We present a novel electroantennal preparation for O. fagi. Antennae of both male and female O. fagi responded to the majority of detectable peaks for host volatiles presented via GC-EAD. Females displayed greater overall sensitivities and less specificity to host volatiles and it is hypothesized that this translates to more generalist olfaction than males. It is clear that olfactory cues are important physiologically though their implications on behaviour are still unknown. The results presented in this study provide a baseline and tools on which to connect the complex and highly time-specific phenology of both F. grandifolia and the destructive pest O. fagi through which olfactory-based lures can be investigated for monitoring systems.

Chemosensory and Behavioural Responses of Ixodes scapularis to Natural Products: Role of Chemosensory Organs in Volatile Detection.

Blacklegged ticks, Ixodes scapularis, represent a significant public health concern due to their vectoring of tick-borne disease. Despite their medical importance, there is still limited knowledge of the chemosensory system of this species, and thus a poor understanding of host-seeking behaviour and chemical ecology. We investigated the electrophysiological sensitivity of adult female blacklegged ticks to attractants and plant-derived compounds via an electrode inserted into the scutum. The response of female ticks to binary mixtures with a constant concentration of a selected attractant (butyric acid) and increasing concentration of volatile organic compounds (VOCs) (geraniol, phenethyl alcohol, ß-citronellol, and citral) was recorded. A strict relationship between increasing volatile concentration and a decreasing response was observed for each VOC. Y-tube bioassays confirmed that tick attraction towards butyric acid decreased with the presence of a VOC, which exerted a deterrent effect. To determine the specific role of sensory appendages involved in the detection of attractant chemical stimuli, we tested tick electrophysiological response after removing appendages that house chemosensory sensilla (foretarsi, pedipalps, or both). The chemosensory response was related to the molecular structure of attractant odorant, and the lack of pedipalps significantly reduced olfactory responses, suggesting they play an important role in detecting attractants. This study provides new insight into the neurophysiological mechanisms underlying tick olfaction and the potential for interactions between attractant and deterrent chemical detection.

Electrotarsogram responses to synthetic odorants by Varroa destructor, a primary parasite of western honey bees (Apis mellifera).

Olfaction is a key sensory modality for many arthropods and could be used as a tool in pest management through manipulation of pest behavior. Management of Varroa destructor, important parasitic mites of honey bees, could be improved through better understanding of the chemical ecology of this host-parasite relationship. We refined techniques of mounting mites to obtain electrophysiological recordings (electrotarsograms) of their responses to synthetic odor stimuli. Results of 271 electrotarsogram recordings from V. destructor revealed responses to 10 odorants relative to solvent controls. Electrotarsogram responses to methyl palmitate, ethyl palmitate, and 2-heptanol were highest at the lowest stimulus loading (10 ng) we tested, suggesting that V. destructor may have acute sensitivity to low concentrations of some odors. Results suggest that odorant origin (e.g., methyl oleate from honey bee larvae, geraniol from adult honey bee alarm pheromone, and α-terpineol, a plant secondary metabolite) can influence the degree of electrophysiological response. Varroa destructor tended to be more responsive to known attractants and repellents relative to previously unexplored odorants and some repellent terpenes. Electrotarsograms offer the potential for screening odors to determine their importance in V. destructor host detection.

Varroa destructor mite electrophysiological responses to honey bee (Apis mellifera) colony volatiles.

Detection and interpretation of chemical cues is essential for Varroa destructor Anderson and Trueman, an important parasite of honey bees (Apis mellifera L.), to complete its life cycle. We collected volatiles from honey bee brood at various developmental stages and screened for V. destructor electrophysiological responses to these with gas chromatography-linked electrotarsal detection. Volatile collections contained several methyl-alkanes that evoked electrophysiological responses from V. destructor. Moreover, odors in honey bee colonies that regulate honey bee colony structure and function were also detected by V. destructor. Collections from mid- to late-stage larvae had detectable levels of low-volatility odors identified as components of the honey bee brood pheromone and branched alkanes likely originating from brood cuticle. Among these, several mid- to heavy-molecular weight compounds elicited high proportional electrophysiological responses by V. destructor relative to their abundance but could not be identified using chemical standards of previously documented honey bee brood odors. We suggest further investigation of these unknown volatiles and future behavioral assays to determine attractiveness/repellency (valence) of those identified through chemical standards.

Plasticity Is Key to Success of Drosophila suzukii (Diptera: Drosophilidae) Invasion.

After its initial discovery in California in 2008, Drosophila suzukii Matsumura has become one of the most important invasive agricultural pest insects across climate zones in much of Asia, Europe, North America, and South America. Populations of D. suzukii have demonstrated notable behavioral and physiological plasticity, adapting to diverse environmental and climatic conditions, interspecific competition, novel food sources, and potential predators. This adaptability and plasticity have enabled rapid range expansion and diversified niche use by D. suzukii, making it a species particularly suited to changing habitats and conditions. This article reviews factors and evidence that influence plasticity in D. suzukii and promotes this species' invasiveness.

Color preference of the spotted wing Drosophila, Drosophila suzukii.

Drosophila suzukii Matsumura (Diptera: Drosophilidae) is a significant invasive pest in soft-skin fruits and berries in Asia, Europe, and North and South America. Many herbivorous insects use multiple cues for host selection, particularly olfactory and visual stimuli. The visual system of closely-related Drosophila melanogaster is well-documented, expressing strong sensitivity to short-wavelength colors (ultraviolet to green) and only limited sensitivity to long-wavelength colors (red to infrared). Our results suggest that D. suzukii have limited ability to distinguish red consistent with visual sensitivity range within the melanogaster subgroup. We propose that color contrast rather than color appearance may be of greater importance in orientation and attraction. We propose that differences in reflectance between light wavelengths important for color opponency are key to color discrimination to provide color contrast between foreground and background, as occurs between fruit and foliage, during host-finding.

Behavioral responses of Ixodes scapularis tick to natural products: development of novel repellents.

The spread of blacklegged ticks (Ixodes scapularis) and growing threat of Lyme disease transmission has increased demand for effective, safe and environmentally friendly repellent products. Plant-derived essential oils are natural products that exhibit insecticidal and repellant activities and represent a promising alternative to synthetic repellants. However, mechanisms by which ticks detect odor stimuli and how such stimuli may function as repellents are not well understood. We examined the repellent activity of selected essential oil components towards I. scapularis in short- and long-term dose-response trials. To determine the specific olfactory organs involved in detection of chemical stimuli, we tested tick behavioral response in repellency bioassays after removing appendages that house chemosensory sensilla (e.g., foretarsi or pedipalps). New prototype formulae were tested in longevity trials repelling up to 95% of tested ticks after 1 h post-application. This study provides new insight regarding tick olfaction and behavior, and innovative methods for selecting appropriate chemicals for development of novel plant-based repellent products for protection from ticks.

Considerations for Insect Learning in Integrated Pest Management.

The past 100 yr have seen dramatic philosophical shifts in our approach to controlling or managing pest species. The introduction of integrated pest management in the 1970s resulted in the incorporation of biological and behavioral approaches to preserve ecosystems and reduce reliance on synthetic chemical pesticides. Increased understanding of the local ecosystem, including its structure and the biology of its species, can improve efficacy of integrated pest management strategies. Pest management strategies incorporating insect learning paradigms to control insect pests or to use insects to control other pests can mediate risk to nontarget insects, including pollinators. Although our understanding of insect learning is in its early stages, efforts to integrate insect learning into pest management strategies have been promising. Due to considerable differences in cognitive abilities among insect species, a case-by-case assessment is needed for each potential application of insect learning within a pest management strategy.

Effect of Color and Contrast of Highbush Blueberries to Host-Finding Behavior by Drosophila suzukii (Diptera: Drosophilidae).

Drosophila suzukii Matsumara (Diptera: Drosophilidae) has become a serious pest in soft-skin fruits and berries, infesting both ripe and ripening fruits. Crop damage in highbush blueberry has been particularly severe. During blueberry fruit development, fruits of various degrees of ripeness are present simultaneously. In addition, foliage color changes as the season progresses. We investigated the influence of blueberry fruit and leaf color on host-finding behavior in D. suzukii. Opposing shifts between reflectance spectra of ripening fruits and senescing leaves increased contrast between ripe fruit and senesced foliage. Developmental changes in contrast between fruit color and leaf color may act as a visual contextual cue in finding suitable host fruits. Opposing shifts in reflectance spectra of ripening fruits and senescing leaves increased the contrast between ripe fruit and senesced foliage. These opposing changes in color may contribute to the attractiveness of blueberry fruit as a late season host for D. suzukii.

Susceptibility of selected boreal fruits and berries to the invasive pest Drosophila suzukii (Diptera: Drosophilidae).

BACKGROUND: Drosophila suzukii Matsumara has recently emerged as a major invasive pest species in soft-skinned fruits in berries throughout North America and Europe. Its distribution has spread so rapidly that little is known of the extent of fruit susceptibility, particularly in boreal regions. Populations of D. suzukii increase dramatically in late summer in boreal regions, concurrent with fruiting seasons for commercially and culturally significant fruits and berries. We tested fruit preference and susceptibility of lingonberry, blueberry, chokecherry, sea buckthorn and raspberry fruits to D. suzukii. RESULTS: Female D. suzukii attempted to oviposit on all fruit types tested. Fruits with lower brix and lower pH levels were preferred in choice tests. Undamaged lingonberries were relatively safe from infestation; however, bruised or frost-damaged fruits were easily penetrated. Sea buckthorn and raspberry fruits were highly preferred. CONCLUSION: Although blueberry growers have experienced severe economic crop losses due to D. suzukii, we have found that blueberries were the least preferred of the fruits tested. This suggests that D. suzukii are largely opportunistic and highlights the importance of fruit phenology in fruit susceptibility. © 2016 Society of Chemical Industry.

Differential Octopaminergic Modulation of Olfactory Receptor Neuron Responses to Sex Pheromones in Heliothis virescens.

Octopamine is an important neuromodulator of neural function in invertebrates. Octopamine increases male moth sensitivity to female sex pheromones, however, relatively little is known as to the role of octopamine in the female olfactory system, nor its possible effects on the reception of non-pheromone odorants. The purpose of this study was to determine relative effects of octopamine on the sensitivity of the peripheral olfactory system in male and female Heliothis virescens. Single sensillum recording was conducted in both sexes following injection with octopamine or Ringer solution, and during odorant stimulation with conspecific female sex pheromone or host plant volatiles. Results indicate that octopamine plays a significant modulatory role in female sex pheromone detection in female moths; and that male and female pheromone detection neurons share distinct pharmacological and physiological similarities in H. virescens despite sexual dimorphism at the antennal level.

Olfactory receptor neuron responses of a longhorned beetle, Tetropium fuscum (Fabr.) (Coleoptera: Cerambycidae), to pheromone, host, and non-host volatiles.

Longhorn wood-boring beetles (Coleoptera: Cerambycidae) use olfactory cues to find mates and hosts for oviposition. Tetropium fuscum (Fabr.) is an invasive longhorned wood-boring beetle originating from Europe that has been established in Nova Scotia, Canada, since at least 1990. This study used single sensillum recordings (SSR) to determine the response of olfactory receptor neurons (ORNs) in the antennal sensilla of male and female T. fuscum to different kinds of olfactory cues, namely host volatiles, non-host volatiles, the aggregation pheromone of T. fuscum (fuscumol), and an aggregation pheromone emitted by other species of longhorn beetles (3-hydroxyhexan-2-one). Each compound had been previously shown to elicit antennal activity in T. fuscum using electroantennography or had been shown to elicit behavioral activity in T. fuscum or other cerambycids. There have been very few SSR studies done on cerambycids, and ours is the first to compare response profiles of pheromone components as well as host and non-host volatiles. Based on SSR studies with other insects, we predicted we would find ORNs that responded to the pheromone alone (pheromone-specialists), as well as ORNs that responded only to host or non-host volatiles, i.e., separation of olfactory cue perception at the ORN level. Also, because male T. fuscum emerge earlier than females and are the pheromone-emitting sex, we predicted that the number of pheromone-sensitive ORNs would be greater in females than males. We found 140 ORNs housed within 97 sensilla that responded to at least one of the 13 compounds. Fuscumol-specific ORNs made up 15% (21/140) of all recordings, but contrary to our prediction, an additional 22 ORNs (16%) responded to fuscumol plus at least one other compound; in total, fuscumol elicited a response from 43/140 (31%) of ORNs with fuscumol-specific ORNs accounting for half of these. Thus, our prediction that pheromone reception would be segregated on specialist ORNs was only partially supported. Our prediction that females would have more ORNs that responded to fuscumol than would males was also not supported, as there was no difference. The stressed-host volatile linalool elicited the most responses of any compound tested, 43% of all recordings.