Host-associated volatile cues drive foraging behavior of Trichopria drosophilae toward Drosophila suzukii- infested fruits

Host-associated volatile cues drive foraging behavior of Trichopria drosophilae toward Drosophila suzukii- infested fruits | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Host-associated volatile cues drive foraging behavior of Trichopria drosophilae toward Drosophila suzukii- infested fruits Hai-Kuan Sun, Fan Yang, Ze-Hua Wang, Ren Li, Guang-Hang Qiao, and 3 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6960668/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 05 Dec, 2025 Read the published version in Journal of Pest Science → Version 1 posted 10 You are reading this latest preprint version Abstract The spotted-wing drosophila ( Drosophila suzukii ), a globally invasive pest, induces distinct shifts in the volatile organic compound (VOC) profiles of infested cherry fruits, enhancing their attractiveness to the pupal parasitoid Trichopria drosophilae . Comparative VOC analysis revealed significant differences among D. suzukii -infested, mechanically damaged, and healthy fruits. Infested cherries emitted elevated levels of key attractants, including esters (ethyl acetate: 0.36 ng/µL; isoamyl acetate: 0.48 ng/µL) and aromatic aldehydes (4-ethylbenzaldehyde: 0.86 ng/µL), which were absent or minimal in controls. Non-metric multidimensional scaling (NMDS) confirmed distinct VOC clustering, with infested fruits chemically diverging from healthy or mechanically damaged samples. Electrophysiological (EAG) assays identified ethyl acetate as the most potent stimulant, while behavioral assays showed concentration-dependent responses: 10 µg/µL isoamyl acetate elicited strong attraction (72% response rate), whereas 3-methylbutanoic acid acted as a repellent (34% response rate). Cage experiments demonstrated that benzyl alcohol, ethyl acetate, and isoamyl acetate significantly increased parasitism rates in D. suzukii pupae compared to controls, with isoamyl acetate showing the strongest effect. These results reveal specific semiochemicals mediating tritrophic interactions and underscore their potential for optimizing T. drosophilae -based biocontrol strategies against D. suzukii . Drosophila suzukii Trichopria drosophilae Volatile organic compounds (VOCs) Tritrophic interactions Biocontrol Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Introduction Plant volatiles serve as pivotal chemical signaling molecules that play a central regulatory role in tri-trophic interactions (plant-herbivore-parasitoid systems) (Vet and Dicke 1992 ; Becker et al. 2015 ; Ali et al. 2023 ), particularly by mediating parasitoid host-searching behavior and thereby influencing trophic cascades in ecosystems (Ayelo et al. 2021b ; Gómez-Cabezas et al. 2023 : Rossetti et al. 2025 ). Among these volatiles, herbivore-induced plant volatiles (HIPVs) are particularly critical, as they facilitate indirect plant defense by attracting natural enemies such as parasitoids and predators, thereby reducing herbivore damage (Xiu et al. 2019 ; Ayelo et al. 2021a ; Yasa et al. 2024 ). Fruit volatile profiles undergo dynamic changes in response to ripening stages (Bi et al. 2023 ) and physical damage (Beck et al. 2008 ). Notably, pathogen or insect infestation significantly alters these profiles, generating chemical cues that guide herbivores to hosts (Mohammed et al. 2019 ; Guo et al. 2023a ). For instance, the parasitoid Aphytis melinus is strongly attracted to D-limonene and β-ocimene emitted from Aonidiella aurantii -infested lime fruits, suggesting these volatiles play a key role in host location (Mohammed et al. 2019 ). Similarly, fungal-infected apples display shifted volatile organic compound (VOC) patterns, characterized by elevated attractants (e.g., ethyl 2-methylbutyrate) and suppressed repellents, ultimately enhancing oviposition preference in Conogethes punctiferalis (yellow peach moth). Key VOCs such as amyl 2-methylbutyrate and heptacosane have been identified as primary drivers of this behavioral response (Guo et al. 2023a ). Therefore, identifying the key bioactive compounds emitted by insect-infested fruits that significantly influence parasitoid behavior is of critical importance. Parasitic wasps integrate host-specific chemical cues with background odors to locate hosts efficiently (Schröder and Hilker 2008 ). For instance, Diachasmimorpha longicaudata , a parasitoid wasp targeting Ceratitis capitata , exhibits strong attraction to volatiles emitted from infested oranges, with six key compounds (D-limonene, acetophenone, linalool, nonanal, decanal, and eugenol) identified as critical attractants (Devescovi et al. 2024 ). Similarly, behavioral studies reveal that Trichopria anastrephae is highly responsive to volatiles from D. suzukii -infested strawberries (containing eggs, larvae, or pupae) and overripe fruits, underscoring the pivotal role of host-associated chemical signals in parasitoid foraging (Krüger et al. 2024 ). Notably, Ganaspis brasiliensis G1, a specialized parasitoid of Drosophila suzukii , dynamically adjusts its response to chemical cues (VOCs and cuticular hydrocarbons), preferentially targeting infested ripening fruits while avoiding decaying substrates. This shift from attraction to repulsion during larval development aligns with its ecological niche as a precision biocontrol agent (Giorgini et al. 2024 ). Collectively, these findings highlight the central role of odor-mediated tritrophic interactions in enhancing parasitoid foraging efficacy within agricultural ecosystems. Trichopria drosophilae is a globally distributed pupal endoparasitoid that targets various fruit fly species, including the invasive pest D. suzukii (Chabert et al. 2012 ; Mazzetto et al. 2016 ; Chen et al. 2018 ; Lisi et al. 2024a ). Unlike traditional fruit flies (e.g., Drosophila melanogaster ) that infest rotting fruits, D. suzukii (spotted-wing drosophila, SWD) possesses a serrated ovipositor, enabling it to damage ripening fruits at early developmental stages, causing severe economic losses to soft-skinned stone fruits such as cherries and blueberries (through inducing rot and mold development in fresh fruits) (Lee et al. 2011 ; Asplen et al. 2015 ; Haye et al. 2016 ; Labbetoul et al. 2025 ). Laboratory studies have demonstrated T. drosophilae 's strong preference for SWD (Wang et al. 2016 ; Yi et al. 2020 ), with enhanced offspring fitness when parasitizing this host (Boycheva Woltering et al. 2019 ; Jarrett et al. 2022 ; Gonzalez-Cabrera et al. 2023 ). Field trials in Mexican orchards further revealed that releasing 4,500 parasitoids per hectare reduced SWD populations by 50% (Gonzalez-Cabrera et al. 2021 ). These findings highlight the potential of T. drosophilae as a biocontrol agent against SWD within an Integrated Pest Management (IPM) framework, offering a sustainable tool to reduce chemical pesticide use and thereby limiting adverse effects on non-target organisms (Desneux et al. 2007 ). However, its implementation must be carefully evaluated in IPM packages where some pesticide applications may still be necessary, to ensure compatibility and effectiveness (e.g. Guo et al. 2023b ; Fouani et al. 2024; Lisi et al. 2024b ). The efficacy of parasitic wasps in field-based biological control is often constrained by both temporal (Pfab et al. 2018 ; Rossi Stacconi et al. 2019 ) and spatial factors (Rossi Stacconi et al. 2018 ; Hogg et al. 2022 ), which explains why their application remains underutilized in the integrated management of D. suzukii . Deciphering the tritrophic chemical communication network among host fruits, D. suzukii , and parasitic wasps could identify key attractant compounds, thereby providing critical insights for field application strategies. This study addresses three pivotal scientific questions: (i) Whether SWD infestation alters the volatile organic compound (VOC) profile of sweet cherries; (ii) The ability of T. drosophilae to discriminate SWD-infested cherries via VOCs; and (iii) Identification of key chemical compounds mediating this behavioral response. Our findings will provide a theoretical foundation for developing VOC-based precision biocontrol strategies against D. suzukii . Materials and methods Insect collection and rearing Drosophila suzukii (spotted-wing drosophila) and Drosophila melanogaster (vinegar fly) were collected in 2022 from the cherry experimental base at the Institute of Forestry and Pomology, Beijing Academy of Agriculture and Forestry Sciences (39°41′N, 116°42′E). Their parasitoid Trichopria drosophilae was concurrently captured at the same location using fermenting fruit-baited traps. Both Drosophila species were reared according to the artificial diet described by Yang et al. 2024 , with all insects maintained in climate-controlled chambers (25 ± 1°C, 70 ± 5% RH, 16:8 h L:D photoperiod). The T. drosophilae population was propagated using D. melanogaster pupae as hosts under identical environmental conditions. Chemicals Ethyl acetate (CAS: 141-78-6, ≥ 99.7%), methyl acetate (CAS: 79-20-9, ≥ 99.0%), isoamyl acetate (CAS: 123-92-2, ≥ 99.5%), ethyl benzene (CAS: 93-89-0, ≥ 99.5%), benzyl alcohol (CAS: 100-51-6, ≥ 99.8%), phenylethanol (CAS: 60-12-8, ≥ 99.5%), ethanol (CAS: 64-17-5, ≥ 99.8%), isovaleric acid (CAS: 503-74-2, ≥ 99.0%), acetic acid (CAS: 64-19-7, ≥ 99.8%), benzaldehyde (CAS: 100-52-7, ≥ 98%), n-hexane (CAS: 110-54-3, ≥ 97%), carene (CAS: 4497-92-1, ≥ 97%). All chemicals were purchased from Shanghai Macklin Biochemical Technology Co., Ltd. Cherry Fruit Treatment Purchased fresh cherries (cultivar: Tieton) were gently washed and carefully inspected for insect eggs, physical damage, or abnormalities to ensure they were free from external defects and pest contamination. To induce D. suzukii infestation, five pairs of adult flies (3–7 days old) were placed in plastic containers containing approximately 50 g of cherries and sealed for 24 hours. After 24 hours, adult flies were removed, and the number of eggs on each cherry was counted under a microscope (average: 10 eggs per cherry). Cherries were then stored in clean plastic containers with fine mesh covers at 25 ± 1°C, 70 ± 5% RH, and a 16 L: 8 D light cycle for 7 days. Once larvae pupated, the pupae were removed, and the fruit volatiles were collected. To distinguish the effects of D. suzukii infestation from physical damage and natural ripening, two controls were used: cherries were punctured 10 times with a sterilized insect needle (0.3 mm diameter) 7 days before volatile collection, and an untreated control group was set under the same conditions without any treatment. Each treatment was repeated eight times, with containers wiped with 75% ethanol for sterility. Collection of Fruit Volatiles Cherries were placed in a dynamic headspace collection bottle (650 mL). The air pump (Model: CAT-8000) was set to a flow rate of 300 mL/min. The air was previously dehumidified with silica gel and filtered through activated carbon. The volatiles were adsorbed in a glass tube packed with 100 mg of Porapak Q adsorbent (Waters Corporation, USA, 80–100 mesh) for a collection period of 24 hours with three replicates. The adsorbed volatiles were eluted with 400 µL of n-hexane, concentrated to 150 µL, and stored in a 2 mL vial at -20°C until gas chromatography-mass spectrometry (GC-MS) analysis. Prior to sample concentration, 2 µL of an internal standard solution (carene) was added. VOC collections were made at a temperature of 25 ± 1°C, 70 ± 5% relative humidity and a photoperiod of 12:12 h (L: D). Chemical Analyses Volatiles emitted by cherries were analyzed using gas chromatography-mass spectrometry (GC-MS) with an Agilent 7890 GC coupled to a 5975 MSD. The GC was equipped with an HP-5 capillary column (30 m × 0.25 mm × 0.25 µm, Agilent Technologies). A 1 µL sample was injected in splitless mode, with the injection port set at 250°C and the ion source at 230°C. Electron ionization (EI) was performed with an ionization energy of 70 eV, using helium as the carrier gas at a flow rate of 1 mL/min. The temperature program was as follows: initial temperature 60°C, held for 1 min; ramped at 4°C/min to 200°C, held for 2 min; ramped at 15°C/min to 260°C, held for 5 min, for a total run time of 47 min. Compounds were identified by matching chromatographic peaks with those in the NIST 8.0 mass spectral library, and confirmed by comparison with authentic standards. The concentrations of volatile compounds released in the treatments (infested by D. suzukii , mechanically damaged, and control) were quantified based on the peak areas of the internal standard. Concentration of unknown volatiles = (Concentration of internal standard × Peak area of unknown) / Peak area of internal standard. Olfactometer Bioassays The olfactory response of female Trichopria drosophilae (3–7 days post-emergence) to host fruit location was measured using a “Y”-shaped olfactometer. The two arms of the olfactometer were 6.0 cm in length, with a straight tube of 9.0 cm in length and an inner diameter of 0.8 cm. The olfactometer was located in a dark opaque chamber (80 cm × 80 cm × 50 cm) to avoid visual interference. Humidied charcoal-filtered air was pushed through the stimuli and olfactometer with a pump at a rate of 100 mL/min. A 40 W incandescent bulb was positioned directly above the box to ensure uniform light intensity across both testing arms. All bioassays were conducted at 25 ± 1°C, 70 ± 5% RH and performed between 09:00 and 11:00 am. After every five wasps were tested, the positions of odor sources in the Y-tube olfactometer were swapped, and a new Y-tube was used. At the end of the experiment, all Y-tubes were collectively cleaned with 95% ethanol for subsequent trials. For the fruit volatiles, three treatment groups were established for comparison: Drosophila suzukii -infested fruit vs. clean air, healthy fruit vs. clean air, and D. suzukii -infested fruit vs. healthy fruit. Subsequently, based on the Electroantennographic (EAG) test results, we randomly selected several compounds and conducted olfactometer tests at different concentrations: 1 µg/µL, 10 µg/µL, and 100 µg/µL. For each treatment, 10 µL of the prepared attractant solution was applied to a filter paper strip (3 cm × 1 cm) and placed in one volatile source bottle. An equal volume of the control solvent (n-hexane) was applied to a separate filter paper strip and placed in the other source bottle. Female T. drosophilae was placed at the base of the “Y” olfactometer, and their behavioral responses were observed and recorded over a 5-minute period. The selection of the odor source in each arm was noted when the female was either within 5 cm of the exit or had passed halfway down an arm and remained there for at least 5 seconds. If no choice was made within the 5-minute observation period, the response was recorded as “no reaction”. Electroantennographic (EAG) The EAG technique was used to assess the antennal selectively and sensitivity of T. drosophilae females to the selected ten volatile organic compounds (VOCs). The antennae of female T. drosophilae were excised at the base using fine scissors. Immediately afterward, approximately 0.5–1 mm was trimmed from the tips of the antennae using a surgical blade. The prepared antennae were then connected to the electrodes using a 2 mm-diameter glass capillary filled with PBS (pH = 7.2) solution. The electrophysiological signals of antennal responses to stimuli were amplified, output, and collected using a data acquisition controller (IDAC). The collected signals were processed and analyzed using EAG-Pro software. Once the baseline signal stabilized, volatile collections were tested. Preliminary experiments were conducted to determine the optimal concentration of standard compounds, which were dissolved in n-hexane. The final concentration gradient was set to 1 µg/µL for the electroantennographic (EAG) response measurements. For each volatile, 10 µL was taken with a micropipette and applied to a clean 1 × 3 cm² filter paper. After 20 s at room temperature to allow evaporation of n-hexane, the paper was placed into a Pasteur pipette (diameter 2 mm, length 12 cm). A control sample was prepared by adding an equal volume of n-hexane to another filter paper under identical conditions. Each volatile was tested on 18 antennae with active responses (one antenna per wasp). There was a 30 s interval between each stimulation, and control measurements were conducted before and after testing each volatile. The relative EAG response value of the antenna to each odorant was calculated as: EAG response value = Volatile response value - (Pre-test control response value + Post-test control response value) / 2. Cage experiment The experiment was conducted in 35 × 35 × 35 cm rearing cages, each equipped with two diagonally placed self-made rubber septum dispensers. For treatment groups, 200 µL of benzyl alcohol, ethyl acetate, or isoamyl acetate (10 µg/µL, selected based on preliminary Y-tube olfactometer screening of bioactive compounds) was applied to the dispensers, while an equal volume of n-hexane served as the solvent control. Twenty-five 1-day-old pupae of D. suzukii were introduced as hosts, followed by the release of five pairs of T. drosophilae adults (3–7 days post-emergence). The dispenser and fly pupae were placed in an open 6 cm diameter Petri dish. A 6 cm Petri dish containing a cotton ball soaked in 10% honey solution was positioned at the center of the cage to provide nutrition for the parasitoid wasps. After 24 h of exposure, pupae were collected to quantify parasitism. After collection, the fly pupae were maintained at 25 ± 1°C, 70 ± 5% relative humidity, and a 16:8 h (light:dark) photoperiod until adult emergence. To confirm parasitism, pupae that failed to eclose were subjected to PCR amplification using species-specific primers to detect T. drosophilae DNA (unpublished content), allowing quantification of parasitized hosts. Each treatment was independently replicated eight times, with fresh hosts and parasitoids used for every replicate. Parasitism rate was calculated as: Parasitism rate (%) = Number of parasitized hosts /Total hosts released ×100% The three volatiles were tested separately, with n-hexane serving as a unified solvent control to eliminate confounding effects. Statistical Analysis Statistical analyses were conducted to evaluate female T. drosophilae behavioral responses through binomial tests examining both host preference (SWD-attacked vs. Undamaged control, SWD-attacked vs. Clean air, and Undamaged control vs. Clean air) and concentration-dependent responses to ten volatile compounds (1, 10, and 100 µg/µL). Volatile organic compound profiles were characterized using NMDS ordination based on Bray-Curtis dissimilarity (stress = 0.183), with statistical significance assessed through PERMANOVA (999 permutations) followed by Benjamini-Hochberg-adjusted pairwise comparisons when appropriate. Electroantennographic responses were analyzed with Kruskal-Wallis tests and Dunn's post hoc tests for significant findings ( p < 0.05), while treatment effects in cage experiments were evaluated using paired t-tests. All statistical analyses were performed using R (Version 4.4.2) (1999–2024 R Core Team). Results Preference of Trichopria drosophilae females for differently treated cherry fruits The binomial tests revealed significant preferences in Trichopria drosophilae for certain odor sources. When given a choice between damaged fruits and healthy fruits, the wasps selected damaged fruits significantly more often (62.9% preference, p = 0.041). Similarly, in the damaged fruits vs air comparison, the wasps showed a stronger attraction to damaged fruits (64.3% preference, p = 0.022). However, no significant preference was observed between healthy fruits and air (55.7% preference, p = 0.403). These results indicate that T. drosophilae exhibited a clear preference for volatiles from damaged cherry fruits over both healthy fruits and air, while showing no significant bias when choosing between healthy fruits and air. Analysis of volatile organic compound (VOC) differences in differently treated cherry fruits A total of 48 volatile compounds were identified in cherry samples. The highest diversity of compounds, including esters, alkenes, and alcohols, was detected in cherries infested by D. suzukii , with 11 compounds being exclusively present in the volatiles of infested cherries. The concentration of 4-ethylbenzaldehyde was significantly higher in D. suzukii -infested cherries (0.86 ± 0.22 ng/µL) compared to the control group (0.20 ± 0.11 ng/µL). In contrast, hexacos-1-ene exhibited a higher concentration in the physical damage group (1.42 ± 0.49 ng/µL) but was reduced in the D. suzukii -infested group (0.35 ± 0.29 ng/µL). Additionally, ester compounds such as ethyl acetate, methyl acetate, and isoamyl acetate were present at higher concentrations in D. suzukii -infested cherries, while they were either undetected or present at lower levels in the control group. These results demonstrate that D. suzukii infestation significantly alters the concentrations of specific compounds in cherry fruits (Table 1 ). Volatile organic compounds were quantified in sweet cherry samples subjected to three experimental treatments: D. suzukii infestation, mechanical injury, and undamaged controls (n = 8 per group). The specific concentrations (ng/µL) of detected volatiles are presented in Table 1 . The ten volatile compounds—ethyl acetate, methyl acetate, isoamyl acetate, ethyl benzoate, benzyl alcohol, phenethyl alcohol, ethanol, 3-methylbutanoic acid, acetic acid, and benzaldehyde—were selected for further investigation due to their significantly higher occurrence frequency and concentration levels in D. suzukii -infested cherries. Most of these compounds were exclusively detected in infested samples, while a few, though present in both infested and healthy cherries but not in physically damaged ones, exhibited markedly elevated concentrations in D. suzukii -infested fruits compared to their trace levels in undamaged or mechanically injured samples. Table 1 Amounts of volatile organic compounds (VOCs) found in collections from cherries subjected to the different treatments: attacked by Drosophila suzukii (SWD-attacked); physical damage and natural fruit ripening (undamaged control). Compounds authenticated by reference standards are indicated with an asterisk (*). Compound CAS Amount (µg/25 g/24 h; mean ± standard error) SWD-attacked N Physically damaged N Undamaged control N Esters Ethyl valerate 539-82-2 0.06 ± 0.04 2 0.09 1 - 0 Isoamyl acetate* 123-92-2 0.48 ± 0.33 7 - 0 - 0 Ethyl benzoate* 93-89-0 0.50 ± 0.31 7 - 0 - 0 Methyl 3,5-dimethylbenzoate 25081-39-4 0.32 ± 0.07 5 0.29 ± 0.10 6 0.09 ± 0.04 6 Methyl 4-ethylbenzoate 7364-20-7 0.23 ± 0.07 6 0.21 ± 0.02 6 0.05 ± 0.01 7 Methyl hexadecanoate 112-39-0 0.16 ± 0.12 6 0.14 ± 0.06 6 0.05 ± 0.05 4 Ethyl acetate* 141-78-6 0.36 ± 0.20 7 - 0 - 0 Methyl stearate 112-61-8 0.26 ± 0.09 4 0.25 ± 0.11 5 0.12 ± 0.04 3 Methyl acetate* 79-20-9 0.25 ± 0.10 6 0.01 1 - 0 isoPropyl Myristate 110-27-0 0.12 ± 0.09 3 0.11 ± 0.03 4 0.04 ± 0.01 2 Methyl 14-methylpentadecanoate 5129-60-2 - 0 0.13 1 0.02 ± 0.01 4 Acetyl tributyl citrate 77-90-7 0.12 1 0.21 ± 0.05 4 0.05 ± 0.02 3 Ethyl isovalerate 108-64-5 0.04 ± 0.02 2 - 0 - 0 Ethyl Palmitate 628-97-7 0.07 ± 0.04 2 - 0 - 0 Ethyl 2-methylbutyrate 7452-79-1 0.02 ± 0.01 2 - 0 - 0 Triethyl citrate 77-93-0 0.06 1 - 0 - 0 Methyl Salicylate 119-36-8 0.03 1 - 0 - 0 Alkenes Styrene 100-42-5 0.12 ± 0.10 7 0.19 ± 0.14 5 0.01 ± 0.01 5 Pinene 7785-70-8 0.22 ± 0.14 7 0.24 ± 0.11 5 0.07 ± 0.04 5 1-Dodecene 112-41-4 0.08 ± 0.03 5 0.05 ± 0.02 4 0.04 ± 0.02 6 1-Hexadecene 629-73-2 0.06 ± 0.01 2 0.02 1 - 0 1-Heptadecene 6765-39-5 0.08 ± 0.06 2 0.05 1 - 0 Squalene 111-02-4 0.73 ± 1.14 5 0.61 ± 0.50 4 0.15 ± 0.18 4 1-Docosanol 1599-67-3 0.02 1 0.07 ± 0.01 2 - 0 1-Eicosene 3452/7/1 0.08 ± 0.04 3 0.10 ± 0.05 3 0.04 ± 0.03 2 1-Hexacosene 18835-33-1 0.35 ± 0.29 5 1.42 ± 0.49 4 0.52 ± 0.02 3 1,3,5,7-Cyclooctatetraene 629-20-9 0.12 ± 0.02 2 0.05 ± 0.04 3 - 0 17-Pentatriacontene 6971-40-0 - 0 0.01 1 - 0 α - Pinene 80-56-8 0.07 1 0.06 ± 0.01 4 0.02 ± 0.01 3 (+)-Camphene 79-92-5 - 0 0.01 ± 0.01 2 - 0 1-Nonadecene 18435-45-5 0.13 1 0.16 1 - 0 β-pinene 18172-67-3 0.10 ± 0.08 2 1 1 - 0 (9Z)-9-Tricosene 27519-02-4 0.06 ± 0.02 2 - 0 0.04 ± 0.03 2 (+)-Limonene 5989-27-5 0.44 ± 0.31 2 0.9 1 0.26 ± 0.14 4 Alcohols 0 Benzyl alcohol* 100-51-6 0.28 ± 0.23 6 - 0 - 0 Phenethyl alcohol* 1960/12/8 0.17 ± 0.18 6 0.17 1 - 0 (+)-Cedrol 77-53-2 0.11 ± 0.08 4 0.07 ± 0.02 4 0.03 ± 0.03 3 Ethanol* 64-17-5 0.37 ± 0.43 5 - 0 - 0 Acids Nonahexacontanoic acid 40710-32-5 0.08 1 0.04 1 0.04 1 3-Methylbutanoic acid* 503-74-2 0.14 ± 0.09 5 - 0 0.07 1 Acetic acid* 64-19-7 0.26 ± 0.09 5 0.02 1 - 0 Stearic acid 1957/11/4 0.02 1 - 0 - 0 Aromatics p-Xylene 106-42-3 0.04 ± 0.01 4 0.04 ± 0.01 3 0.02 ± 0.02 4 Naphthalene 91-20-3 0.39 ± 0.10 5 0.64 ± 0.55 3 0.08 ± 0.03 3 o-xylene 95-47-6 0.16 ± 0.15 2 0.05 ± 0.02 4 0.03 ± 0.04 4 Aldehydes Decanal 112-31-2 0.81 ± 0.49 5 1.46 ± 1.25 4 0.22 ± 0.17 6 Benzaldehyde* 100-52-7 0.72 ± 0.74 7 - 0 0.01 1 4-Ethylbenzaldehyde 4748-78-1 0.86 ± 0.22 4 0.95 ± 0.34 6 0.20 ± 0.11 4 VOC components are classified in groups and N indicates the number of samples in which a compound was found. Non-metric multidimensional scaling (NMDS) based on Bray-Curtis dissimilarity matrices revealed significant spatial separation in volatile organic compound (VOC) profiles among the three treatment groups (stress = 0.183, acceptable per Kruskal’s criterion). Non-metric multidimensional scaling (NMDS) analysis revealed distinct clustering patterns among the three experimental groups (Fig. 2 ). Drosophila suzukii -infested fruits (depicted as red dots) exhibited significant clustering in the negative NMDS1 dimension (mean ± SE: -0.530 ± 0.104), whereas undamaged control fruits (green dots) occupied the positive NMDS1 quadrant (0.565 ± 0.126). Fruits subjected to mechanical injury (blue dots) demonstrated intermediate positioning along the NMDS1 axis (-0.0355 ± 0.101), suggesting a transitional state between infestation and control conditions. These results indicate that VOC profiles of SWD-attacked and undamaged control fruits are distinctly separated along NMDS1, while physically damaged fruits exhibit an intermediate profile, suggesting a gradient in VOC composition based on treatment type. The key characteristics of spatial distribution revealed that the D. suzukii -attacked (SWD-attacked) group exhibited specific separation, with its mean NMDS1 value (-0.530) differing by 1.095 units from that of the healthy group (0.565), indicating directional changes in volatiles induced by pest infestation (Fig. 2 ). In contrast, the physically damaged group showed a heterogeneous response, as evidenced by the significantly higher standard error of the NMDS2 axis (0.209) compared to the healthy group (0.151), suggesting greater randomness in volatile release patterns (Fig. 2 ). These findings highlight distinct spatial distribution patterns in volatile compounds between pest-infested, physically damaged, and healthy cherry fruits. The volatile compound composition significantly differed among the treatment groups (R² = 0.328, F = 5.125, p = 0.001) (Table 2 ). Specifically, the D. suzukii -attacked (SWD-attacked) group showed highly significant differences compared to the undamaged control group (R² = 0.397, F = 9.220, p = 0.0015), and the physically damaged group also exhibited significant differences relative to the undamaged control group (R² = 0.282, F = 5.508, p = 0.0015) (Table 2 ). However, no significant differences were observed between the SWD-attacked group and the physically damaged group (R² = 0.124, F = 1.985, p = 0.094) (Table 2 ). These results indicated that both D. suzukii infestation and physical damage distinctly alter the volatile profiles of cherries, with D. suzukii infestation eliciting a more pronounced effect compared to undamaged controls. Table 2 Mean NMDS coordinates ± standard error of VOCs in cherry fruits under different treatments and results of pairwise comparisons Treatment NMDS1 (Mean ± SE) NMDS2 (Mean ± SE) Pairwise comparison groups R² F value Corrected p-values SWD-attacked -0.530 ± 0.104 -0.192 ± 0.111 vs Undamaged control 0.3971 9.22 0.0015 Physically damaged -0.0355 ± 0.101 0.0058 ± 0.209 vs SWD-attacked 0.1241 1.9845 0.0940 Undamaged control 0.565 ± 0.126 0.186 ± 0.151 vs Physically damaged 0.2823 5.5075 0.0015 EAG responses of Trichopria drosophilae females to cherry VOCs Through analysis, ten compounds with significantly increased volatile emissions in the D. suzukii -infested treatment were selected to further investigate the electrophysiological responses of T. drosophilae to these compounds. The electrophysiological responses (EAG) to ten volatile compounds varied significantly (χ² = 65.76, df = 9, p < 0.001). Ethyl acetate elicited the strongest response (mean ± SE: 0.0393 ± 0.0057), followed by benzaldehyde (0.0233 ± 0.0044) and methyl acetate (0.0170 ± 0.0037). Ethyl benzoate (0.0136 ± 0.0024) and isoamyl acetate (0.0120 ± 0.0025) showed moderate responses, with isoamyl acetate being significantly higher than phenethyl alcohol (0.0034 ± 0.0021, p = 0.022) and ethanol (-0.0006 ± 0.0012, p = 0.001). Acetic acid (0.0078 ± 0.0018) and 3-methylbutanoic acid (0.0079 ± 0.0024) induced weaker responses, while benzyl alcohol (0.0045 ± 0.0025) showed minimal activity. These findings indicate that esters (ethyl acetate, ethyl benzoate, isoamyl acetate, methyl acetate) and aromatic aldehydes (benzaldehyde) generally evoked stronger EAG responses than alcohols (benzyl alcohol, ethanol, phenethyl alcohol) or carboxylic acids (acetic acid, 3-Methylbutanoic acid), with ethyl acetate being the most potent stimulant (Fig. 3 ). Preference of Trichopria drosophilae females for VOCs at different concentrations Additionally, we further investigated whether T. drosophilae exhibited behavioral responses to these ten compounds. The Y-tube olfactometer tests revealed significant behavioral responses to specific volatile compounds. Notably, isoamyl acetate elicited strong attraction at 10 µg/µL concentration (72.0% response rate, p = 0.0026), while ethyl acetate showed dose-dependent effects with significant responses at both 100% (70.0%, p = 0.0066) and 10% (66%, p = 0.0328) concentrations. Benzyl alcohol demonstrated moderate attraction at 10% concentration (66%, p = 0.0328). Conversely, 3-methylbutanoic acid showed significant repellency at 10% concentration (34% response rate, p = 0.0328). Control compounds (ethanol, acetic acid, etc.) generally showed neutral responses (44–54% response rates, p > 0.05). The complete response patterns suggest concentration-dependent effects, with intermediate concentrations (10%) often producing the strongest behavioral responses compared to higher (100%) or lower (1%) concentrations. Oviposition Response of Trichopria drosophilae Females to VOCs Behavioral response assays of the ten target volatile compounds identified three candidates for validation in cage experiments. The parasitism rates under different chemical treatments showed significant variations compared to the controls, n-hexane. Benzyl alcohol significantly increased parasitism rates compared to controls (t 14 = -2.34, p = 0.035). Similarly, ethyl acetate showed a significant treatment effect (t 14 = -2.21, p = 0.044). The most pronounced effect was observed in the isoamyl acetate group, where parasitism rates were significantly higher in treated samples (t 14 = -2.71, p = 0.017). These results demonstrate that both benzyl alcohol, ethyl acetate and isoamyl acetate significantly promote parasitism, with isoamyl acetate showing the strongest enhancing effect among the tested compounds (Fig. 5 ). Discussion Our results demonstrate that Trichopria drosophilae specifically responds to Drosophila suzukii -induced HIPVs, showing strong attraction to infested fruits but not to healthy ones. Chemical analyses revealed distinct volatile profiles in infested fruits, particularly enriched esters and aromatic aldehydes, which were further confirmed as key attractants through electrophysiological and behavioral assays. Importantly, these HIPVs significantly enhanced parasitism rates, supporting their potential application in D. suzukii biological control. Evidence shows that T. drosophilae selectively targets D. suzukii -infested cherry fruits (62.9% vs. healthy; 64.3% vs. air), confirming its HIPV detection capability. Healthy fruits and air showed no difference in attraction (55.7%, p = 0.403), indicating that the parasitoid's response depends on herbivore-induced volatiles rather than fruit emissions. Volatile profiling revealed distinct chemical shifts in infested fruits (e.g., elevated esters and 4-ethylbenzaldehyde), with NMDS clearly separating infested and healthy samples (NMDS1: -0.530 vs. 0.565). Mechanically damaged fruits emitted intermediate volatiles, suggesting HIPV induction involves herbivore-specific factors (e.g., salivary enzymes or microbial symbionts). This mechanistic parallel is well-established in other systems, particularly Chilo suppressalis -infested rice where herbivore oral secretions trigger HIPV emission to recruit parasitoids(Yu et al. 2024 ), collectively demonstrating HIPVs' evolutionarily conserved function in mediating tritrophic interactions through herbivore-specific chemical cues. Importantly, herbivore-associated microbial communities to D. suzukii (Labbetoul et al. 2025 ) might modulate HIPVs released, representing a sophisticated co-evolutionary adaptation where microbial modulation of volatile profiles directly impacts parasitoid host-seeking efficiency while navigating complex host defense systems. These insights advance our understanding of HIPV-mediated ecological interactions and highlight potential applications for enhancing biological control strategies in integrated pest management. Trichopria drosophilae exhibits a unique high sensitivity to ester compounds. This aligns with findings that acetate esters emitted by microorganisms (e.g., Saccharomyces cerevisiae ) attract parasitoids such as T. drosophilae , likely facilitating host location by mirroring their hosts' olfactory preferences (Đurović et al. 2021 ). Similarly, GC-EAD analysis revealed that other female parasitoids ( Leptopilina boulardi and Trichopria anastrephae ) respond antennally to fruit esters (ethyl butanoate, methyl hexanoate, ethyl hexanoate) from D. suzukii -infested strawberries, confirming esters as critical foraging cues (Triñanes et al. 2022 ). However, the role of ester concentration in behavioral responses is context-dependent: while low concentrations may guide parasitoids, high concentrations of compounds like isoamyl acetate can deter D. suzukii females, possibly signaling overripe fruit unsuitable for oviposition (Cha et al. 2012 ; Revadi et al. 2015 ). These findings collectively highlight the dual role of ester volatiles in mediating trophic interactions, where their ecological function depends on both chemical identity and environmental context. Our study provides a clear example that strong electrophysiological responses to a volatile (e.g., 3-methylbutanoic acid at 1 µg/µL) do not necessarily predict behavioral attraction. Specifically, 3-methylbutanoic acid elicited antennal responses at 1 µg/µL but acted as a repellent at 10 µg/µL in “Y”-shaped olfactometer. This finding contrasts with observations in Trissolcus basalis , an egg parasitoid of Nezara viridula , where the same compound was attractive at similar concentrations when emitted from buckwheat flowers. Notably, Trissolcus basalis displayed strong antennal responses to 3-methylbutanoic acid at 100 µg/µL, suggesting that behavioral outcomes may diverge even when electrophysiological detection occurs (Foti et al. 2017 ). Concentration-dependent effects likely play a critical role—while certain low doses may attract or stimulate antennal detection, intermediate or high concentrations could either enhance attraction or trigger repellency, with no consistent linear pattern across species or compounds. For instance, (Z)-3-hexenol attracts Trichogramma chilonis at 0.1 µg/µL but repels it at 10 µg/µL, while (E)-2-hexenal exhibits the opposite trend. Similarly, nonanal shows no significant effect at 0.1 or 10 µg/µL but acts as a repellent at 1 µg/µL, further underscoring the nonlinear relationship between concentration and behavioral response (Wang et al. 2025 ). The same volatile compounds (e.g., aromatic and terpenoid compounds) from companion plants ( Desmodium spp. and Brachiaria ) repel fall armyworm ( Spodoptera frugiperda ) while attracting its parasitoid wasps, demonstrating the dual ecological function of these volatiles in the "Push-Pull" cropping system (Sobhy et al. 2022 ). Together, these results emphasize that the ecological role of a volatile compound cannot be generalized across species or contexts, and future research should explore how dose-dependent effects and volatile blends shape multitrophic interactions in agroecosystems. While this study provides valuable insights into T. drosophilae ’s behavioral responses, certain limitations should be noted. The laboratory conditions using single compounds may not fully represent the complexity of field environments, where volatile interactions and ecological factors likely play important roles. The ecological context of volatile emission must be considered. In natural environments, 3-methylbutanoic acid is part of a complex blend—such as in D. suzukii -infested cherry fruit volatiles, where it attracts parasitoid females, whereas its isolated presentation in Y-tube assays elicits repellency. Additionally, the molecular mechanisms underlying these olfactory responses remain to be explored. Future studies should further investigate these aspects under more realistic conditions, examine potential variations across different host plants and environmental contexts, and evaluate the practical effectiveness of these findings for field applications in D. suzukii management. Such work would help bridge the gap between laboratory research and real-world implementation. Our study demonstrated that T. drosophilae exhibited high specificity toward VOCs (especially esters) released by cherry infested with D. suzukii , highlighting the potential of these volatile cues in developing targeted biological control strategies. These findings provide a theoretical foundation for utilizing VOC-based attractants to enhance parasitoid efficiency in D. suzukii management. However, further research integrating chemical ecology and field trials is essential to optimize lure formulations (Muskat and Patel 2021; Nieri et al. 2022 ) and elucidate the underlying mechanisms governing parasitoid-host interactions. Such efforts will bridge the gap between laboratory findings and practical applications, ultimately contributing to sustainable pest control in agroecosystems. Declarations Author contributions S.N.W, Y.Y.W conceived and designed research. H.K.S conducted experiments and wrote the manuscript. F.Y analyzed data, prepared figures and wrote the manuscript. R.L, Z.H.W and G.H.Q revised the manuscript. Z.H.W and Q.F.L assisted in experiments. All authors read and approved the manuscript. Acknowledgements We express our sincere gratitude to Ting Geng from the Institute of Plant Protection, Chinese Academy of Agricultural Sciences (IPP-CAAS) for his expert assistance in capturing the diagnostic images of Trichopria drosophilae . Special thanks are extended to Yu Gao for her skillful contribution to the graphical illustrations. We also acknowledge the sustained support provided by the Institute of Forestry and Pomology, Beijing Academy of Agriculture and Forestry Sciences (BAAFS) throughout this research project. Funding This work was supported by the Promotion and Innovation of Beijing Academy of Agriculture and Forestry Sciences (KJCX20250901, KJCX20240403) and the Young Elite Scientists Sponsorship Program by the BAST (BYESS2023474). Data availability The datasets generated during or analyzed during the current study are available from the corresponding author upon reasonable request. Conflict of interest The authors have no relevant financial or non-financial interests to disclose. Ethical approval This statement is not applicable. 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Cite Share Download PDF Status: Published Journal Publication published 05 Dec, 2025 Read the published version in Journal of Pest Science → Version 1 posted Editorial decision: Revision requested 24 Aug, 2025 Reviews received at journal 29 Jul, 2025 Reviews received at journal 22 Jul, 2025 Reviewers agreed at journal 14 Jul, 2025 Reviewers agreed at journal 11 Jul, 2025 Reviewers agreed at journal 09 Jul, 2025 Reviewers invited by journal 09 Jul, 2025 Editor assigned by journal 26 Jun, 2025 Submission checks completed at journal 26 Jun, 2025 First submitted to journal 23 Jun, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6960668","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":483080999,"identity":"ad4e41ef-0a13-40a9-8c9c-c1c08127c978","order_by":0,"name":"Hai-Kuan Sun","email":"","orcid":"","institution":"College of Plant Protection, Hebei Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"Hai-Kuan","middleName":"","lastName":"Sun","suffix":""},{"id":483081000,"identity":"7c43bf57-510e-4a19-822a-1b76192ef5ee","order_by":1,"name":"Fan Yang","email":"","orcid":"","institution":"Institute of Plant Protection, Beijing Academy of Agriculture and Forestry Sciences","correspondingAuthor":false,"prefix":"","firstName":"Fan","middleName":"","lastName":"Yang","suffix":""},{"id":483081001,"identity":"9d7902c2-7eb0-4e18-99d5-ae9528541818","order_by":2,"name":"Ze-Hua Wang","email":"","orcid":"","institution":"Institute of Plant Protection, Beijing Academy of Agriculture and Forestry Sciences","correspondingAuthor":false,"prefix":"","firstName":"Ze-Hua","middleName":"","lastName":"Wang","suffix":""},{"id":483081002,"identity":"65eaea8a-35a5-446c-8cc6-1426b2238d71","order_by":3,"name":"Ren Li","email":"","orcid":"","institution":"Institute of Plant Protection, Beijing Academy of Agriculture and Forestry Sciences","correspondingAuthor":false,"prefix":"","firstName":"Ren","middleName":"","lastName":"Li","suffix":""},{"id":483081003,"identity":"1b0ce4cc-db68-4199-9290-c9272452c25b","order_by":4,"name":"Guang-Hang Qiao","email":"","orcid":"","institution":"Institute of Plant Protection, Beijing Academy of Agriculture and Forestry Sciences","correspondingAuthor":false,"prefix":"","firstName":"Guang-Hang","middleName":"","lastName":"Qiao","suffix":""},{"id":483081004,"identity":"85f91022-c708-4ff9-9f71-7c7043290606","order_by":5,"name":"Qiu-Fei Li","email":"","orcid":"","institution":"Institute of Plant Protection, Beijing Academy of Agriculture and Forestry Sciences","correspondingAuthor":false,"prefix":"","firstName":"Qiu-Fei","middleName":"","lastName":"Li","suffix":""},{"id":483081005,"identity":"dc73feb2-9960-41b4-a8e8-b1d2e318d975","order_by":6,"name":"Yu-Yu Wang","email":"","orcid":"","institution":"College of Plant Protection, Hebei Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"Yu-Yu","middleName":"","lastName":"Wang","suffix":""},{"id":483081006,"identity":"50091fbc-cc55-4012-a460-36324e0c1474","order_by":7,"name":"Shan-Ning Wang","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA7UlEQVRIiWNgGAWjYLACxgYgwQzEH4DYgCQtjDNgWg4QpQWki4cYLQbHzx5++XOHXZ45O+/h17ZtNvLmDLwHH3/Ap+VMXpo175nkYstmvjTr3LY0w50NfMkG+GwxO5BjZszYxpy44TCPmXFu22HGDQd4zCTwajn/xszwZ1s9RItl2397oBbzH3i13MgxfsDbdhikxfgxY9uBRJAteL1vf+ONGTNv23GwLYw955KTQXolzuDRItmfY/zxZ1t14obzZ4w//Cizs91wvMfwQwUeLUDAJgFnMLIxQJIBAcD8AcH4Q1j5KBgFo2AUjDwAAIrxVSEdNBicAAAAAElFTkSuQmCC","orcid":"","institution":"Institute of Plant Protection, Beijing Academy of Agriculture and Forestry Sciences","correspondingAuthor":true,"prefix":"","firstName":"Shan-Ning","middleName":"","lastName":"Wang","suffix":""}],"badges":[],"createdAt":"2025-06-24 02:23:15","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6960668/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6960668/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s10340-025-01987-y","type":"published","date":"2025-12-05T15:58:21+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":86550378,"identity":"41e381a5-ba87-4ac1-aa8d-ba7f435888d1","added_by":"auto","created_at":"2025-07-12 03:32:35","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":17283,"visible":true,"origin":"","legend":"\u003cp\u003ePercent \u003cem\u003eTrichopria drosophilae\u003c/em\u003e females in each arm of an olfactometer tube in choice bioassays with volatile cues produced by blueberries attacked by \u003cem\u003eDrosophila suzukii\u003c/em\u003e (SWD-attacked), undamaged control or clean air. The panels show the arm choice at the end of a 5-min test period, respectively. P values for the binomial test are reported. Red \"ns\" denotes no significant difference (\u003cem\u003ep\u003c/em\u003e ≥ 0.05), while * indicates significant differences at \u003cem\u003ep\u003c/em\u003e\u0026lt; 0.05.\u003c/p\u003e","description":"","filename":"image1.png","url":"https://assets-eu.researchsquare.com/files/rs-6960668/v1/033714f98a627802735e24ea.png"},{"id":86550769,"identity":"cf92a402-31c1-46db-90bb-9ab88518f290","added_by":"auto","created_at":"2025-07-12 03:40:35","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":37545,"visible":true,"origin":"","legend":"\u003cp\u003eNon-metric multidimensional scaling (NMDS) ordination based on Bray–Curtis dissimilarities of the volatile organic compounds (VOCs) from cherries attacked by \u003cem\u003eDrosophila suzukii \u003c/em\u003e(SWD-attacked, N = 8, red dots); VOCs from physically-damaged cherries (N = 8, blue dots) and from undamaged fruit (Control, N = 8, green dots). Stress value = 0.183\u003c/p\u003e","description":"","filename":"image2.png","url":"https://assets-eu.researchsquare.com/files/rs-6960668/v1/8e71e6016c430b5b4cff119f.png"},{"id":86550767,"identity":"2d70c685-43f3-4d60-8ae6-4ac0a447f7a7","added_by":"auto","created_at":"2025-07-12 03:40:35","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":37098,"visible":true,"origin":"","legend":"\u003cp\u003eElectroantennogram (EAG) responses of \u003cem\u003eTrichopria drosophilae\u003c/em\u003e females to different VOCs. Data represent mean ± SE. Different lowercase letters above bars indicate significant differences (\u003cem\u003ep\u003c/em\u003e \u0026lt; 0.05, Kruskal-Wallis test followed by Dunn's post-hoc test).\u003c/p\u003e","description":"","filename":"image3.png","url":"https://assets-eu.researchsquare.com/files/rs-6960668/v1/7a17a31929bd4205ba3ca40d.png"},{"id":86550374,"identity":"25c9d9bc-50dd-4671-a660-f4fc9c8e0d2c","added_by":"auto","created_at":"2025-07-12 03:32:35","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":59465,"visible":true,"origin":"","legend":"\u003cp\u003eOlfactory preference of \u003cem\u003eTrichopria drosophilae\u003c/em\u003e females in response to volatile compounds at varying concentrations (1, 10, and 100 μg/μL) in Y-tube olfactometer assays. Numbers beside bars indicate the count of females selecting test compounds vs. control. Red \"ns\" denotes no significant difference (\u003cem\u003ep\u003c/em\u003e ≥ 0.05), while * and ** indicate significant differences at \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.05 and \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.01, respectively.\u003c/p\u003e","description":"","filename":"image4.png","url":"https://assets-eu.researchsquare.com/files/rs-6960668/v1/adb1a3ae5b592e5940041cf8.png"},{"id":86550373,"identity":"418a24dd-9ce4-4fac-bbb5-651022aa337d","added_by":"auto","created_at":"2025-07-12 03:32:35","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":59650,"visible":true,"origin":"","legend":"\u003cp\u003eExperimental setup and parasitism rates in cage assays testing volatile effects on \u003cem\u003eTrichopria drosophilae\u003c/em\u003e parasitizing \u003cem\u003eDrosophila suzukii pupae\u003c/em\u003e. (A) Schematic representation of the cage assay design. (B) Mean (± SE) parasitism rates of \u003cem\u003eT. drosophilae\u003c/em\u003e on \u003cem\u003eD. suzukii\u003c/em\u003e pupae after exposure to 10 mg/mL benzyl alcohol, ethyl acetate, or isoamyl acetate dissolved in hexane, or hexane alone (control). Asterisks denote significant differences from the control group (\u003cem\u003ep\u003c/em\u003e\u0026lt; 0.05).\u003c/p\u003e","description":"","filename":"image5.png","url":"https://assets-eu.researchsquare.com/files/rs-6960668/v1/007f4059d3e62fdafe6349ea.png"},{"id":97724673,"identity":"72386042-87de-4a23-8efd-8b852a549740","added_by":"auto","created_at":"2025-12-08 16:13:17","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1478074,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6960668/v1/5b55e407-3588-4873-8893-feb81f28469f.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Host-associated volatile cues drive foraging behavior of Trichopria drosophilae toward Drosophila suzukii- infested fruits","fulltext":[{"header":"Introduction","content":"\u003cp\u003ePlant volatiles serve as pivotal chemical signaling molecules that play a central regulatory role in tri-trophic interactions (plant-herbivore-parasitoid systems) (Vet and Dicke \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e1992\u003c/span\u003e; Becker et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Ali et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), particularly by mediating parasitoid host-searching behavior and thereby influencing trophic cascades in ecosystems (Ayelo et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2021b\u003c/span\u003e; G\u0026oacute;mez-Cabezas et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2023\u003c/span\u003e: Rossetti et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Among these volatiles, herbivore-induced plant volatiles (HIPVs) are particularly critical, as they facilitate indirect plant defense by attracting natural enemies such as parasitoids and predators, thereby reducing herbivore damage (Xiu et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Ayelo et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2021a\u003c/span\u003e; Yasa et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eFruit volatile profiles undergo dynamic changes in response to ripening stages (Bi et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) and physical damage (Beck et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Notably, pathogen or insect infestation significantly alters these profiles, generating chemical cues that guide herbivores to hosts (Mohammed et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Guo et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2023a\u003c/span\u003e). For instance, the parasitoid \u003cem\u003eAphytis melinus\u003c/em\u003e is strongly attracted to D-limonene and β-ocimene emitted from \u003cem\u003eAonidiella aurantii\u003c/em\u003e-infested lime fruits, suggesting these volatiles play a key role in host location (Mohammed et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Similarly, fungal-infected apples display shifted volatile organic compound (VOC) patterns, characterized by elevated attractants (e.g., ethyl 2-methylbutyrate) and suppressed repellents, ultimately enhancing oviposition preference in \u003cem\u003eConogethes punctiferalis\u003c/em\u003e (yellow peach moth). Key VOCs such as amyl 2-methylbutyrate and heptacosane have been identified as primary drivers of this behavioral response (Guo et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2023a\u003c/span\u003e). Therefore, identifying the key bioactive compounds emitted by insect-infested fruits that significantly influence parasitoid behavior is of critical importance.\u003c/p\u003e\u003cp\u003eParasitic wasps integrate host-specific chemical cues with background odors to locate hosts efficiently (Schr\u0026ouml;der and Hilker \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). For instance, \u003cem\u003eDiachasmimorpha longicaudata\u003c/em\u003e, a parasitoid wasp targeting \u003cem\u003eCeratitis capitata\u003c/em\u003e, exhibits strong attraction to volatiles emitted from infested oranges, with six key compounds (D-limonene, acetophenone, linalool, nonanal, decanal, and eugenol) identified as critical attractants (Devescovi et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Similarly, behavioral studies reveal that \u003cem\u003eTrichopria anastrephae\u003c/em\u003e is highly responsive to volatiles from \u003cem\u003eD. suzukii\u003c/em\u003e-infested strawberries (containing eggs, larvae, or pupae) and overripe fruits, underscoring the pivotal role of host-associated chemical signals in parasitoid foraging (Kr\u0026uuml;ger et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Notably, \u003cem\u003eGanaspis brasiliensis\u003c/em\u003e G1, a specialized parasitoid of \u003cem\u003eDrosophila suzukii\u003c/em\u003e, dynamically adjusts its response to chemical cues (VOCs and cuticular hydrocarbons), preferentially targeting infested ripening fruits while avoiding decaying substrates. This shift from attraction to repulsion during larval development aligns with its ecological niche as a precision biocontrol agent (Giorgini et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Collectively, these findings highlight the central role of odor-mediated tritrophic interactions in enhancing parasitoid foraging efficacy within agricultural ecosystems.\u003c/p\u003e\u003cp\u003e\u003cem\u003eTrichopria drosophilae\u003c/em\u003e is a globally distributed pupal endoparasitoid that targets various fruit fly species, including the invasive pest \u003cem\u003eD. suzukii\u003c/em\u003e (Chabert et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Mazzetto et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Chen et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Lisi et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2024a\u003c/span\u003e). Unlike traditional fruit flies (e.g., \u003cem\u003eDrosophila melanogaster\u003c/em\u003e) that infest rotting fruits, \u003cem\u003eD. suzukii\u003c/em\u003e (spotted-wing drosophila, SWD) possesses a serrated ovipositor, enabling it to damage ripening fruits at early developmental stages, causing severe economic losses to soft-skinned stone fruits such as cherries and blueberries (through inducing rot and mold development in fresh fruits) (Lee et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Asplen et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Haye et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Labbetoul et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Laboratory studies have demonstrated \u003cem\u003eT. drosophilae\u003c/em\u003e's strong preference for SWD (Wang et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Yi et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), with enhanced offspring fitness when parasitizing this host (Boycheva Woltering et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Jarrett et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Gonzalez-Cabrera et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Field trials in Mexican orchards further revealed that releasing 4,500 parasitoids per hectare reduced SWD populations by 50% (Gonzalez-Cabrera et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). These findings highlight the potential of T. drosophilae as a biocontrol agent against SWD within an Integrated Pest Management (IPM) framework, offering a sustainable tool to reduce chemical pesticide use and thereby limiting adverse effects on non-target organisms (Desneux et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). However, its implementation must be carefully evaluated in IPM packages where some pesticide applications may still be necessary, to ensure compatibility and effectiveness (e.g. Guo et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2023b\u003c/span\u003e; Fouani et al. 2024; Lisi et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2024b\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe efficacy of parasitic wasps in field-based biological control is often constrained by both temporal (Pfab et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Rossi Stacconi et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) and spatial factors (Rossi Stacconi et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Hogg et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), which explains why their application remains underutilized in the integrated management of \u003cem\u003eD. suzukii\u003c/em\u003e. Deciphering the tritrophic chemical communication network among host fruits, \u003cem\u003eD. suzukii\u003c/em\u003e, and parasitic wasps could identify key attractant compounds, thereby providing critical insights for field application strategies. This study addresses three pivotal scientific questions: (i) Whether SWD infestation alters the volatile organic compound (VOC) profile of sweet cherries; (ii) The ability of \u003cem\u003eT. drosophilae\u003c/em\u003e to discriminate SWD-infested cherries via VOCs; and (iii) Identification of key chemical compounds mediating this behavioral response. Our findings will provide a theoretical foundation for developing VOC-based precision biocontrol strategies against \u003cem\u003eD. suzukii\u003c/em\u003e.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003eInsect collection and rearing\u003c/h2\u003e\u003cp\u003e\u003cem\u003eDrosophila suzukii\u003c/em\u003e (spotted-wing drosophila) and \u003cem\u003eDrosophila melanogaster\u003c/em\u003e (vinegar fly) were collected in 2022 from the cherry experimental base at the Institute of Forestry and Pomology, Beijing Academy of Agriculture and Forestry Sciences (39\u0026deg;41\u0026prime;N, 116\u0026deg;42\u0026prime;E). Their parasitoid \u003cem\u003eTrichopria drosophilae\u003c/em\u003e was concurrently captured at the same location using fermenting fruit-baited traps. Both \u003cem\u003eDrosophila\u003c/em\u003e species were reared according to the artificial diet described by Yang et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2024\u003c/span\u003e, with all insects maintained in climate-controlled chambers (25\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u0026deg;C, 70\u0026thinsp;\u0026plusmn;\u0026thinsp;5% RH, 16:8 h L:D photoperiod). The \u003cem\u003eT. drosophilae\u003c/em\u003e population was propagated using \u003cem\u003eD. melanogaster\u003c/em\u003e pupae as hosts under identical environmental conditions.\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eChemicals\u003c/h3\u003e\n\u003cp\u003eEthyl acetate (CAS: 141-78-6, \u0026ge;\u0026thinsp;99.7%), methyl acetate (CAS: 79-20-9, \u0026ge;\u0026thinsp;99.0%), isoamyl acetate (CAS: 123-92-2, \u0026ge;\u0026thinsp;99.5%), ethyl benzene (CAS: 93-89-0, \u0026ge;\u0026thinsp;99.5%), benzyl alcohol (CAS: 100-51-6, \u0026ge;\u0026thinsp;99.8%), phenylethanol (CAS: 60-12-8, \u0026ge;\u0026thinsp;99.5%), ethanol (CAS: 64-17-5, \u0026ge;\u0026thinsp;99.8%), isovaleric acid (CAS: 503-74-2, \u0026ge;\u0026thinsp;99.0%), acetic acid (CAS: 64-19-7, \u0026ge;\u0026thinsp;99.8%), benzaldehyde (CAS: 100-52-7, \u0026ge;\u0026thinsp;98%), n-hexane (CAS: 110-54-3, \u0026ge;\u0026thinsp;97%), carene (CAS: 4497-92-1, \u0026ge;\u0026thinsp;97%). All chemicals were purchased from Shanghai Macklin Biochemical Technology Co., Ltd.\u003c/p\u003e\n\u003ch3\u003eCherry Fruit Treatment\u003c/h3\u003e\n\u003cp\u003ePurchased fresh cherries (cultivar: Tieton) were gently washed and carefully inspected for insect eggs, physical damage, or abnormalities to ensure they were free from external defects and pest contamination. To induce \u003cem\u003eD. suzukii\u003c/em\u003e infestation, five pairs of adult flies (3\u0026ndash;7 days old) were placed in plastic containers containing approximately 50 g of cherries and sealed for 24 hours. After 24 hours, adult flies were removed, and the number of eggs on each cherry was counted under a microscope (average: 10 eggs per cherry). Cherries were then stored in clean plastic containers with fine mesh covers at 25\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u0026deg;C, 70\u0026thinsp;\u0026plusmn;\u0026thinsp;5% RH, and a 16 L: 8 D light cycle for 7 days. Once larvae pupated, the pupae were removed, and the fruit volatiles were collected. To distinguish the effects of \u003cem\u003eD. suzukii\u003c/em\u003e infestation from physical damage and natural ripening, two controls were used: cherries were punctured 10 times with a sterilized insect needle (0.3 mm diameter) 7 days before volatile collection, and an untreated control group was set under the same conditions without any treatment. Each treatment was repeated eight times, with containers wiped with 75% ethanol for sterility.\u003c/p\u003e\n\u003ch3\u003eCollection of Fruit Volatiles\u003c/h3\u003e\n\u003cp\u003eCherries were placed in a dynamic headspace collection bottle (650 mL). The air pump (Model: CAT-8000) was set to a flow rate of 300 mL/min. The air was previously dehumidified with silica gel and filtered through activated carbon. The volatiles were adsorbed in a glass tube packed with 100 mg of Porapak Q adsorbent (Waters Corporation, USA, 80\u0026ndash;100 mesh) for a collection period of 24 hours with three replicates. The adsorbed volatiles were eluted with 400 \u0026micro;L of n-hexane, concentrated to 150 \u0026micro;L, and stored in a 2 mL vial at -20\u0026deg;C until gas chromatography-mass spectrometry (GC-MS) analysis. Prior to sample concentration, 2 \u0026micro;L of an internal standard solution (carene) was added. VOC collections were made at a temperature of 25\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u0026deg;C, 70\u0026thinsp;\u0026plusmn;\u0026thinsp;5% relative humidity and a photoperiod of 12:12 h (L: D).\u003c/p\u003e\n\u003ch3\u003eChemical Analyses\u003c/h3\u003e\n\u003cp\u003eVolatiles emitted by cherries were analyzed using gas chromatography-mass spectrometry (GC-MS) with an Agilent 7890 GC coupled to a 5975 MSD. The GC was equipped with an HP-5 capillary column (30 m \u0026times; 0.25 mm \u0026times; 0.25 \u0026micro;m, Agilent Technologies). A 1 \u0026micro;L sample was injected in splitless mode, with the injection port set at 250\u0026deg;C and the ion source at 230\u0026deg;C. Electron ionization (EI) was performed with an ionization energy of 70 eV, using helium as the carrier gas at a flow rate of 1 mL/min. The temperature program was as follows: initial temperature 60\u0026deg;C, held for 1 min; ramped at 4\u0026deg;C/min to 200\u0026deg;C, held for 2 min; ramped at 15\u0026deg;C/min to 260\u0026deg;C, held for 5 min, for a total run time of 47 min.\u003c/p\u003e\u003cp\u003eCompounds were identified by matching chromatographic peaks with those in the NIST 8.0 mass spectral library, and confirmed by comparison with authentic standards. The concentrations of volatile compounds released in the treatments (infested by \u003cem\u003eD. suzukii\u003c/em\u003e, mechanically damaged, and control) were quantified based on the peak areas of the internal standard.\u003c/p\u003e\u003cp\u003eConcentration of unknown volatiles = (Concentration of internal standard \u0026times; Peak area of unknown) / Peak area of internal standard.\u003c/p\u003e\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e\u003ch2\u003eOlfactometer Bioassays\u003c/h2\u003e\u003cp\u003eThe olfactory response of female \u003cem\u003eTrichopria drosophilae\u003c/em\u003e (3\u0026ndash;7 days post-emergence) to host fruit location was measured using a \u0026ldquo;Y\u0026rdquo;-shaped olfactometer. The two arms of the olfactometer were 6.0 cm in length, with a straight tube of 9.0 cm in length and an inner diameter of 0.8 cm. The olfactometer was located in a dark opaque chamber (80 cm \u0026times; 80 cm \u0026times; 50 cm) to avoid visual interference. Humidied charcoal-filtered air was pushed through the stimuli and olfactometer with a pump at a rate of 100 mL/min. A 40 W incandescent bulb was positioned directly above the box to ensure uniform light intensity across both testing arms. All bioassays were conducted at 25\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u0026deg;C, 70\u0026thinsp;\u0026plusmn;\u0026thinsp;5% RH and performed between 09:00 and 11:00 am. After every five wasps were tested, the positions of odor sources in the Y-tube olfactometer were swapped, and a new Y-tube was used. At the end of the experiment, all Y-tubes were collectively cleaned with 95% ethanol for subsequent trials.\u003c/p\u003e\u003cp\u003eFor the fruit volatiles, three treatment groups were established for comparison: \u003cem\u003eDrosophila suzukii\u003c/em\u003e-infested fruit vs. clean air, healthy fruit vs. clean air, and \u003cem\u003eD. suzukii\u003c/em\u003e-infested fruit vs. healthy fruit. Subsequently, based on the Electroantennographic (EAG) test results, we randomly selected several compounds and conducted olfactometer tests at different concentrations: 1 \u0026micro;g/\u0026micro;L, 10 \u0026micro;g/\u0026micro;L, and 100 \u0026micro;g/\u0026micro;L. For each treatment, 10 \u0026micro;L of the prepared attractant solution was applied to a filter paper strip (3 cm \u0026times; 1 cm) and placed in one volatile source bottle. An equal volume of the control solvent (n-hexane) was applied to a separate filter paper strip and placed in the other source bottle.\u003c/p\u003e\u003cp\u003eFemale \u003cem\u003eT. drosophilae\u003c/em\u003e was placed at the base of the \u0026ldquo;Y\u0026rdquo; olfactometer, and their behavioral responses were observed and recorded over a 5-minute period. The selection of the odor source in each arm was noted when the female was either within 5 cm of the exit or had passed halfway down an arm and remained there for at least 5 seconds. If no choice was made within the 5-minute observation period, the response was recorded as \u0026ldquo;no reaction\u0026rdquo;.\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eElectroantennographic (EAG)\u003c/h3\u003e\n\u003cp\u003eThe EAG technique was used to assess the antennal selectively and sensitivity of \u003cem\u003eT. drosophilae\u003c/em\u003e females to the selected ten volatile organic compounds (VOCs).\u003c/p\u003e\u003cp\u003eThe antennae of female \u003cem\u003eT. drosophilae\u003c/em\u003e were excised at the base using fine scissors. Immediately afterward, approximately 0.5\u0026ndash;1 mm was trimmed from the tips of the antennae using a surgical blade. The prepared antennae were then connected to the electrodes using a 2 mm-diameter glass capillary filled with PBS (pH\u0026thinsp;=\u0026thinsp;7.2) solution. The electrophysiological signals of antennal responses to stimuli were amplified, output, and collected using a data acquisition controller (IDAC). The collected signals were processed and analyzed using EAG-Pro software.\u003c/p\u003e\u003cp\u003eOnce the baseline signal stabilized, volatile collections were tested. Preliminary experiments were conducted to determine the optimal concentration of standard compounds, which were dissolved in n-hexane. The final concentration gradient was set to 1 \u0026micro;g/\u0026micro;L for the electroantennographic (EAG) response measurements. For each volatile, 10 \u0026micro;L was taken with a micropipette and applied to a clean 1 \u0026times; 3 cm\u0026sup2; filter paper. After 20 s at room temperature to allow evaporation of n-hexane, the paper was placed into a Pasteur pipette (diameter 2 mm, length 12 cm). A control sample was prepared by adding an equal volume of n-hexane to another filter paper under identical conditions. Each volatile was tested on 18 antennae with active responses (one antenna per wasp). There was a 30 s interval between each stimulation, and control measurements were conducted before and after testing each volatile. The relative EAG response value of the antenna to each odorant was calculated as: EAG response value\u0026thinsp;=\u0026thinsp;Volatile response value - (Pre-test control response value\u0026thinsp;+\u0026thinsp;Post-test control response value) / 2.\u003c/p\u003e\n\u003ch3\u003eCage experiment\u003c/h3\u003e\n\u003cp\u003eThe experiment was conducted in 35 \u0026times; 35 \u0026times; 35 cm rearing cages, each equipped with two diagonally placed self-made rubber septum dispensers. For treatment groups, 200 \u0026micro;L of benzyl alcohol, ethyl acetate, or isoamyl acetate (10 \u0026micro;g/\u0026micro;L, selected based on preliminary Y-tube olfactometer screening of bioactive compounds) was applied to the dispensers, while an equal volume of n-hexane served as the solvent control. Twenty-five 1-day-old pupae of \u003cem\u003eD. suzukii\u003c/em\u003e were introduced as hosts, followed by the release of five pairs of \u003cem\u003eT. drosophilae\u003c/em\u003e adults (3\u0026ndash;7 days post-emergence). The dispenser and fly pupae were placed in an open 6 cm diameter Petri dish. A 6 cm Petri dish containing a cotton ball soaked in 10% honey solution was positioned at the center of the cage to provide nutrition for the parasitoid wasps. After 24 h of exposure, pupae were collected to quantify parasitism. After collection, the fly pupae were maintained at 25\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u0026deg;C, 70\u0026thinsp;\u0026plusmn;\u0026thinsp;5% relative humidity, and a 16:8 h (light:dark) photoperiod until adult emergence. To confirm parasitism, pupae that failed to eclose were subjected to PCR amplification using species-specific primers to detect \u003cem\u003eT. drosophilae\u003c/em\u003e DNA (unpublished content), allowing quantification of parasitized hosts. Each treatment was independently replicated eight times, with fresh hosts and parasitoids used for every replicate. Parasitism rate was calculated as:\u003c/p\u003e\u003cp\u003eParasitism rate (%)\u0026thinsp;=\u0026thinsp;Number of parasitized hosts /Total hosts released \u0026times;100%\u003c/p\u003e\u003cp\u003eThe three volatiles were tested separately, with n-hexane serving as a unified solvent control to eliminate confounding effects.\u003c/p\u003e\u003cdiv id=\"Sec11\" class=\"Section2\"\u003e\u003ch2\u003eStatistical Analysis\u003c/h2\u003e\u003cp\u003eStatistical analyses were conducted to evaluate female \u003cem\u003eT. drosophilae\u003c/em\u003e behavioral responses through binomial tests examining both host preference (SWD-attacked vs. Undamaged control, SWD-attacked vs. Clean air, and Undamaged control vs. Clean air) and concentration-dependent responses to ten volatile compounds (1, 10, and 100 \u0026micro;g/\u0026micro;L). Volatile organic compound profiles were characterized using NMDS ordination based on Bray-Curtis dissimilarity (stress\u0026thinsp;=\u0026thinsp;0.183), with statistical significance assessed through PERMANOVA (999 permutations) followed by Benjamini-Hochberg-adjusted pairwise comparisons when appropriate. Electroantennographic responses were analyzed with Kruskal-Wallis tests and Dunn's post hoc tests for significant findings (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05), while treatment effects in cage experiments were evaluated using paired t-tests. All statistical analyses were performed using R (Version 4.4.2) (1999\u0026ndash;2024 R Core Team).\u003c/p\u003e\u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003e\u003cb\u003ePreference of\u003c/b\u003e \u003cb\u003eTrichopria drosophilae\u003c/b\u003e \u003cb\u003efemales for differently treated cherry fruits\u003c/b\u003e\u003c/p\u003e\u003cp\u003eThe binomial tests revealed significant preferences in \u003cem\u003eTrichopria drosophilae\u003c/em\u003e for certain odor sources. When given a choice between damaged fruits and healthy fruits, the wasps selected damaged fruits significantly more often (62.9% preference, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.041). Similarly, in the damaged fruits vs air comparison, the wasps showed a stronger attraction to damaged fruits (64.3% preference, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.022). However, no significant preference was observed between healthy fruits and air (55.7% preference, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.403). These results indicate that \u003cem\u003eT. drosophilae\u003c/em\u003e exhibited a clear preference for volatiles from damaged cherry fruits over both healthy fruits and air, while showing no significant bias when choosing between healthy fruits and air.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cdiv id=\"Sec13\" class=\"Section2\"\u003e\u003ch2\u003eAnalysis of volatile organic compound (VOC) differences in differently treated cherry fruits\u003c/h2\u003e\u003cp\u003eA total of 48 volatile compounds were identified in cherry samples. The highest diversity of compounds, including esters, alkenes, and alcohols, was detected in cherries infested by \u003cem\u003eD. suzukii\u003c/em\u003e, with 11 compounds being exclusively present in the volatiles of infested cherries. The concentration of 4-ethylbenzaldehyde was significantly higher in \u003cem\u003eD. suzukii\u003c/em\u003e-infested cherries (0.86\u0026thinsp;\u0026plusmn;\u0026thinsp;0.22 ng/\u0026micro;L) compared to the control group (0.20\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11 ng/\u0026micro;L). In contrast, hexacos-1-ene exhibited a higher concentration in the physical damage group (1.42\u0026thinsp;\u0026plusmn;\u0026thinsp;0.49 ng/\u0026micro;L) but was reduced in the \u003cem\u003eD. suzukii\u003c/em\u003e-infested group (0.35\u0026thinsp;\u0026plusmn;\u0026thinsp;0.29 ng/\u0026micro;L). Additionally, ester compounds such as ethyl acetate, methyl acetate, and isoamyl acetate were present at higher concentrations in \u003cem\u003eD. suzukii\u003c/em\u003e-infested cherries, while they were either undetected or present at lower levels in the control group. These results demonstrate that \u003cem\u003eD. suzukii\u003c/em\u003e infestation significantly alters the concentrations of specific compounds in cherry fruits (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eVolatile organic compounds were quantified in sweet cherry samples subjected to three experimental treatments: \u003cem\u003eD. suzukii\u003c/em\u003e infestation, mechanical injury, and undamaged controls (n\u0026thinsp;=\u0026thinsp;8 per group). The specific concentrations (ng/\u0026micro;L) of detected volatiles are presented in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e. The ten volatile compounds\u0026mdash;ethyl acetate, methyl acetate, isoamyl acetate, ethyl benzoate, benzyl alcohol, phenethyl alcohol, ethanol, 3-methylbutanoic acid, acetic acid, and benzaldehyde\u0026mdash;were selected for further investigation due to their significantly higher occurrence frequency and concentration levels in \u003cem\u003eD. suzukii\u003c/em\u003e-infested cherries. Most of these compounds were exclusively detected in infested samples, while a few, though present in both infested and healthy cherries but not in physically damaged ones, exhibited markedly elevated concentrations in \u003cem\u003eD. suzukii\u003c/em\u003e-infested fruits compared to their trace levels in undamaged or mechanically injured samples.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eAmounts of volatile organic compounds (VOCs) found in collections from cherries subjected to the different treatments: attacked by \u003cem\u003eDrosophila suzukii\u003c/em\u003e (SWD-attacked); physical damage and natural fruit ripening (undamaged control). Compounds authenticated by reference standards are indicated with an asterisk (*).\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"8\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eCompound\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eCAS\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"6\" nameend=\"c8\" namest=\"c3\"\u003e\u003cp\u003eAmount (\u0026micro;g/25 g/24 h; mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard error)\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eSWD-attacked\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eN\u003c/em\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePhysically damaged\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cem\u003eN\u003c/em\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003eUndamaged control\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003e\u003cem\u003eN\u003c/em\u003e\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eEsters\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEthyl valerate\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e539-82-2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.06\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eIsoamyl acetate*\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e123-92-2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.48\u0026thinsp;\u0026plusmn;\u0026thinsp;0.33\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEthyl benzoate*\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e93-89-0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.50\u0026thinsp;\u0026plusmn;\u0026thinsp;0.31\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMethyl 3,5-dimethylbenzoate\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e25081-39-4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.32\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.29\u0026thinsp;\u0026plusmn;\u0026thinsp;0.10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.09\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMethyl 4-ethylbenzoate\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e7364-20-7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.23\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.21\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.05\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMethyl hexadecanoate\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e112-39-0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.16\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.14\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.05\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEthyl acetate*\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e141-78-6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.36\u0026thinsp;\u0026plusmn;\u0026thinsp;0.20\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMethyl stearate\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e112-61-8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.26\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.25\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.12\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMethyl acetate*\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e79-20-9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.25\u0026thinsp;\u0026plusmn;\u0026thinsp;0.10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.01\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eisoPropyl Myristate\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e110-27-0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.12\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.11\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.04\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMethyl 14-methylpentadecanoate\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5129-60-2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.02\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAcetyl tributyl citrate\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e77-90-7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.21\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.05\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEthyl isovalerate\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e108-64-5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.04\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEthyl Palmitate\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e628-97-7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.07\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEthyl 2-methylbutyrate\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e7452-79-1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.02\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTriethyl citrate\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e77-93-0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMethyl Salicylate\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e119-36-8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.03\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eAlkenes\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eStyrene\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e100-42-5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.12\u0026thinsp;\u0026plusmn;\u0026thinsp;0.10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.19\u0026thinsp;\u0026plusmn;\u0026thinsp;0.14\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.01\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePinene\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e7785-70-8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.22\u0026thinsp;\u0026plusmn;\u0026thinsp;0.14\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.24\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.07\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e1-Dodecene\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e112-41-4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.05\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.04\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e1-Hexadecene\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e629-73-2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.06\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e1-Heptadecene\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e6765-39-5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.05\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSqualene\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e111-02-4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.73\u0026thinsp;\u0026plusmn;\u0026thinsp;1.14\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.61\u0026thinsp;\u0026plusmn;\u0026thinsp;0.50\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.15\u0026thinsp;\u0026plusmn;\u0026thinsp;0.18\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e1-Docosanol\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1599-67-3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.07\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e1-Eicosene\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e3452/7/1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.10\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.04\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e1-Hexacosene\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e18835-33-1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.35\u0026thinsp;\u0026plusmn;\u0026thinsp;0.29\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.42\u0026thinsp;\u0026plusmn;\u0026thinsp;0.49\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.52\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e1,3,5,7-Cyclooctatetraene\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e629-20-9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.12\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.05\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e17-Pentatriacontene\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e6971-40-0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.01\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eα - Pinene\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e80-56-8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.06\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.02\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e(+)-Camphene\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e79-92-5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.01\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e1-Nonadecene\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e18435-45-5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.16\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eβ-pinene\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e18172-67-3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.10\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e(9Z)-9-Tricosene\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e27519-02-4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.06\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.04\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e(+)-Limonene\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5989-27-5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.44\u0026thinsp;\u0026plusmn;\u0026thinsp;0.31\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.26\u0026thinsp;\u0026plusmn;\u0026thinsp;0.14\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eAlcohols\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBenzyl alcohol*\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e100-51-6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.28\u0026thinsp;\u0026plusmn;\u0026thinsp;0.23\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePhenethyl alcohol*\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1960/12/8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.17\u0026thinsp;\u0026plusmn;\u0026thinsp;0.18\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.17\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e(+)-Cedrol\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e77-53-2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.11\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.07\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.03\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEthanol*\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e64-17-5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.37\u0026thinsp;\u0026plusmn;\u0026thinsp;0.43\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eAcids\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNonahexacontanoic acid\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e40710-32-5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.08\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.04\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.04\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e3-Methylbutanoic acid*\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e503-74-2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.14\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAcetic acid*\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e64-19-7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.26\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eStearic acid\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1957/11/4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eAromatics\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ep-Xylene\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e106-42-3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.04\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.04\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.02\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNaphthalene\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e91-20-3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.39\u0026thinsp;\u0026plusmn;\u0026thinsp;0.10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.64\u0026thinsp;\u0026plusmn;\u0026thinsp;0.55\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eo-xylene\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e95-47-6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.16\u0026thinsp;\u0026plusmn;\u0026thinsp;0.15\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.05\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.03\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eAldehydes\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDecanal\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e112-31-2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.81\u0026thinsp;\u0026plusmn;\u0026thinsp;0.49\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.46\u0026thinsp;\u0026plusmn;\u0026thinsp;1.25\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.22\u0026thinsp;\u0026plusmn;\u0026thinsp;0.17\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBenzaldehyde*\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e100-52-7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.72\u0026thinsp;\u0026plusmn;\u0026thinsp;0.74\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.01\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e4-Ethylbenzaldehyde\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4748-78-1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.86\u0026thinsp;\u0026plusmn;\u0026thinsp;0.22\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.95\u0026thinsp;\u0026plusmn;\u0026thinsp;0.34\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.20\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eVOC components are classified in groups and \u003cem\u003eN\u003c/em\u003e indicates the number of samples in which a compound was found.\u003c/p\u003e\u003cp\u003eNon-metric multidimensional scaling (NMDS) based on Bray-Curtis dissimilarity matrices revealed significant spatial separation in volatile organic compound (VOC) profiles among the three treatment groups (stress\u0026thinsp;=\u0026thinsp;0.183, acceptable per Kruskal\u0026rsquo;s criterion). Non-metric multidimensional scaling (NMDS) analysis revealed distinct clustering patterns among the three experimental groups (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). \u003cem\u003eDrosophila suzukii\u003c/em\u003e-infested fruits (depicted as red dots) exhibited significant clustering in the negative NMDS1 dimension (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SE: -0.530\u0026thinsp;\u0026plusmn;\u0026thinsp;0.104), whereas undamaged control fruits (green dots) occupied the positive NMDS1 quadrant (0.565\u0026thinsp;\u0026plusmn;\u0026thinsp;0.126). Fruits subjected to mechanical injury (blue dots) demonstrated intermediate positioning along the NMDS1 axis (-0.0355\u0026thinsp;\u0026plusmn;\u0026thinsp;0.101), suggesting a transitional state between infestation and control conditions.\u003c/p\u003e\u003cp\u003eThese results indicate that VOC profiles of SWD-attacked and undamaged control fruits are distinctly separated along NMDS1, while physically damaged fruits exhibit an intermediate profile, suggesting a gradient in VOC composition based on treatment type.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eThe key characteristics of spatial distribution revealed that the \u003cem\u003eD. suzukii\u003c/em\u003e-attacked (SWD-attacked) group exhibited specific separation, with its mean NMDS1 value (-0.530) differing by 1.095 units from that of the healthy group (0.565), indicating directional changes in volatiles induced by pest infestation (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). In contrast, the physically damaged group showed a heterogeneous response, as evidenced by the significantly higher standard error of the NMDS2 axis (0.209) compared to the healthy group (0.151), suggesting greater randomness in volatile release patterns (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). These findings highlight distinct spatial distribution patterns in volatile compounds between pest-infested, physically damaged, and healthy cherry fruits.\u003c/p\u003e\u003cp\u003eThe volatile compound composition significantly differed among the treatment groups (R\u0026sup2; = 0.328, F\u0026thinsp;=\u0026thinsp;5.125, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.001) (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Specifically, the \u003cem\u003eD. suzukii\u003c/em\u003e-attacked (SWD-attacked) group showed highly significant differences compared to the undamaged control group (R\u0026sup2; = 0.397, F\u0026thinsp;=\u0026thinsp;9.220, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.0015), and the physically damaged group also exhibited significant differences relative to the undamaged control group (R\u0026sup2; = 0.282, F\u0026thinsp;=\u0026thinsp;5.508, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.0015) (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). However, no significant differences were observed between the SWD-attacked group and the physically damaged group (R\u0026sup2; = 0.124, F\u0026thinsp;=\u0026thinsp;1.985, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.094) (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). These results indicated that both \u003cem\u003eD. suzukii\u003c/em\u003e infestation and physical damage distinctly alter the volatile profiles of cherries, with \u003cem\u003eD. suzukii\u003c/em\u003e infestation eliciting a more pronounced effect compared to undamaged controls.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eMean NMDS coordinates\u0026thinsp;\u0026plusmn;\u0026thinsp;standard error of VOCs in cherry fruits under different treatments and results of pairwise comparisons\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"7\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTreatment\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNMDS1 (Mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SE)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNMDS2 (Mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SE)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003ePairwise comparison groups\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eR\u0026sup2;\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eF value\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003eCorrected p-values\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSWD-attacked\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c2\"\u003e\u003cp\u003e-0.530\u0026thinsp;\u0026plusmn;\u0026thinsp;0.104\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e\u003cp\u003e-0.192\u0026thinsp;\u0026plusmn;\u0026thinsp;0.111\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003evs Undamaged control\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.3971\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e9.22\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e0.0015\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePhysically damaged\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c2\"\u003e\u003cp\u003e-0.0355\u0026thinsp;\u0026plusmn;\u0026thinsp;0.101\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e\u003cp\u003e0.0058\u0026thinsp;\u0026plusmn;\u0026thinsp;0.209\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003evs SWD-attacked\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.1241\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e1.9845\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e0.0940\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eUndamaged control\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c2\"\u003e\u003cp\u003e0.565\u0026thinsp;\u0026plusmn;\u0026thinsp;0.126\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e\u003cp\u003e0.186\u0026thinsp;\u0026plusmn;\u0026thinsp;0.151\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003evs Physically damaged\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.2823\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e5.5075\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e0.0015\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003cb\u003eEAG responses of\u003c/b\u003e \u003cb\u003eTrichopria drosophilae\u003c/b\u003e \u003cb\u003efemales to cherry VOCs\u003c/b\u003e\u003c/p\u003e\u003cp\u003eThrough analysis, ten compounds with significantly increased volatile emissions in the \u003cem\u003eD. suzukii\u003c/em\u003e-infested treatment were selected to further investigate the electrophysiological responses of \u003cem\u003eT. drosophilae\u003c/em\u003e to these compounds. The electrophysiological responses (EAG) to ten volatile compounds varied significantly (χ\u0026sup2; = 65.76, df\u0026thinsp;=\u0026thinsp;9, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). Ethyl acetate elicited the strongest response (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SE: 0.0393\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0057), followed by benzaldehyde (0.0233\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0044) and methyl acetate (0.0170\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0037). Ethyl benzoate (0.0136\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0024) and isoamyl acetate (0.0120\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0025) showed moderate responses, with isoamyl acetate being significantly higher than phenethyl alcohol (0.0034\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0021, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.022) and ethanol (-0.0006\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0012, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.001). Acetic acid (0.0078\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0018) and 3-methylbutanoic acid (0.0079\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0024) induced weaker responses, while benzyl alcohol (0.0045\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0025) showed minimal activity. These findings indicate that esters (ethyl acetate, ethyl benzoate, isoamyl acetate, methyl acetate) and aromatic aldehydes (benzaldehyde) generally evoked stronger EAG responses than alcohols (benzyl alcohol, ethanol, phenethyl alcohol) or carboxylic acids (acetic acid, 3-Methylbutanoic acid), with ethyl acetate being the most potent stimulant (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003cb\u003ePreference of\u003c/b\u003e \u003cb\u003eTrichopria drosophilae\u003c/b\u003e \u003cb\u003efemales for VOCs at different concentrations\u003c/b\u003e\u003c/p\u003e\u003cp\u003eAdditionally, we further investigated whether \u003cem\u003eT. drosophilae\u003c/em\u003e exhibited behavioral responses to these ten compounds. The Y-tube olfactometer tests revealed significant behavioral responses to specific volatile compounds. Notably, isoamyl acetate elicited strong attraction at 10 \u0026micro;g/\u0026micro;L concentration (72.0% response rate, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.0026), while ethyl acetate showed dose-dependent effects with significant responses at both 100% (70.0%, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.0066) and 10% (66%, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.0328) concentrations. Benzyl alcohol demonstrated moderate attraction at 10% concentration (66%, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.0328).\u003c/p\u003e\u003cp\u003eConversely, 3-methylbutanoic acid showed significant repellency at 10% concentration (34% response rate, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.0328). Control compounds (ethanol, acetic acid, etc.) generally showed neutral responses (44\u0026ndash;54% response rates, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.05). The complete response patterns suggest concentration-dependent effects, with intermediate concentrations (10%) often producing the strongest behavioral responses compared to higher (100%) or lower (1%) concentrations.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003cb\u003eOviposition Response of\u003c/b\u003e \u003cb\u003eTrichopria drosophilae\u003c/b\u003e \u003cb\u003eFemales to VOCs\u003c/b\u003e\u003c/p\u003e\u003cp\u003eBehavioral response assays of the ten target volatile compounds identified three candidates for validation in cage experiments. The parasitism rates under different chemical treatments showed significant variations compared to the controls, n-hexane. Benzyl alcohol significantly increased parasitism rates compared to controls (t\u003csub\u003e14\u003c/sub\u003e = -2.34, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.035). Similarly, ethyl acetate showed a significant treatment effect (t\u003csub\u003e14\u003c/sub\u003e = -2.21, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.044). The most pronounced effect was observed in the isoamyl acetate group, where parasitism rates were significantly higher in treated samples (t\u003csub\u003e14\u003c/sub\u003e = -2.71, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.017). These results demonstrate that both benzyl alcohol, ethyl acetate and isoamyl acetate significantly promote parasitism, with isoamyl acetate showing the strongest enhancing effect among the tested compounds (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eOur results demonstrate that \u003cem\u003eTrichopria drosophilae\u003c/em\u003e specifically responds to \u003cem\u003eDrosophila suzukii\u003c/em\u003e-induced HIPVs, showing strong attraction to infested fruits but not to healthy ones. Chemical analyses revealed distinct volatile profiles in infested fruits, particularly enriched esters and aromatic aldehydes, which were further confirmed as key attractants through electrophysiological and behavioral assays. Importantly, these HIPVs significantly enhanced parasitism rates, supporting their potential application in \u003cem\u003eD. suzukii\u003c/em\u003e biological control.\u003c/p\u003e\u003cp\u003eEvidence shows that \u003cem\u003eT. drosophilae\u003c/em\u003e selectively targets \u003cem\u003eD. suzukii\u003c/em\u003e-infested cherry fruits (62.9% vs. healthy; 64.3% vs. air), confirming its HIPV detection capability. Healthy fruits and air showed no difference in attraction (55.7%, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.403), indicating that the parasitoid's response depends on herbivore-induced volatiles rather than fruit emissions. Volatile profiling revealed distinct chemical shifts in infested fruits (e.g., elevated esters and 4-ethylbenzaldehyde), with NMDS clearly separating infested and healthy samples (NMDS1: -0.530 vs. 0.565). Mechanically damaged fruits emitted intermediate volatiles, suggesting HIPV induction involves herbivore-specific factors (e.g., salivary enzymes or microbial symbionts). This mechanistic parallel is well-established in other systems, particularly \u003cem\u003eChilo suppressalis\u003c/em\u003e-infested rice where herbivore oral secretions trigger HIPV emission to recruit parasitoids(Yu et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), collectively demonstrating HIPVs' evolutionarily conserved function in mediating tritrophic interactions through herbivore-specific chemical cues. Importantly, herbivore-associated microbial communities to \u003cem\u003eD. suzukii\u003c/em\u003e (Labbetoul et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2025\u003c/span\u003e) might modulate HIPVs released, representing a sophisticated co-evolutionary adaptation where microbial modulation of volatile profiles directly impacts parasitoid host-seeking efficiency while navigating complex host defense systems. These insights advance our understanding of HIPV-mediated ecological interactions and highlight potential applications for enhancing biological control strategies in integrated pest management.\u003c/p\u003e\u003cp\u003e\u003cem\u003eTrichopria drosophilae\u003c/em\u003e exhibits a unique high sensitivity to ester compounds. This aligns with findings that acetate esters emitted by microorganisms (e.g., \u003cem\u003eSaccharomyces cerevisiae\u003c/em\u003e) attract parasitoids such as \u003cem\u003eT. drosophilae\u003c/em\u003e, likely facilitating host location by mirroring their hosts' olfactory preferences (Đurović et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Similarly, GC-EAD analysis revealed that other female parasitoids (\u003cem\u003eLeptopilina boulardi\u003c/em\u003e and \u003cem\u003eTrichopria anastrephae\u003c/em\u003e) respond antennally to fruit esters (ethyl butanoate, methyl hexanoate, ethyl hexanoate) from \u003cem\u003eD. suzukii\u003c/em\u003e-infested strawberries, confirming esters as critical foraging cues (Tri\u0026ntilde;anes et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). However, the role of ester concentration in behavioral responses is context-dependent: while low concentrations may guide parasitoids, high concentrations of compounds like isoamyl acetate can deter \u003cem\u003eD. suzukii\u003c/em\u003e females, possibly signaling overripe fruit unsuitable for oviposition (Cha et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Revadi et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). These findings collectively highlight the dual role of ester volatiles in mediating trophic interactions, where their ecological function depends on both chemical identity and environmental context.\u003c/p\u003e\u003cp\u003eOur study provides a clear example that strong electrophysiological responses to a volatile (e.g., 3-methylbutanoic acid at 1 \u0026micro;g/\u0026micro;L) do not necessarily predict behavioral attraction. Specifically, 3-methylbutanoic acid elicited antennal responses at 1 \u0026micro;g/\u0026micro;L but acted as a repellent at 10 \u0026micro;g/\u0026micro;L in \u0026ldquo;Y\u0026rdquo;-shaped olfactometer. This finding contrasts with observations in \u003cem\u003eTrissolcus basalis\u003c/em\u003e, an egg parasitoid of \u003cem\u003eNezara viridula\u003c/em\u003e, where the same compound was attractive at similar concentrations when emitted from buckwheat flowers. Notably, \u003cem\u003eTrissolcus basalis\u003c/em\u003e displayed strong antennal responses to 3-methylbutanoic acid at 100 \u0026micro;g/\u0026micro;L, suggesting that behavioral outcomes may diverge even when electrophysiological detection occurs (Foti et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Concentration-dependent effects likely play a critical role\u0026mdash;while certain low doses may attract or stimulate antennal detection, intermediate or high concentrations could either enhance attraction or trigger repellency, with no consistent linear pattern across species or compounds. For instance, (Z)-3-hexenol attracts \u003cem\u003eTrichogramma chilonis\u003c/em\u003e at 0.1 \u0026micro;g/\u0026micro;L but repels it at 10 \u0026micro;g/\u0026micro;L, while (E)-2-hexenal exhibits the opposite trend. Similarly, nonanal shows no significant effect at 0.1 or 10 \u0026micro;g/\u0026micro;L but acts as a repellent at 1 \u0026micro;g/\u0026micro;L, further underscoring the nonlinear relationship between concentration and behavioral response (Wang et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The same volatile compounds (e.g., aromatic and terpenoid compounds) from companion plants (\u003cem\u003eDesmodium\u003c/em\u003e spp. and \u003cem\u003eBrachiaria\u003c/em\u003e) repel fall armyworm (\u003cem\u003eSpodoptera frugiperda\u003c/em\u003e) while attracting its parasitoid wasps, demonstrating the dual ecological function of these volatiles in the \"Push-Pull\" cropping system (Sobhy et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Together, these results emphasize that the ecological role of a volatile compound cannot be generalized across species or contexts, and future research should explore how dose-dependent effects and volatile blends shape multitrophic interactions in agroecosystems.\u003c/p\u003e\u003cp\u003eWhile this study provides valuable insights into \u003cem\u003eT. drosophilae\u003c/em\u003e\u0026rsquo;s behavioral responses, certain limitations should be noted. The laboratory conditions using single compounds may not fully represent the complexity of field environments, where volatile interactions and ecological factors likely play important roles. The ecological context of volatile emission must be considered. In natural environments, 3-methylbutanoic acid is part of a complex blend\u0026mdash;such as in \u003cem\u003eD. suzukii\u003c/em\u003e-infested cherry fruit volatiles, where it attracts parasitoid females, whereas its isolated presentation in Y-tube assays elicits repellency. Additionally, the molecular mechanisms underlying these olfactory responses remain to be explored. Future studies should further investigate these aspects under more realistic conditions, examine potential variations across different host plants and environmental contexts, and evaluate the practical effectiveness of these findings for field applications in \u003cem\u003eD. suzukii\u003c/em\u003e management. Such work would help bridge the gap between laboratory research and real-world implementation.\u003c/p\u003e\u003cp\u003eOur study demonstrated that \u003cem\u003eT. drosophilae\u003c/em\u003e exhibited high specificity toward VOCs (especially esters) released by cherry infested with \u003cem\u003eD. suzukii\u003c/em\u003e, highlighting the potential of these volatile cues in developing targeted biological control strategies. These findings provide a theoretical foundation for utilizing VOC-based attractants to enhance parasitoid efficiency in \u003cem\u003eD. suzukii\u003c/em\u003e management. However, further research integrating chemical ecology and field trials is essential to optimize lure formulations (Muskat and Patel 2021; Nieri et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) and elucidate the underlying mechanisms governing parasitoid-host interactions. Such efforts will bridge the gap between laboratory findings and practical applications, ultimately contributing to sustainable pest control in agroecosystems.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAuthor contributions\u0026nbsp;\u003c/strong\u003eS.N.W, Y.Y.W conceived and designed research. H.K.S conducted experiments and wrote the manuscript. F.Y analyzed data, prepared figures and wrote the manuscript. R.L, Z.H.W and G.H.Q revised the manuscript. Z.H.W and Q.F.L assisted in experiments. All authors read and approved the manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgements\u0026nbsp;\u003c/strong\u003eWe express our sincere gratitude to Ting Geng from the Institute of Plant Protection, Chinese Academy of Agricultural Sciences (IPP-CAAS) for his expert assistance in capturing the diagnostic images of \u003cem\u003eTrichopria drosophilae\u003c/em\u003e. Special thanks are extended to Yu Gao for her skillful contribution to the graphical illustrations. We also acknowledge the sustained support provided by the Institute of Forestry and Pomology, Beijing Academy of Agriculture and Forestry Sciences (BAAFS) throughout this research project.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u0026nbsp;\u003c/strong\u003eThis work was supported by the Promotion and Innovation of Beijing Academy of Agriculture and Forestry Sciences (KJCX20250901, KJCX20240403) and the Young Elite Scientists Sponsorship Program by the BAST (BYESS2023474).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData availability\u0026nbsp;\u003c/strong\u003eThe datasets generated during or analyzed during the current study are available from the corresponding author upon reasonable request.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflict of interest\u0026nbsp;\u003c/strong\u003eThe authors have no relevant financial or non-financial interests to disclose.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthical approval\u0026nbsp;\u003c/strong\u003eThis statement is not applicable.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAli MY, Naseem T, Holopainen JK et al (2023) Tritrophic interactions among arthropod natural enemies, herbivores and plants considering volatile blends at different scale levels. 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Bioscience 58:308\u0026ndash;316. https://doi.org/10.1641/B580406\u003c/li\u003e\n\u003cli\u003eSobhy IS, Tamiru A, Chiriboga Morales X et al (2022) Bioactive volatiles from push-pull companion crops repel fall armyworm and attract its parasitoids. Front Ecol Evol 10:. https://doi.org/10.3389/fevo.2022.883020\u003c/li\u003e\n\u003cli\u003eTri\u0026ntilde;anes F, Gonz\u0026aacute;lez A, de la Vega GJ (2022) Olfactory responses of \u003cem\u003eDrosophila suzukii\u003c/em\u003e parasitoids to chemical cues from SWD-infested fruit. bioRxiv. https://doi.org/10.1101/2022.09.08.507209\u003c/li\u003e\n\u003cli\u003eVet LEM, Dicke M (1992) Ecology of infochemical use by natural enemies in a tritrophic context. Annu Rev Entomol 37:141\u0026ndash;172. https://doi.org/10.1146/annurev.en.37.010192.001041\u003c/li\u003e\n\u003cli\u003eWang JL, Yi T, Wang MW et al (2025) Herbivore-induced maize volatiles: dual functions in repelling fall armyworm and attracting natural enemies. 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Insects 15:984. https://doi.org/10.3390/insects15120984\u003c/li\u003e\n\u003cli\u003eYasa V, Suroshe SS, Nebapure SM (2024) Behavioral response of zigzag ladybird beetle \u003cem\u003eCheilomenes sexmaculata\u003c/em\u003e to the HIPVs induced by cotton aphid, \u003cem\u003eAphis gossypii\u003c/em\u003e. Arthropod-Plant Interact 18:771\u0026ndash;780. https://doi.org/10.1007/s11829-024-10087-0\u003c/li\u003e\n\u003cli\u003eYi CD, Cai PM, Lin J, et al (2020) Life history and host preference of \u003cem\u003eTrichopria drosophilae\u003c/em\u003e from Southern China, one of the effective pupal parasitoids on the \u003cem\u003eDrosophila\u003c/em\u003e species. Insects 11:103. https://doi.org/10.3390/insects11020103\u003c/li\u003e\n\u003cli\u003eYu S, Gong L, Han YC, et al (2024) Oral secretions from striped stem borer (\u003cem\u003eChilo suppressalis\u003c/em\u003e) induce defenses in rice. Pest Manag Sci 80:6437\u0026ndash;6449. https://doi.org/10.1002/ps.8376\u003cstrong\u003e\u003c/strong\u003e\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"journal-of-pest-science","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"pest","sideBox":"Learn more about [Journal of Pest Science](https://www.springer.com/journal/10340)","snPcode":"10340","submissionUrl":"https://submission.nature.com/new-submission/10340/3","title":"Journal of Pest Science","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Drosophila suzukii, Trichopria drosophilae, Volatile organic compounds (VOCs), Tritrophic interactions, Biocontrol","lastPublishedDoi":"10.21203/rs.3.rs-6960668/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6960668/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe spotted-wing drosophila (\u003cem\u003eDrosophila suzukii\u003c/em\u003e), a globally invasive pest, induces distinct shifts in the volatile organic compound (VOC) profiles of infested cherry fruits, enhancing their attractiveness to the pupal parasitoid \u003cem\u003eTrichopria drosophilae\u003c/em\u003e. Comparative VOC analysis revealed significant differences among \u003cem\u003eD. suzukii\u003c/em\u003e-infested, mechanically damaged, and healthy fruits. Infested cherries emitted elevated levels of key attractants, including esters (ethyl acetate: 0.36 ng/\u0026micro;L; isoamyl acetate: 0.48 ng/\u0026micro;L) and aromatic aldehydes (4-ethylbenzaldehyde: 0.86 ng/\u0026micro;L), which were absent or minimal in controls. Non-metric multidimensional scaling (NMDS) confirmed distinct VOC clustering, with infested fruits chemically diverging from healthy or mechanically damaged samples. Electrophysiological (EAG) assays identified ethyl acetate as the most potent stimulant, while behavioral assays showed concentration-dependent responses: 10 \u0026micro;g/\u0026micro;L isoamyl acetate elicited strong attraction (72% response rate), whereas 3-methylbutanoic acid acted as a repellent (34% response rate). Cage experiments demonstrated that benzyl alcohol, ethyl acetate, and isoamyl acetate significantly increased parasitism rates in \u003cem\u003eD. suzukii\u003c/em\u003e pupae compared to controls, with isoamyl acetate showing the strongest effect. These results reveal specific semiochemicals mediating tritrophic interactions and underscore their potential for optimizing \u003cem\u003eT. drosophilae\u003c/em\u003e-based biocontrol strategies against \u003cem\u003eD. suzukii\u003c/em\u003e.\u003c/p\u003e","manuscriptTitle":"Host-associated volatile cues drive foraging behavior of Trichopria drosophilae toward Drosophila suzukii- infested fruits","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-07-12 03:32:30","doi":"10.21203/rs.3.rs-6960668/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-08-24T13:04:18+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-07-29T07:29:39+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-07-22T18:42:01+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"146263120035927624344305317919760855508","date":"2025-07-14T07:29:07+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"182710925268273228597065886750849588224","date":"2025-07-11T13:41:36+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"24640515597791715604083958622822426448","date":"2025-07-09T16:17:32+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-07-09T08:48:49+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-06-26T14:06:20+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-06-26T14:03:45+00:00","index":"","fulltext":""},{"type":"submitted","content":"Journal of Pest Science","date":"2025-06-24T02:14:08+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"journal-of-pest-science","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"pest","sideBox":"Learn more about [Journal of Pest Science](https://www.springer.com/journal/10340)","snPcode":"10340","submissionUrl":"https://submission.nature.com/new-submission/10340/3","title":"Journal of Pest Science","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"1e2ea99a-ba33-476a-870f-e13970c32146","owner":[],"postedDate":"July 12th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2025-12-08T16:10:23+00:00","versionOfRecord":{"articleIdentity":"rs-6960668","link":"https://doi.org/10.1007/s10340-025-01987-y","journal":{"identity":"journal-of-pest-science","isVorOnly":false,"title":"Journal of Pest Science"},"publishedOn":"2025-12-05 15:58:21","publishedOnDateReadable":"December 5th, 2025"},"versionCreatedAt":"2025-07-12 03:32:30","video":"","vorDoi":"10.1007/s10340-025-01987-y","vorDoiUrl":"https://doi.org/10.1007/s10340-025-01987-y","workflowStages":[]},"version":"v1","identity":"rs-6960668","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6960668","identity":"rs-6960668","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}