Field application of tea volatiles mediating host selection of Aleurocanthus spiniferus

Field application of tea volatiles mediating host selection of Aleurocanthus spiniferus | 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 Field application of tea volatiles mediating host selection of Aleurocanthus spiniferus Zhi-Fei Jia, Li-Yun Huang, Ya-Nan Bian, Kai Song, Dan-Dan Li, and 3 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4333561/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Aleurocanthus spiniferus is a major pest of tea trees; both nymphs and adults suck plant sap and induce tea sooty mould. Pest occurrence varies significantly among different tea cultivars, even within the same plantation environment. It was speculated that adult A. spiniferus may be differentially attracted by different cultivars. This study aims to identify the bioactive components in tea volatiles that mediate host selection of A. spiniferus and to utilize them for effective field trapping. Through field investigations and Y-tube olfactometer tests, we identifiedtwo highly preferred tea cultivars (‘Huangjinya’ and ‘Fuding white tea’) and two cultivars (‘Baiye No. 1’ and ‘Longjing 43’) that were not. Behavioral assays and gas chromatography-mass spectrometry (GC-MS) analysis identified four attractive compounds [hexanol, ( E )-2-hexenal, linalool, and ( E , E )- α -farnesene] and two repellent compounds [nonanal and ( Z )-3-hexenol] from tea volatiles of the four. Among these compounds, four [nonanol, ( Z )-3-hexenol, linalool and ( E , E )- α -farnesene] were able to elicit obvious electrophysiological (EAG) response at 100 μ g/ μ L. In field trials, the synergistic bait trap equipped with two types of attractants, blend 4 [the mixture of linalool and ( E , E )- α -farnesene] at 100 μ g/ μ L and blend 6 [the mixture of linalool and ( Z )-3-hexenol] at 100 μ g/ μ L, had a significant attractant effect with a long effective period. Two synthetic mixtures of tea volatiles, blend 4 and blend 6, had the potential to be developed as commercial plant-based attractants for adult A. spiniferus . This study contributes to the development of sustainable, environmentally friendly management strategies for a pest that is difficult to prevent and control. Aleurocanthus spiniferus tea cultivars volatiles behavioral assays EAG field trials Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Figure 9 Figure 10 Figure 11 Key Message 1. A. spiniferus occurrence varies significantly among different tea cultivars within the same plantation. 2. It was speculated that adult A. spiniferus may be differentially attracted by different cultivars. 3. The odor components in tea volatiles play a crucial role in the host selection of adult A. spiniferus. 4. Two attractants based on tea volatiles were found to be highly effective in trapping adult A. spiniferus. 1. INTRODUCTION Camellia sinensis (L.) Kuntze (Theaceae), also known as tea, is an important economic crop, widely cultivated in tropical and subtropical regions such as China, Japan, and Kenya (Zhao et al. 2020 ). Aleurocanthus spiniferus (Quaintance) (Hemiptera: Aleyrodidae) is a major pest of tea plants (Tang et al. 2015 ). Adults generally congregate to feed on tea shoots and lay eggs. In addition, first-instar nymphs can be active, while all other nymphal stages are stationary and feed on leaves, directly affecting tea yield and quality. Tea sooty mould, Neocapnodium theae , is caused by the excrement of this pest, which reduces the photosynthetic capability of tea leaves and affects tea plant growth (Byrne and Bellows 1991 ). Aleurocanthus spiniferus is widely distributed in China, Japan, and Indonesia (Uesugi et al. 2016 ; Bragard et al. 2018 ; Yang et al. 2022 ), which has become a common and major pest in Chinese tea plantations. Currently, chemical pesticides are often used to control A. spiniferus (Hazarika et al. 2001 ; Tian et al. 2020 ). Therefore, safer and more effective prevention and control methods are urgently needed. Using insecticides or repellent components from plant volatiles can effectively prevent pests from selecting their hosts. Host plant location and selection is a crucial task for herbivorous insects in their search for resources, mates, and oviposition sites; and olfaction typically plays an important role in this process (Song et al. 2022 ). Adults detect volatile organic compounds (VOCs) emitted by plants in a specific way, discriminating between different resistant plants or cultivars based on the types and proportions of volatile compounds (Ranger et al. 2005 ; Huang et al. 2022 ). Volatiles from non-host plants or resistant cultivars contain repellent components to prevent herbivorous insect infestation (Yuan et al. 2022 ). For example, Rosmarinus officinalis , a non-host plant with specifically bioactive compounds in its volatiles that repelling Ectropis obliqua adults (Zhang et al. 2015 ); ocimene, a dominant component of resistant wheat varieties repelled Sitodiplosis mosellana females (Huang et al. 2022 ). Similarly, attractive components in host volatile blends are the crucial reason why host plants or susceptible cultivars are more susceptible to pest infestation (Birkett et al. 2004 ; Yang et al. 2016 ; Xiu et al. 2019 ). A key volatile, ( Z )-3-hexenyl-acetate, mediate the host and oviposition preference of Spodoptera frugiperda on maize (Wang et al. 2023 ). Some specific compounds have been successfully used for population monitoring and integrated pest management (Li et al. 2014 ; Chen et al. 2019 ; Wang et al. 2020 ). A binary repellent containing DMDS and 1,8-cineole, for instance, applied by a slow-release method against Empoasca onukii in a tea plantation (Cai et al. 2020 ). In tea plantations, there are observable differences in the occurrence of A. spiniferus on four tea cultivars. ‘Huangjinya’ and ‘Fuding white tea’ are more susceptible and suitable for A. spiniferus (Tian et al. 2018 ; Lu et al. 2023 ; He et al. 2023 ), while ‘Baiye No. 1’ and ‘Longjing 43’ are common resistant cultivars (Liang et al. 2012 ). To elucidate the potential role of volatiles in host selection, this study evaluated the selection preferences of adult A. spiniferus towards four tea cultivars with different resistance properties. Differences in volatile components among the four cultivars with different resistances were analyzed using GC-MS. The odor components in tea volatiles, which play a crucial role in the host selection behavior of adult A. spiniferus , were analyzed by electrophysiological (EAG) and behavioral measurements. Finally, these compounds and their mixtures were used as attractants in different tea plantations in two years to identify the most effective attractants. This study provides theoretical support for the development of monitoring and control technology for A. spiniferus . 2. MATERIALS AND METHODS 2.1 Tea cultivars and insect collection Four tea cultivars, ‘Baiye No. 1’, ‘Longjing 43’, ‘Fuding white tea’, and ‘Huangjinya’, were used for headspace volatile analysis, assessment of behavioral preferences and population dynamics of adult A. spiniferus . Adult A. spiniferus were collected from the tea cultivars ‘Huangjinya’ that were maintained in the tea plantations managed by Shandong Qianrun Ecological Agriculture Development Co., Ltd. in Tai’an, Shandong Province, China (32°08′N, 117°43′E). Before experiments, adults were acclimated for one day on four different tea cultivars in environmental incubators at 26 ± 2°C with 70 ± 5% RH and a 16L:8D photoperiod. Insect preference tests for two cultivars, the test insects were kept on the corresponding cultivars, 50/50 for each. Insects for the other laboratory tests were kept on ‘Huangjinya’. Due to the low sex ratio of males, we are unable to obtain enough males for subsequent experiments. Therefore, females and males were mixed for behavioral preference and EAG tests. 2.2 Population dynamics of A. spiniferus on four tea cultivars The experiment was conducted in the same tea plantations as described above from May to November 2022. As typical of their habit of tending to aggregate towards tender leaves and buds, adult A. spiniferus were mainly active in the upper branches of the tea plants. Therefore, the second and third leaves below the bud were selected for counting the density of adults and eggs in the four tea cultivars. The number of nymphs and pseudopupae on the middle and lower leaves of the tea plant was investigated. Specifically, one hundred leaves were randomly selected for counting the number of A. spiniferus adults, eggs, nymphs, and pseudopupae as a biological replicate, respectively. There were four replicates for each tea cultivar. 2.3 Y-tube test for behavioral responses of adult A. spiniferus to tea shoots and synthetic volatile compounds. The Y-tube olfactometer test has been refined with reference to relevant studies (Han and Chen 2002 ), and the apparatus is shown in Fig. 1 . Bioassays were conducted using a dual-choice glass Y-tube olfactometer, which consisted of a central tube and two lateral arms (15 cm long, 13 mm diameter) with an inside angle of 60°. For each paired bioassay, two different odor sources were placed in two odor bottles, which were connected to an activated carbon-filtered, humidified air source and the two arms of a Y-tube through Teflon® tubing. The airflow rate through each arm was 100 mL/min by the pre-experiment, measured using an LZB-3WB rotor meter. Two-year-old tea plants were used for tea plant testing. Two treatments were set up in the experiment: healthy plants and whitefly-infested plants. Pretreatment for whitefly-infested plants was as follows: eight healthy tea plants were placed in a nylon screen cage (50 cm × 50 cm × 50 cm) with 500 (± 10) A. spiniferus adults per plant for each cultivar. Adults were removed after 4 hours of feeding, and tea plants were used when nymphs had developed to the 2nd-3rd instar. Eight healthy tea plants for each cultivar were placed in a nylon screen cage (50 cm × 50 cm × 50 cm) as a healthy plant treatment. To prepare the test chemicals, six standard synthetic chemicals were dissolved in hexane at three different concentrations (1, 10, and 100 µ g/ µ L). Odorants (10 µ L) were added dropwise to filter paper strips (2 × 2 cm), then allowed to evaporate for 20 s and placed in an odor bottle. Filter paper strips (2 × 2 cm) containing hexane were placed in another odor bottle. Experimental adults were starved for 1 hour before the bioassay. A total of 80 insects were tested per bioassay, and each individual was only used once. In each test, a choice was recorded if the adult moved 5 cm past the Y-junction within ten minutes and remained there for at least one minute. Otherwise, it was recorded as no-choice. At the end of the odor source test, the Y-tube, odor bottles, and other glass components were cleaned with acetone and dried before reuse. The Y-tube olfactometer was placed in the laboratory with diffuse light at a temperature of 26 ± 2°C and a relative humidity of 70% ± 5% at about 09:30 − 15:30 hrs. 2.4 Volatile collection and analysis Headspace volatiles from tea plants were collected using the dynamic headspace absorption technique (Sun et al. 2010 ; Zhang et al. 2013 ). The Y-tube tests showed that the pre-infection of A. spiniferus nymphs did not significantly affect the preference of the subsequent adult A. spiniferus. Six 2-year-old healthy tea plants without nymphs were placed in the glass cylinder chamber. The inlet, outlet, and all interfaces of the glass cylinder chamber were sealed with sealing film (Bemis, Inc., Neenah, Wisconsin, USA). The airflow entered the glass cylinder chamber after passing through the silicone desiccant, activated carbon, and rotameter at 200 mL/min. The gas-carrying volatiles were extracted after collection onto the glass tube containing Super Q absorbent (35 mg; 80/100 mesh; Alltech Associates., Inc., Deerfield, IL, USA). All components of the volatile collection apparatus were connected by Teflon® tubing. After 1 hour of collection, the trapped tea shoot volatiles were extracted from the Super Q tube with chromatographically pure hexane (500 µ L). The eluting solution was stored in a glass vial to which decanoic acid ethyl ester (5 µ L) at a concentration of 10 − 4 g/mL was added as an internal standard. The sample was stored at -20°C for qualitative and quantitative analysis of volatile components. The GC was an Agilent 7890B model, and the MS was the 7000D model (Agilent, Inc., PaloAlto, CA, USA). Samples were analyzed on an HP-5ms (30 m × 0.25 mm i.d., 0.25 µ m film thickness). The solvent delay was 3 minutes. The inlet temperature was 230°C, and the GC/MS interface temperature was 280°C. The analysis program was as follows: the oven temperature was programmed at 40°C for 0 minutes, increased at 3°C/min to 190°C for 2 min, then increased at 10°C/min to 290°C and held for 3 minutes. Helium (99.999% purity) was used as the carrier gas at a flow rate of 1 ml per minute. The mass spectrometer was operated in the electron-impact (EI, at 70 eV) scan mode. The scan rate was twice per second. Qualitative analysis of the volatile components was processed by Agilent MassHunter Qualitative Analysis Navigator software. Compounds were identified by comparing the obtained spectra with the spectra of reference compounds from the National Institute of Standards and Technology (NIST17), as well as retention times in the relevant literature on tea shoot volatiles (Mu et al. 2012 ; Sun et al. 2014 ; Cai et al. 2014 ; Cai et al. 2015 ; Magsi et al. 2021 ). The contents of each volatile compound were calculated by comparing their GC total ion current peak areas with the peak area of the internal standard. 2.5 Chemicals Hexane (analytical pure) served as solvent and negative control was purchased from Tianjin Kaitong Chemical Reagent Co. Ltd. (Tianjin, China). Hexanol (99%, CAS: 111-27‐3), linalool (98%, CAS: 78‐70‐6), ( Z )-3-hexenol (98%, CAS: 928‐96‐1), nonanal (96%, CAS: 124‐19‐6), ( E )-2-hexenal (98%, CAS: 6728‐26‐3) was purchased from Macklin Inc. (Shanghai, China), ( E , E )- α -farnesene (96%, CAS: 21499‐64‐9) was synthesized by the College of Chemistry, Shandong Agricultural University. 2.6 EAG experiments The EAG test was based on the same method used for the determination of antennally-active volatiles for tea geometrids (Yuan et al. 2022 ). The EAG system (Syntech, Ltd., Hilversum, The Netherlands) consisted of an IDAC-4 interface box, a CS-55 air stimulus controller, and a desktop computer. To prepare the test chemicals, six standard chemical compounds were dissolved in hexane at five different concentrations (1, 10, 100, 250, and 500 µ g/ µ L). Exactly 10 µ L of the tested solution was placed onto a white filter paper strip (0.5 × 6 cm), the solvent was allowed to evaporate for 30 seconds, then the strip was carefully transferred to a Pasteur pipette cartridge (14.5 cm long). Hexane (10 µ L) was used as the control using the same method. The adult A. spiniferus was chilled in an ice box and removed immediately after the induction of torpor. An antenna was dissected under the stereoscopic microscope (Nikon, Ltd., Japan), and then the reference and recording electrodes were attached to the base and tip of the antenna using conductive adhesive, respectively. To perform the stimulation, the tip of a Pasteur pipette cartridge was inserted into a small hole in the wall of a steel tube (14 cm long, 8 mm diameter), approximately 3 mm diameter, located approximately 11 cm from the end of the tube. A regulated airflow (10 mL/s) entered the steel tube and was directed toward the antenna for chemical exposure. The antenna was located approximately 1 cm from the end of the steel tube. Each antenna was randomly stimulated for testing with three standard chemicals. The stimulation sequence consisted of a control stimulus, followed by three test chemical stimuli, and then another control stimulus. The duration of each stimulation was set to 0.5 s, with an interval of 60 s between two consecutive stimuli (Delorme and Payne 1990 ). A total of 15 antennas were measured for each concentration level of volatiles. Only one antenna was randomly selected per adult. All EAG tests were conducted at a temperature of 26 ± 2°C and a relative humidity of 70 ± 5% from 09:30 − 15:30 hrs. 2.7 Field trial In 1st October 2022, the field trial was conducted in the tea plantation in Gongjiazhuang Village, Jinan City, Shandong Province, China (36°31′N, 117°41′E) with low-density of A. spiniferus adult. In 9th October 2023, it was conducted at the same tea plantation as described in population dynamics experiment, with high-density of A. spiniferus adult, coincident with the third peak of A. spiniferus adult occurrence in tea plantations. The average temperature and relative humidity during the field trials were 8–15°C and 55–70% RH, respectively, and the weather was sunny or cloudy with a gentle breeze. Attractants for tests were formulated as six single-component and seven different mixed-component blends (Table 1 ) diluted with hexane to three concentrations (1, 10, and 100 µ g/ µ L) and adsorbed onto a rubber dispenser (Keyun, Co. Ltd., Henan, China), each containing 50 mg of the active ingredient (a.i.). The following treatments were included: (i) synergistic bait traps: attractants + yellow sticky traps (25×20 cm, Lvpusen Technology, Ltd., Quanzhou, China); (ii) yellow sticky traps; (iii) single attractants: attractants + white sticky traps (25×20 cm, Lvpusen Technology, Ltd., Quanzhou, China); and white sticky trap was used as a control. All sticky traps were attached to bamboo poles, and placed 5–10 cm from the tea shoots; the spacing between each sticky trap was greater than 20 m to minimize the interference between compounds. The trial was conducted in three plots of approximately 1 ha each, with the above four treatments randomized between plots. The distance between the two plots was at least 15 m. The number of adult A. spiniferus on sticky traps was counted on days 3, 7, 14, and 23 in 2022 and on days 7, 14, 23, and 30 in 2023 after the start of the experiment, and sticky traps were replaced after each survey. Table 1 Composition of seven blends formulated as potential attractants for adult Aleurocanthus spiniferus Code Component Blend 1 Hexanol, nonanal, ( E )-2-hexenal, ( Z )-3-hexenol, linalool, ( E , E )- α -farnesene Blend 2 Hexanol, ( Z )-3-hexenol, linalool, ( E , E )- α -farnesene Blend 3 Nonanal, ( E )-2-hexenal Blend 4 Linalool, ( E , E )- α -farnesene Blend 5 ( Z )-3-hexenol, ( E , E )- α -farnesene Blend 6 ( Z )-3-hexenol, linalool Blend 7 Hexanol, ( Z )-3-hexenol 2.8 Data analysis All data were analyzed in Graphpad Prism 9 software (GraphPad Software, San Diego, CA, USA). Data on population dynamics, volatile quantities, relative EAG response, and field trials were analyzed by one-way analysis of variance (ANOVA). The normality of the residuals was tested for all data using the Shapiro-Wilk test. Means were compared using Tukey’s-b multiple range test or the Games-Howell test, depending on whether the treatment and control variances were equal (Shapiro-Wilks test, P > 0.05) or unequal (Shapiro-Wilks test, P < 0.05). The Y-tube olfactometer behavioral test data were analyzed using a chi-squared test ( P < 0.05) to determine differences between pairs of treatments. Non-choice individuals were recorded but excluded from the statistical analysis. Data on lure longevity time was analyzed by two-tailed student’s t test. 3. RESULTS 3.1 Population dynamics of A. spiniferus on four tea varieties The peak flight season occurred in June. The number of A. spiniferus adults and eggs peaked on 14 June 2022, with significant differences in occurrence among different cultivars (adults: F = 34.71, P < 0.001; eggs: F = 63.34, P < 0.001). The number of adults on ‘Huangjinya’ reached 2597.20 ± 386.99, which was significantly higher than that of ‘Fuding white tea’ (807.60 ± 57.86), ‘Baiye No.1’ (156.20 ± 14.21), and ‘Longjing 43’ (149.60 ± 20.25) (Fig. 2 a). Egg abundance showed the same trend, with 3919.80 ± 293.71 for ‘Huangjinya’, slightly lower for ‘Fuding white tea’ at 2587.60 ± 321.15, and only 504.80 ± 45.04 and 275.00 ± 21.47 for ‘Baiye No.1’ and ‘Longjing 43’ (Fig. 2 c). On 14 June, nymph abundance showed significant differences among the different cultivars ( F = 102.16, P < 0.001), with ‘Huangjinya’ and ‘Fuding white tea’ was 3111.80 ± 225.70 and 1359.80 ± 142.46, respectively; while ‘Baiye No.1’ and ‘Longjing 43’ were only 277.20 ± 29.14 and 189.40 ± 18.29, respectively (Fig. 2 b). Pseudopupae abundance on ‘Huangjinya’ and ‘Fuding white tea’ peaked on 31 May (‘Huangjinya’: 2328.40 ± 168.65; ‘Fuding white tea’: 1439.40 ± 143.56) and 14 June (‘Huangjinya’: 1960.00 ± 272.04; ‘Fuding white tea’: 1697.20 ± 206.26), at which time the number of pseudopupae on ‘Baiye No.1’ (31 May: 250.40 ± 19.15; 14 June: 343.80 ± 57.64) and ‘Longjing 43’ (31 May: 200.20 ± 23.31; 14 June: 266.80 ± 22.92) was relatively low (Fig. 2 d). Based on almost one year of investigation, it was tentatively determined that the selectivity of A. spiniferus on four tea cultivars under the same environment in a tea plantation was as follows: ‘Huangjinya’ > ‘Fuding white tea’ > ‘Baiye No.1’ > ‘Longjing 43’. 3.2 Preference of A. spiniferus among four tea cultivars. To further clarify the preference of adult A. spiniferus among the tested tea cultivars, we conducted Y-tube olfactometer experiments. The results showed that the pre-infection of A. spiniferus nymphs did not significantly affect the preference of the subsequent adult A. spiniferus (Fig. 3 a). The preference of adult A. spiniferus for ‘Huangjinya’ was significantly higher than that for the other three cultivars. When paired with ‘Baiye No.1’ and ‘Longjing 43’, adult A. spiniferus tended to choose ‘Fuding white tea’. There was no significant difference in the preference of adult A. spiniferus between ‘Baiye No.1’ and ‘Longjing 43’ (Fig. 3 b and 3 c). 3.3 Volatile profiles of tested tea cultivars. A total of 14 compounds were identified, 11 of which were common to all four cultivars. The content of hexanol ( F = 10.86, P = 0.003), ( E )-2-hexenal ( F = 4.49, P = 0.04), and 7-methyl-heptadecane ( F = 7.16, P = 0.012) in the volatiles of the shoots from four cultivars showed significant differences. Hexanol and ( E )-2-hexenal were dominant in two resistant cultivars (‘Baiye No. 1’ and ‘Longjing 43’). There was no evidence that 7-methyl-heptadecane is a behaviorally active volatile organic compound in tea plants, thus this compound was not included in this study. Additionally, linalool was only present in the susceptible ‘Fuding white tea’. Nonanol was only absent in the resistant ‘Longjing 43’. ( E , E )- α -farnesene was only present in the susceptible ‘Huangjinya’, while ( Z )-3-hexenol was only absent in the ‘Huangjinya’ (Fig. 4 and Table 2 ). IS, internal standard, decanoic acid ethyl ester Table 2 Identified volatile organic compounds (VOCs) emitted from four tea shoots (‘Huangjinya’, ‘Fuding white tea’, ‘Baiye No. 1’, and ‘Longjing 43’) and mean emission amount a (ng plant − 1 h − 1 ) Volatile compounds Retention time (min) Concentration (mean ± SE) (ng plant − 1 h − 1 ) Baiye No. 1 Fuding white tea Huangjinya Longjing 43 1. Hexanol 3.2 6711.92 ± 561.98 a 3727.79 ± 959.26 ab 1667.67 ± 322.83 b 5633.52 ± 688.38 a 2. ( E )-2-Hexenal 7.1 235.80 ± 49.13 ab 167.61 ± 21.43 ab 90.83 ± 21.33 b 249.50 ± 37.39 a 3. ( Z )-3-Hexenol 7.4 48.28 ± 4.37 a 29.11 ± 4.10 a ND 32.96 ± 4.83 a 4. Benzaldehyde 8.0 567.62 ± 124.79 a 406.04 ± 84.73 a 172.94 ± 88.29 a 235.60 ± 35.35 a 5. 2-Ethyl-1-hexanol 11.8 228.87 ± 64.03 a 250.96 ± 51.94 a 220.90 ± 61.29 a 425.11 ± 67.08 a 6. ( E )-Ocimene 12.9 1045.18 ± 398.30 a 789.52 ± 158.49 a 566.01 ± 154.63 a 539.97 ± 169.47 a 7. Linalool 15.6 ND 96.29 ± 23.63 ND ND 8. Nonanol 15.9 60.37 ± 18.47 a 34.35 ± 4.22 a 14.54 ± 3.72 a ND 9. ( Z )-3-Hexenyl butanoate 19.1 1025.41 ± 248.76 a 858.78 ± 149.65 a 689.06 ± 179.84 a 784.68 ± 237.15 a 10. 7-Methyl-heptadecane 26.9 681.58 ± 202.02 b 636.01 ± 82.64 b 1815.75 ± 137.89 a 1355.68 ± 336.37 ab 11. ( E , E )- α -Farnesene 30.9 ND ND 34.25 ± 2.60 ND 12. Unknown 34.1 732.53 ± 236.59 b 2160.95 ± 319.25 ab 3763.17 ± 585.63 a 3819.90 ± 680.01 a 13. Cadalene 40.5 814.17 ± 256.77 a 963.19 ± 174.40 a 766.72 ± 159.95 a 1713.63 ± 303.49 a 14. Caffeine 46.2 227.83 ± 47.99 a 303.41 ± 41.73 a 252.29 ± 50.87 a 415.13 ± 64.31 a a Data in the table are presented as mean ± SE ( n = 3), and those followed by different letters in the same row indicate significant differences at 0.05 level by Tukey’s-b multiple range test. “ND” denotes that compounds were not detected 3.4 Preference of A. spiniferus for synthetic compounds of tea volatiles. All six compounds were able to elicit an intense response in adult A. spiniferus , and the response was concentration-dependent. Adults were significantly attracted to hexanol at 100 µ g/ µ L; to linalool at 100 µ g/ µ L; to ( E )-2-hexenal at 100 µ g/ µ L, 10 µ g/ µ L; to ( E , E )- α -farnesene at 100 µ g/ µ L, 10 µ g/ µ L, 1 µ g/ µ L. In contrast, they were repelled by both nonanal and ( Z )-3-hexenol at all concentrations tested concentrations ranging from 1 to 100 µ g/ µ L (Fig. 5 ). 3.5 EAG responses elicited by synthetic compounds of tea volatiles. Among six synthetic compounds of tea volatiles tested, four were able to elicit an obvious EAG response at 100 µ g/ µ L, including nonanol (relative EAG value = 0.12 mV) (Fig. 6 b), ( Z )-3-hexenol (relative EAG value = 0.14 mV) (Fig. 6 d), linalool (relative EAG value = 0.13 mV) (Fig. 6 e), and ( E , E )- α -farnesene (relative EAG value = 0.11 mV) (Fig. 6 f). The other two compounds, hexanol and ( E )-2-hexenal, elicited very weak or no response at all tested concentrations (relative EAG value < 0.1 mV) (Fig. 6 a and 6 c). 3.6 Field trials of synthetic compounds and blends of tea volatiles. In October 2022 and 2023, the field trials were conducted in two different tea gardens in Jinan and Tai’an, respectively. The result showed that the synergistic bait trap (attractant + yellow sticky trap) had the best attraction effect. The trapping effect of synergistic bait traps containing blend 3 at 100 µ g/ µ L, blend 4 at 100 µ g/ µ L, and blend 6 at 100 µ g/ µ L were significantly better than those of the other three treatments (CK, single attractant, and yellow sticky trap) in 2022 (Figs. 7 and 9 ). The trapping effect of synergistic bait traps containing ( E )-2-hexenal at 1 µ g/ µ L, ( E , E )- α -farnesene at 1 µ g/ µ L, blend 1 at 1 µ g/ µ L, blend 3 at 10 µ g/ µ L, blend 4 at 100 µ g/ µ L, blend 5 at 100 µ g/ µ L, and blend 6 at 100 µ g/ µ L were significantly better than those of the other three treatments in 2023 (Figs. 8 and 10 ). The lure longevity time of two attractants, blend 4 [the mixture of linalool and ( E , E )- α -farnesene] and blend 6 [the mixture of ( Z )-3-hexenol and linalool] at a concentration of 100 µ g/ µ L, were assessed in 2022 and 2023. From 3 to 14 days after application in 2022 and from 7 to 23 days after application in 2023, compared to using yellow sticky traps alone, the number of adult A. spiniferus was significantly higher in yellow sticky traps containing two types of attractants, blend 4 and blend 6 (Fig. 11 ). From 14 to 23 days after application in 2022 and from 23 to 30 days after application in 2023, there was no significant difference in the number of adults captured by each of the two treatments compared to the yellow sticky trap alone (Fig. 11 ). 4. DISCUSSION It is well known that the type or variety of host plants affects the growth, development, and reproduction of herbivorous insects (Golizadeh et al. 2016 ; Sarkar et al. 2021 ). Therefore, the selection of optimal host plants is an important factor in the life of herbivorous insects. In this study, A. spiniferus adults were more inclined to choose ‘Huangjinya’ and ‘Fuding white tea’ compared with ‘Baiye No.1’ and ‘Longjing 43’, and laid more eggs, in the same environment of a tea plantation (Fig. 2 ). ‘Huangjinya’ is a novel chlorophyll-deficient albino cultivar that is susceptible to common pests of tea plants (e.g., E. onukii and Toxoptera aurantii ) (Tian et al. 2018 ; He et al. 2023 ). ‘Fuding white tea’ is more suitable for herbivorous insects (e.g., Dendrothrips minowai , T. aurantii , and E. onukii ) due to its high free amino acid content (Lu et al. 2023 ). Although the nutritional level of plants often affects their suitability and resistance to herbivorous insects, insects search for and locate distant host plants using their sense of smell and vision in the early stages of pest infestation (Bruce et al. 2005 , 2011). External morphological features of tea plants may be an important factor in the attraction of adult A. spiniferus , such as the unique yellowish-green leaves of ‘Huangjinya’, which may be more attractive to insects with a preference for yellow (Li et al. 2018 ). However, the results of indoor Y-shaped olfactometer experiments showed that adult A. spiniferus were still strongly attracted by ‘Huangjinya’ and ‘Fuding white tea’ even without visual cues. Therefore, olfaction plays a crucial role in the host-finding process of adult A. spiniferus . When choosing between healthy plants and whitefly-infested plants of ‘Longjing 43’, 62.5 percent of adults did not make a choice (Fig. 3 a). Odor may be an major factor contributing to the low population density on ‘Longjing 43’. Plants release varying amounts of volatile compounds from their leaves to regulate the behavior of various community members around them (Baldwin et al. 2006 ; Soler et al. 2007 ). The comparison of the headspace volatiles of tea shoots from four cultivars showed that ‘Longjing 43’ and ‘Baiye No. 1’ released more attractive compounds, hexanol and ( E )-2-hexenal, compared with the other two cultivars. This is consistent with previously reported research; hexanol is a positive signal for attracting Nysomyia neivai (Machado et al. 2002); ( E )-2-hexenal has exhibited an attractive effect on herbivorous insects (e.g., Plutella xylostella and Bemisia tabaci ) (Li et al. 2014 ; Yang et al. 2016 ; Onnink et al. 2017 ), which can be used to lure E. onukii into pheromone-baited traps (Mu et al. 2012 ). It is worth noting that hexanol and ( E )-2-hexenal did not elicit an obvious EAG response in adult A. spiniferus (Fig. 6 a and 6 c). Therefore, the antennae may not serve as the primary sensory organ for their detection. Chemoreceptors are present in body hairs, appendages, mouthparts, and wings of insects (Sánchez-Gracia et al. 2009 ). Odorant-binding proteins (OBPs) and chemosensory proteins (CSPs), which are involved in the binding and transport of odor molecules, are also expressed in tissues outside the antennae (Yi et al. 2015 ; Lizana et al. 2022 ). Tissue-specific transcriptome analysis revealed that CSPs are widely expressed in both the head (including the antennae) and body tissues of adult orange spiny whiteflies (Gao et al. 2022 ). Linalool have been only detected in ‘Fuding white tea’, and laboratory tests have confirmed that at a concentration of 100 µ g/ µ L, it can elicit obvious EAG responses and show attractive effects (Figs. 5 and 6 e). Natural linalool usually has two isomers: the levorotatory isomer (S)-(+)-linalool and the dextrorotatory isomer (R)-(−)-linalool (Aprotosoaie et al. 2014 ). (S)-(+)-linalool is mainly attracting pollinators, whereas (R)-(−)-linalool is mainly enhances resistance to herbivores and pathogens (Zhang et al. 2023 ). Unfortunately, we have not been able to identify which isomer of linalool was released from ‘Fuding white tea’ by GC-MS. Further work is needed to demonstrate the conformation as well as the specific roles of the linalool released from the non-flowering ‘Fuding white tea’. In this study, only trace amounts [34.25 ± 2.60 (ng plant − 1 h − 1 )] of ( E , E )- α -farnesene were detected in ‘Huangjinya’, and it showed significant attractive effects at the concentrations of 1, 10, and 100 µ g/ µ L. ( E , E )- α -farnesene is a common compound of herbivore-induced plant volatiles from many plants such as tea, soybean, pear, apple, and poplar, and it is a positive signal for attracting natural enemies (e.g., Aphidius ervi , Coleomegilla maculate , and Chrysoperla carnea ) (Du et al. 1998 ; Zhu et al. 1999 ; Lin et al. 2017 ; Wang et al. 2019 ). Consequently, we reasoned that the presence of ( E , E )- α -farnesene may have both positive and negative effects, as it attracts natural enemies while making the tea plant vulnerable to attacks from A. spiniferus . Except for the susceptible ‘Huangjinya’, ( Z )-3-hexenol was detected in the headspace volatiles of the other three cultivars (Table 2 ). Laboratory tests confirmed that ( Z )-3-hexenol had a repellent effect at concentrations of 1, 10, and 100 µ g/ µ L (Fig. 5 ). The accumulating evidence on the C6-volatile ( Z )‐3‐hexenol supports its crucial role in mediating indirect defense responses in plants. Tea plants release ( Z )-3-hexenol as a defense response against herbivores (Dong et al. 2011 ; Sun et al. 2014 ; Cai et al. 2014 ). This compound contributes to the unique grassy aroma of green tea (Jing et al. 2019 ). We hypothesized that healthy tea plants emit trace amounts of ( Z )-3-hexenol as an insect repellent; conversely, undamaged ‘Huangjinya’ do not release the aforementioned repellent and are therefore favored by adult A. spiniferus . Another compound with a repellent effect verified by behavioral experiments is nonanal, but it was not detected in the repellent variety ‘Longjing 43’. This could be attributed to the fact that most insects use mixtures of volatiles instead of single components for host identification (Delorme and Payne 1990 ; Bruce et al. 2005 ). For example, female Aulacphora foveicollis responded positively to the volatile mixture of Solena amplexicaulis fruits due to the synergistic effect of nonanal and ( E , Z )-2,6-nonadienals (Karmakar et al. 2020 ). The application of plant-derived semiochemicals to control tea pests has been widely reported, such as the application of a mixture of dimethyl disulfide (DMDS) and 1,8-eucalyptin to repel E. onukii in tea fields and the application of ( E )-2-hexenal, ( Z )-3-hexenol, ( Z )-hexenyl acetate, ( E )-ocimene, linalool, and geraniol as E. onukii attractants (Chen et al. 2019 ; Cai et al. 2020 ). Field trials had shown a significant improvement in trapping efficiency when combined with yellow sticky traps. Yellow sticky traps are commonly used for whitefly monitoring and control (Moreau and Isman 2011 ). Extensive control of adult A. spiniferus using yellow sticky traps has been reported in Japan (Uesugi and Sato 2013 ). The results of field trials showed that, the addition of attractants greatly increased the attractiveness of the yellow sticky traps. It is worth noting that more attractants had a significant luring effect in the tea plantation with high-density of adult A. spiniferus . Previous studies have shown that when populations were low, enlarging the entry area can increased Drosophila suzukii captures and improved selectivity for D. suzukii (Renkema et al. 2014 ). When areas affected by extremely dense bark beetle populations, the use of anti-attractants did not prove effective (Jakuš et al. 2022 ). Suggesting that population density must be taken into account in the testing of attractant effects. In the lure longevity time experiment in October 2023, the number of adult A. spiniferus trapped by yellow sticky traps gradually declined, it is consistent with the results of population dynamics (Fig. 2 a). The field trial was conducted when the third peak of A. spiniferus adult occurrence, subsequently, the number of adult A. spiniferus gradually decreased. This is an indication that the lure longevity time may be affected by the season; the long-term monitoring tests will be required (Wang et al. 2024 ). The field trials revealed two anomalies: (i) ( Z )-3-hexenol and nonanal, which were repellent in the laboratory test, did not show repellent effects in the field experiments, but instead showed a positive response; (ii) in Tai’an (2023) but not Jinan (2022), the lowest dose of ( E )-2-hexenal, ( E , E )- α -farnesene and blend 1 increased the catches significantly, it was not consistent with laboratory tests. Background odors in the field can interfere with the attractiveness of plant volatile attractants to herbivorous insects (Xu et al. 2017 ). Similarities were found in a screening study for D. suzukii attractant volatiles, when the co-attractiveness of some of the individual compounds were tested under different background odor environments, they appears to change, suggesting that background odor can influence detection of potential attractants (Cha et al. 2018 ). In the other study, five traps were put at different sites in a cornfield that had record of emergence of Anomala corpulenta , only the baits at site 1–4 effectively lured and trapped (Qu et al. 2024 ). Suggesting that for more accurate conclusions, field trials need to be carried out at different locations. 5. CONCLUSION Our study demonstrates that plant volatiles mediate host selection of adult A. spiniferus among tea cultivars. Blend 4 [the mixture of linalool and ( E , E )- α -farnesene] and blend 6 [the mixture of linalool and ( Z )-3-hexenol], two combinations of attractants based on tea shoot volatiles, were found to be highly effective and long-lasting in trapping adult A. spiniferus in two years. They have the potential to be developed as commercial attractants for the prevention and monitoring of adult A. spiniferus . Declarations Author’s contribution YYX, ZZC and ZFJ conceived the idea and designed the research; ZFJ, LYH, YNB and KS conducted the experiments and analyzed the data; ZFJ wrote the manuscript; ZQL, YYX, ZZC and DDL edited and reviewed the manuscript. All authors commented and approved the manuscript. Acknowledgments We sincerely thank all the staff and students of the Laboratory of Insect Ecology and Physiology (Shandong Agricultural University), as well as Shandong Qianrun Ecological Agriculture Development Co., Ltd., for providing us with the experimental fields. We would like to thank DBMediting for professional english language editing services. Funding This research was funded by the National Key Research & Development Program of China (2023YFD1700405) and the Modern Tea Industry Technology System of Shandong Province (SDAIT-19-04). Competing interests The authors acknowledge no conflict of interest concerning the publication of this research article. Data availability The datasets generated and analysed during the current study are available from the corresponding author on reasonable request. References Aprotosoaie AC, Hancianu M, Costache II, Miron A (2014) Linalool: A review on a key odorant molecule with valuable biological properties. Flavour Frag J 29:193−219. https://doi.org/10.1002/ffj.3197 Baldwin IT, Halitschke R, Paschold A, von Dahl CC, Preston CA (2006) Volatile signaling in plant−plant interactions:"Talking trees" in the genomics era. Science 311:812−815. https://science.org/doi/10.1126/science.1118446 Birkett MA, Bruce TJA, Martin JL, Smart LE, Oakley J, Wadhams LJ (2004) Responses of female orange wheat blossom midge, Sitodiplosis mosellana , to wheat panicle volatiles. J Chem Ecol 30:1319−1328. https://doi.org/10.1023/b:joec.0000037742.05022.9f Bragard C, Dehnen‐Schmutz K, Di Serio F et al (2018). Pest categorisation of Aleurocanthus spp. . EFSA J 16:5436. https://doi.org/10.2903/j.efsa.2018.5436 Bruce TJA, Pickett JA (2011) Perception of plant volatile blends by herbivoro-us insects－Finding the right mix. Phytochemistry 72:1605−1611. https://doi.org/10.1016/j.phytochem.2011.04.011 Bruce TJA, Wadhams LJ, Woodcock CM (2005) Insect host location: Avolatile situation. Trends Plant Sci 10:269−274. https://doi.org/10.1016/j.tplants.2005.04.003 Byrne DN and Bellows TS (1991) Whitefly biology. Annu Rev Entomol 36:431–457. https://doi.org/10.1146/annurev.en.36.010191.002243 Cai XM, Sun XL, Dong WX, Wang GC, Chen ZM (2014) Herbivore species, infestation time, and herbivore density affect induced volatiles in tea plants. Chemoecology 24:1−14. https://doi.org/10.1007/s00049-013-0141-2 Cai XM, Xu XX, Bian L, Luo ZX, Xin ZJ, Chen ZM (2015) Attractiveness of host volatiles combined with background visual cues to the tea leafhopper, Empoasca vitis . Entomol Exp Appl 157:291−299. https://doi.org/10.1111/eea.12364 Cai XM, Luo ZX, Meng ZN et al (2020) Primary screening and application of repellent plant volatiles to control tea leafhopper, Empoasca onukii Matsuda. Pest Manag Sci 76:1304−1312. https://doi.org/10.1002/ps.5641 Cha DH, Loeb GM, Linn CEJ, Hesler SP, Landolt PJ (2018) A multiple-choice bioassay approach for rapid screening of key attractant volatiles. Environ Entomol 47:946−950. https://doi.org/10.1093/ee/nvy054 Chen K, Huang MX, Shi QC, Xie X, Jin LH, Xu WM, Li XY (2019) Screening of a potential leafhopper attractants and their applications in tea plantations. J Environ Sci Heal B 54:858−865.https://doi.org/10.1080/03601234.2019.1633856 Delorme JD, Payne TL (1990) Antennal olfactory responses of black turpentine beetle, Dendroctonus terebrans (Olivier), to bark beetle pheromones and host terpenes. J Chem Ecol 16:1321−1329. https://doi.org/10.1007/bf01021029 Dong F, Yang Z, Baldermann S, Sato Y, Asai T, Watanabe N (2011) Herbivore-induced volatiles from tea ( Camellia sinensis ) plants and their involveme-nt in intraplant communication and changes in endogenous nonvolatile me-tabolites. J Agric Food Chem 59:13131−13135. https://doi.org/10.1021/jf203396a Du YJ, Poppy GM, Powell W, Pickett JA, Wadhams LJ, Woodcock CM (1998)Identification of semiochemicals released during aphid feeding that attract parasitoid Aphidius ervi . J Chem Ecol 24:1355−1368. https://doi.org/10.1023/a:1021278816970 Gao YQ, Chen ZZ, Liu MY et al (2022) Characterization of antennal chemos-ensilla and associated chemosensory genes in the orange spiny whitefly, A-leurocanthus spiniferus (Quaintanca). Front Physiol 13:847895. https://doi.org/10.3389/fphys.2022.847895 Golizadeh A, Abedi Z, Borzoui E, Golikhajeh N, Jafary M (2016) Susceptibilit-y of five sugar beet cultivars to the black bean aphid, Aphis fabae scopoli(Hemiptera: Aphididae). Neotrop Entomol 45:427–432. https://doi.org/10.1007/s13744-016-0383-0 Han BY and Chen ZM (2002) Behavioral and electrophysiological responses ofnatural enemies to synomones from tea shoots and kairomones from tea a-phids, Toxoptera aurantii . J Chem Ecol 28:2203−2219. https://doi.org/10.1023/a:1021045231501 Hazarika LK, Puzari KC, Wahab S (2001) Biological control of tea pests. In: Upadhyay RK, Mukerji KG and Chamola BP (ed) Biocontrol potential and its exploitation in sustainable agriculture. Springer, Boston, pp 159–180. He Y, Lu C, Jiang W, Chen WL, Fan JJ, Niu SZ, Zhao DG (2023) Adaptability of Toxoptera aurantii (Hemiptera: Aphididae) to different tea cultivars. Agriculture 13:2039. https://doi.org/10.3390/agriculture13102039 Huang QT, Han XQ, Zhang GJ, Zhu-Salzman K, Cheng WN (2022) Plant vol-atiles mediate host selection of Sitodiplosis mosellana (Diptera: Cecidomyi-idae) among wheat varieties. J Agric Food Chem 70:10466−10475. https://doi.org/10.1021/acs.jafc.2c03244 Jakuš R, Modlinger R, Kašpar J et al (2022) Testing the efficiency of the pus-h-and-pull strategy during severe Ips typographus outbreak and extreme dr-ought in Norway spruce stands. Forests 13:2175. https://doi.org/10.3390/f13122175 Jing TT, Zhang N., Gao T et al (2019) Glucosylation of (Z)‐3‐hexenol informs intraspecies interactions in plants: A case study in Camellia sinensis . Plant, Cell & Environment 42:1352–1367. https://doi.org/10.1111/pce.13479 Karmakar A, Mitra P, Koner A, Das S, Barik A (2020) Fruit volatiles of creeping cucumber ( Solena amplexicaulis ) attract a generalist insect herbivore. J Chem Ecol 46:275−287. https://doi.org/10.1007/s10886-020-01154-w Li CF, Ma JQ, Huang DJ, Ma CL, Jin JQ, Yao MZ, Chen L (2018) Comprehensive dissection of metabolic changes in albino and green tea cultivars. J Agric Food Chem 66:2040−2048. https://doi.org/10.1021/acs.jafc.7b05623 Li YF, Zhong ST, Qin YC, Zhang SQ, Gao ZL, Dang ZH, Pan WL (2014) Identification of plant chemicals attracting and repelling whiteflies. Arthropod-Plant Inte 8:183−190. https://doi.org/10.1007/s11829-014-9302-7 Liang LY, Liu LF, Yu XP, Han BY (2012) Evaluation of the resistance of different tea cultivars to tea aphids by EPG technique. J Integr Agr 11:2028−2034. https://doi.org/10.1016/S2095-3119(12)60460-2 Lin J, Wang D, Chen X et al (2017) An (E,E)-α-farnesene synthase gene of s-oybean has a role in defence against nematodes and is involved in synthe-sizing insect-induced volatiles. Plant Biotechnol J 15:510−519. https://doi.org/10.1111/pbi.12649 Lizana P, Mutis A, Quiroz A, Venthur H (2022). Insights into chemosensory p-roteins from non-model insects: Advances and perspectives in the context of pest management. Front Physiol 13:924750. https://doi.org/10.3389/fphys.2022.924750 Lu C, Shen N, Jiang W, et al (2023) Different tea germplasms distinctly influence the adaptability of Toxoptera aurantii (Hemiptera: Aphididae). Insects 14:695. https://doi.org/10.3390/insects14080695 Machado VE, Silva FBD, Goulart TM, Pinto MC (2022) Behavioral responses of sand fly Nyssomyia neivai (Psychodidae: Phlebotominae) to 1-hexanol and light. Acta Trop 236:106680. https://doi.org/10.1016/j.actatropica.2022.106680 Magsi FH, Luo ZX, Zhao YJ, Li ZQ, Cai XM, Bian L, Chen ZM (2021) Electrophysiological and behavioral responses of Dasychira baibarana (Lepidoptera: Lymantriidae) to tea plant volatiles. Environ Entomol 50:589−598. https://doi.org/10.1093/ee/nvab016 Moreau TL and Isman MB (2011) Trapping whiteflies? A comparison of greenhouse whitefly ( Trialeurodes vaporariorum ) responses to trap crops and yellow sticky traps. Pest Manag Sci 67:408−413. https://doi.org/10.1002/ps.2078 Mu D, Cui L, Ge J et al (2012) Behavioral responses for evaluating the attractiveness of specific tea shoot volatiles to the tea green leafhopper, Empoaca vitis . Insect Sci 19:229−238. https://doi.org/10.1111/j.1744-7917.2011.01476.x Onnink CM, Williams AA, Williams JS (2017) Electrophysiological and behavioral evidence for (E)-2-hexenal as a female-attracting pheromone produced by disturbed Megacopta cribraria (Hemiptera: Plataspidae). J Entomol Sci 52:39−51. https://doi.org/10.18474/JES16-18.1 Qu CJ, Shi JT, Wang CZ, Guo H, Jiang XJ (2024) Identification of an adult attractant for Anomala corpulenta by the reverse chemical ecology approach. J Pest Sci. https://doi.org/10.1007/s10340-023-01740-3 Ranger CM, Winter REK, Backus EA, Rottinghaus GE, Ellersieck MR, Johnson DW (2005) Discrimination by the potato leafhopper (Hemiptera: Cicadellidae) of host volatiles from resistant and susceptible alfalfa, Medicago sativa L. Environ Entomol 34:271−280. https://doi.org/10.1603/0046-225X-34.2.271 Renkema JM, Buitenhuis R, Hallett RH (2014) Optimizing trap design and trapping protocols for Drosophila suzukii (Diptera: Drosophilidae). J Econ Entomol 107:2107−2118. https://doi.org/10.1603/ec14254 Sánchez-Gracia A, Vieira FG, Rozas J (2009) Molecular evolution of the majorchemosensory gene families in insects. Heredity 103: 208−216. https://doi.org/10.1038/hdy.2009.55 Sarkar S, Babu A, Chakraborty K, Deka B, Kashyap B, Adhikary B (2021) Study on varietal preference and nutritional indices of tea looper Hyposidra talaca (Lepidoptera: Geometridae) on ten selected tea varieties. Int J Trop Insect Sc 41:3087−3098. https://doi.org/10.1007/s42690-021-00502-x Soler R, Harvey JA, Kamp AFD et al (2007) Root herbivores in fluence the behaviour of an aboveground parasitoid through changes in plant-volatile signals. Oikos 116:367−376. https://doi.org/10.1111/j.0030-1299.2007.15501.x Song CF, Ma L, Zhao JY, Xue ZS, Yan XZ, Hao C (2022) Electrophysiological and behavioral responses of Plutella xylostella (Lepidoptera:Plutellidae) to volatiles from a non-host plant, geranium, Pelargonium hortorum (Geraniaceae). J Agric Food Chem 70:5982−5992. https://doi.org/10.1021/acs.jafc.1c08165 Sun XL, Wang GC, Cai XM, Jin S, Gao Y, Chen ZM (2010) The tea weevil, Myllocerinus aurolineatus , is attracted to volatiles induced by conspecifics. J Chem Ecol 36:388−395. https://doi.org/10.1007/s10886-010-9771-9 Sun XL, Wang GC, Gao Y, Zhang XZ, Xin ZJ, Chen ZM (2014) Volatiles emitted from tea plants infested by Ectropis obliqua larvae are attractive to conspecific moths. J Chem Ecol 40:1080−1089. https://doi.org/10.1007/s10886-014-0502-5 Tang XT, Tao HH, Du YZ (2015) Microsatellite-based analysis of the genetic structure and diversity of Aleurocanthus spiniferus (Hemiptera: aleyrodidae)from tea plants in China. Gene 560:107–113. https://doi.org/10.1016/j.gene.2015.01.050 Tian Y, Zhao Y, Zhang L, Mu W, Zhang ZQ (2018) Morphological, physiolog-ical, and biochemical responses of two tea cultivars to Empoasca onukii (Hemiptera: Cicadellidae) infestation. J Econ Entomol 111:899−908. https://doi.org/10.1093/jee/toy011 Tian YY, Chen ZJ, Huang XQ, Zhang LX, Zhang ZQ (2020) Evaluation of botanicals for management of piercing–sucking pests and the effect on beneficial arthropod populations in tea trees Camellia sinensis (L.) O. Kuntze (Theaceae). J Insect Sci 20: 27. https://doi.org/10.1093/jisesa/ieaa101 Uesugi R and Sato Y (2013) Counting adults on a yellow sticky trap is useful for monitoring the tea spiny whitefly, Aleurocanthus camelliae (Hemiptera: Aleyrodidae), at the early invasion stage. JPN J Appl Entomol Z 57:35−41. https://doi.org/10.1303/jjaez.2013.35 Uesugi R, Sato Y, Han BY, Huang ZD, Yara K, Furuhashi K (2016) Molecular evidence for multiple phylogenetic groups within two species of invasive spiny whiteflies and their parasitoid wasp. Bull Entomol Res 106:328–340. https://doi.org/10.1017/S0007485315001030 Wang HM, Bai PH, Zhang J, Zhang XM, Hui Q, Zheng HX, Zhang XH (2020) Attraction of bruchid beetles Callosobruchus chinensis (L.)(Coleoptera: Bruchidae) to host plant volatiles. J Integr Agr 19:3035−3044. https://doi.org/10.1016/S2095-3119(20)63237-3 Wang J, Wei J, Yi T, Li YY, Xu T, Chen L, Xu H (2023) A green leaf volat-ile, (Z)-3-hexenyl-acetate, mediates differential oviposition by Spodoptera f-rugiperda on maize and rice. BMC Biol 21:140. https://doi.org/10.1186/s12915-023-01642-x Wang LY, Yang CX, Mei XD, Guo BB, Yang XL, Zhang T, Ning J (2024) Identification of sex pheromone in Macdunnoughia crassisigna Warren (Lepidoptera: Noctuidae) and field optimization of the sex attractant. Pest Manage Sci 80:577−585. https://doi.org/10.1002/ps.7783 Wang X, Zeng L, Liao Y, Li J, Tang J, Yang Z (2019) Formation of α-farnes-ene in tea ( Camellia sinensis ) leaves induced by herbivore-derived woundi-ng and its effect on neighboring tea plants. Int J Mol Sci 20:4151. https://doi.org/10.3390/ijms20174151 Xiu CL, Xu B, Pan HS, Zhang W, Yang YZ, Lu YH (2019) Volatiles from Sophora japonica flowers attract Harmonia axyridis adults (Coleoptera: Coccinellidae). J Integr Agr 18:873−883. https://doi.org/10.1016/S2095-3119(18)61927-6 Xu XX, Cai XM, Bian L, Luo ZX, Li ZQ, Chen ZM (2017) Does backgroun-d odor in tea gardens mask attractants? Screening and application of attra-ctants for Empoasca onukii Matsuda. J Econ Entomol 110:2357−2363. https://doi.org/10.1093/jee/tox269 Yang G, Zhang YN, Gurr GM, Vasseur L, You M (2016). Electroantennogram and behavioral responses of Cotesia plutellae to plant volatiles. Insect Sci 23:245−252. https://doi.org/10.1111/1744-7917.12308 Yang TB, Song XH, Zhong Y, Wang B, Zhou CQ (2022) Field investigation‐ and dietary metabarcoding‐based screening of arthropods that prey on primary tea pests. Ecol Evol 12:e9060. https://doi.org/10.1002/ece3.9060 Yi X, Zhang Y, Wang P, Qi JW, Hu MY, Zhong GH (2015) Ligands binding and molecular simulation: the potential investigation of a biosensor based on an insect odorant binding protein. Int J Biol Sci 11:75−87. https://www.ijbs.com/v11p0075.htm Yuan GG, Zhao LC, Du YW, Yu H, Shi XB, Chen WC,Chen G (2022) Repel-lence or attraction: secondary metabolites in pepper mediate attraction and defense against Spodoptera litura . Pest Manage Sci 78:4859−4870. https://doi.org/10.1002/ps.7107 Zhang L, Su QF, Wang LS, Lv MW, Hou YX, Li SS (2023) Linalool: A ubiquitous floral volatile mediating the communication between plants and insects. J Syst Evol 61:538−549. https://doi.org/10.1111/jse.12930 Zhang ZQ, Sun XL, Xin ZJ, Luo ZX, Gao Y, Bian L, Chen ZM (2013) Ident-ification and field evaluation of non-host volatiles disturbing host location by the tea geometrid, Ectropis obliqua . J Chem Ecol 39:1284−1296. https://doi.org/10.1007/s10886-013-0344-6 Zhang ZQ, Bian L, Sun XL, Luo ZX, Xin ZJ, Luo FJ, Chen ZM (2015) Ele-ctrophysiological and behavioural responses of the tea geometrid Ectropis obliqua (Lepidoptera: Geometridae) to volatiles from a non-host plant, ros-emary, Rosmarinus officinalis (Lamiaceae). Pest Manag Sci 71:96−104. https://doi.org/10.1002/ps.3771 Zhao MY, Zhang N, Gao T et al (2020) Sesquiterpene glucosylation mediated by glucosyltransferase UGT91Q2 is involved in the modulation of cold str-ess tolerance in tea plants. New Phytol 226:362–372. https://doi.org/10.1111/nph.16364 Zhu JW, Cosse AA, Obrycki JJ, Boo KS, Baker TC (1999) Olfactory reactions of the twelve-spotted lady beetle, Coleomegilla maculata and the green lacewing, Chrysoperla carnea to semiochemicals released from their prey and host plant: electroantennogram and behavioral responses. J Chem Ecol 25:1163−1177. https://doi.org/10.1023/a:1020846212465 Additional Declarations No competing interests reported. Cite Share Download PDF Status: Posted Version 1 posted 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. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. 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-4333561","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":296614423,"identity":"37085032-da8a-4c78-950b-17fabca3b597","order_by":0,"name":"Zhi-Fei Jia","email":"","orcid":"","institution":"Shandong Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"Zhi-Fei","middleName":"","lastName":"Jia","suffix":""},{"id":296614425,"identity":"5968816e-6d2d-42f5-8c24-ce8b94cc1738","order_by":1,"name":"Li-Yun Huang","email":"","orcid":"","institution":"Wuzhou Institute of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Li-Yun","middleName":"","lastName":"Huang","suffix":""},{"id":296614427,"identity":"e4671fa8-26e0-42ae-9b4b-19383ee0fd35","order_by":2,"name":"Ya-Nan Bian","email":"","orcid":"","institution":"Shandong Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"Ya-Nan","middleName":"","lastName":"Bian","suffix":""},{"id":296614428,"identity":"0ca495cc-5c28-490d-a561-2f156e444db3","order_by":3,"name":"Kai Song","email":"","orcid":"","institution":"Shandong Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"Kai","middleName":"","lastName":"Song","suffix":""},{"id":296614429,"identity":"59656708-b5e7-4045-9be4-2aed3c80a933","order_by":4,"name":"Dan-Dan Li","email":"","orcid":"","institution":"Shandong Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"Dan-Dan","middleName":"","lastName":"Li","suffix":""},{"id":296614430,"identity":"9e53bf35-f68b-4c11-b097-4868b75545ff","order_by":5,"name":"Zhao-Qun Li","email":"","orcid":"","institution":"Chinese Academy of Agricultural Science","correspondingAuthor":false,"prefix":"","firstName":"Zhao-Qun","middleName":"","lastName":"Li","suffix":""},{"id":296614431,"identity":"ee75509b-ca5f-4ab5-9074-a9b8af4be270","order_by":6,"name":"Yong-Yu Xu","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAuklEQVRIiWNgGAWjYLCCDwzMYFqCaB2MM0jWwsxDkhaDGzlm0jZ/rOUNDjAfvM3DYJdHUIvkDKCW3LZ0ww0H2JKteRiSiwlq4ZcAaWk4zLjhAI+ZNA/DgcQGQlrYQFos/hy233CA/xtxWsC2MLAdTgTawkacFsmeZ8WWvW3pyTMPsxlbzjFIJqzF4Hjyxhs//ljb9h1vfnjjTYUdYS0MDBwmkOgAR40BYfVAwP74A1HqRsEoGAWjYOQCAAuxNuKxrW6rAAAAAElFTkSuQmCC","orcid":"","institution":"Shandong Agricultural University","correspondingAuthor":true,"prefix":"","firstName":"Yong-Yu","middleName":"","lastName":"Xu","suffix":""},{"id":296614432,"identity":"8ab4f43c-b9ec-467f-a526-8d751dede7b7","order_by":7,"name":"Zhen-Zhen Chen","email":"","orcid":"","institution":"Shandong Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"Zhen-Zhen","middleName":"","lastName":"Chen","suffix":""}],"badges":[],"createdAt":"2024-04-27 10:26:09","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4333561/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4333561/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":55814469,"identity":"344ab173-5f99-4b2c-975f-ed20149ec2ec","added_by":"auto","created_at":"2024-05-03 18:18:57","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":173501,"visible":true,"origin":"","legend":"\u003cp\u003eDiagram of the Y-tube olfactometer apparatus used to test the preference of \u003cem\u003eAleurocanthus spiniferus\u003c/em\u003e for synthetic chemicals\u003c/p\u003e","description":"","filename":"Picture1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-4333561/v1/e4125eafdb6eea9114df5865.jpg"},{"id":55814471,"identity":"019de476-c095-467a-a235-a9eddbd8e530","added_by":"auto","created_at":"2024-05-03 18:18:58","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":541100,"visible":true,"origin":"","legend":"\u003cp\u003ePopulation dynamics of \u003cem\u003eAleurocanthus spiniferus\u003c/em\u003e on four tea cultivars (‘Huangjinya’, ‘Fuding white tea’, ‘Baiye No. 1’, and ‘Longjing 43’) in Tai’an in 2022. (a) adults; (b) nymphs; (c) eggs; (d) pseudopupae. Data are shown as mean ± SE (\u003cem\u003en\u003c/em\u003e =4). Bars indicate standard deviations\u003c/p\u003e","description":"","filename":"Picture2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-4333561/v1/aa4cdafe52a1db2df309a2a5.jpg"},{"id":55814470,"identity":"2e1c3b28-05c5-4662-a19f-86af85207dc7","added_by":"auto","created_at":"2024-05-03 18:18:57","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":295049,"visible":true,"origin":"","legend":"\u003cp\u003e(a) Preference of \u003cem\u003eAleurocanthus spiniferus\u003c/em\u003e adults for healthy and whitefly-infested tea shoots. (b) Responses of \u003cem\u003eAleurocanthus spiniferus\u003c/em\u003e adults to four healthy tea plant cultivar shoots in the Y-tube olfactometer. (c) Responses of \u003cem\u003eAleurocanthus spiniferus\u003c/em\u003eadults to four whitefly-infested tea cultivar shoots in the Y-tube olfactometer. Numbers in brackets represent the percentage of \u003cem\u003eAleurocanthus spiniferus\u003c/em\u003e that were not selected. Asterisks and n.s. indicate significant (*\u003cem\u003eP\u003c/em\u003e\u0026lt; 0.05, **\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.01, ***\u003cem\u003eP \u003c/em\u003e\u0026lt; 0.001) and non-significant preference between the two tea cultivars by chi-square test, respectively\u003c/p\u003e","description":"","filename":"Picture3.jpg","url":"https://assets-eu.researchsquare.com/files/rs-4333561/v1/e9989085f2c79f936d987c69.jpg"},{"id":55814474,"identity":"d8110cf1-0962-4659-87b3-896f51df8dd1","added_by":"auto","created_at":"2024-05-03 18:18:58","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":357931,"visible":true,"origin":"","legend":"\u003cp\u003eRepresentative GC-MS traces of headspace volatiles were released from four tea shoots (‘Huangjinya’, ‘Fuding white tea’, ‘Baiye No. 1’, and ‘Longjing 43’). The identities of the compounds are listed numerically in Table 2\u003c/p\u003e\n\u003cp\u003eIS, internal standard, decanoic acid ethyl ester\u003c/p\u003e","description":"","filename":"Picture4.jpg","url":"https://assets-eu.researchsquare.com/files/rs-4333561/v1/72d30ab712ca2da51224297e.jpg"},{"id":55814473,"identity":"003730bd-9916-487e-b521-ab853fddc345","added_by":"auto","created_at":"2024-05-03 18:18:58","extension":"jpg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":334708,"visible":true,"origin":"","legend":"\u003cp\u003ePreference of adult \u003cem\u003eAleurocanthus spiniferus\u003c/em\u003eto 10 \u003cem\u003eμ\u003c/em\u003eL of hexane solution of six tea shoot volatiles at different concentrations (1, 10, and 100 \u003cem\u003eμ\u003c/em\u003eg/\u003cem\u003eμ\u003c/em\u003eL). All compounds used in the Y-tube olfactometer assays were diluted with hexane. Asterisks indicate significant (*\u003cem\u003eP \u003c/em\u003e\u0026lt; 0.05, **\u003cem\u003eP \u003c/em\u003e\u0026lt; 0.01, ***\u003cem\u003eP \u003c/em\u003e\u0026lt; 0.001) preference between the two odor sources by chi-square test\u003c/p\u003e","description":"","filename":"Picture5.jpg","url":"https://assets-eu.researchsquare.com/files/rs-4333561/v1/430b8a01365bc4481c85e1af.jpg"},{"id":55814475,"identity":"bd0cf04d-07df-42d7-962d-e3d0190bcbbd","added_by":"auto","created_at":"2024-05-03 18:18:58","extension":"jpg","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":304891,"visible":true,"origin":"","legend":"\u003cp\u003eEAG responses of adult \u003cem\u003eAleurocanthus spiniferus \u003c/em\u003eto hexanol (a), nonanal (b), (\u003cem\u003eE\u003c/em\u003e)-2-hexenal (c), (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol (d), linalool (e), and (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene (f) at different concentrations (1, 10, 100, 250, and 500 \u003cem\u003eμ\u003c/em\u003eg/\u003cem\u003eμ\u003c/em\u003eL). Bars indicate standard deviations. Different letters above bars indicate significant differences in relative EAG responses (mV) at different concentrations. \u003cem\u003eP \u003c/em\u003e\u0026lt; 0.05, one-way analysis of variance, Tukey’s-b multiple range test\u003c/p\u003e","description":"","filename":"Picture6.jpg","url":"https://assets-eu.researchsquare.com/files/rs-4333561/v1/de39816b36da274dcc56499e.jpg"},{"id":55814479,"identity":"d3f4c9a6-99e0-438a-b7b1-92ad17cfcf9d","added_by":"auto","created_at":"2024-05-03 18:18:58","extension":"jpg","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":633552,"visible":true,"origin":"","legend":"\u003cp\u003eMean number of adult \u003cem\u003eAleurocanthus spiniferus\u003c/em\u003e in the sticky traps containing six synthetic compounds, hexanol (a), nonanal (b), (\u003cem\u003eE\u003c/em\u003e)-2-hexenal (c), (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol (d), linalool (e), and (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene (f), at different concentrations (1, 10, and 100 \u003cem\u003eμ\u003c/em\u003eg/\u003cem\u003eμ\u003c/em\u003eL) when the traps were placed after 3 d in Jinan in October 2022. Bars indicate standard deviations. Different letters above bars indicate significant differences in the number of adult \u003cem\u003eAleurocanthus spiniferus\u003c/em\u003e at different treatments. \u003cem\u003eP \u003c/em\u003e\u0026lt; 0.05, one-way analysis of variance, Tukey’s-b multiple range test\u003c/p\u003e","description":"","filename":"Picture7.jpg","url":"https://assets-eu.researchsquare.com/files/rs-4333561/v1/20737bf28a3e58d375a3b3a9.jpg"},{"id":55814477,"identity":"84ca7cea-dba1-4bb5-ba0d-84775662392d","added_by":"auto","created_at":"2024-05-03 18:18:58","extension":"jpg","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":644754,"visible":true,"origin":"","legend":"\u003cp\u003eMean number of adult \u003cem\u003eAleurocanthus spiniferus\u003c/em\u003e in the sticky traps containing six synthetic compounds, hexanol (a), nonanal (b), (\u003cem\u003eE\u003c/em\u003e)-2-hexenal (c), (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol (d), linalool (e), and (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene (f), at different concentrations (1, 10, and 100 \u003cem\u003eμ\u003c/em\u003eg/\u003cem\u003eμ\u003c/em\u003eL) when the traps were placed after 7 d in Tai’an in October 2023. Bars indicate standard deviations. Different letters above bars indicate significant differences in the number of adult \u003cem\u003eAleurocanthus spiniferus\u003c/em\u003eat different treatments. \u003cem\u003eP \u003c/em\u003e\u0026lt; 0.05, one-way analysis of variance, Tukey’s-b multiple range test\u003c/p\u003e","description":"","filename":"Picture8.jpg","url":"https://assets-eu.researchsquare.com/files/rs-4333561/v1/b06b0486376e909a79304af0.jpg"},{"id":55814476,"identity":"fe89143d-2efa-457d-84ab-6f0b31a3e44b","added_by":"auto","created_at":"2024-05-03 18:18:58","extension":"jpg","order_by":9,"title":"Figure 9","display":"","copyAsset":false,"role":"figure","size":489500,"visible":true,"origin":"","legend":"\u003cp\u003eMean number of adult \u003cem\u003eAleurocanthus spiniferus\u003c/em\u003e caught on sticky traps baited with seven synthetic blends at different concentrations (1, 10, and 100 \u003cem\u003eμ\u003c/em\u003eg/\u003cem\u003eμ\u003c/em\u003eL) when the traps were placed after 3 d in Jinan in October 2022. (a) blend 1, (b) blend 2, (c) blend 3, (d) blend 4, (e) blend 5, (f) blend 6, and (g) blend 7. Bars indicate standard deviations. Different letters above bars indicate significant differences in the number of adult \u003cem\u003eAleurocanthus camellia\u003c/em\u003e at different treatments. \u003cem\u003eP \u003c/em\u003e\u0026lt; 0.05, one-way analysis of variance, Tukey’s-b multiple range test\u003c/p\u003e","description":"","filename":"Picture9.jpg","url":"https://assets-eu.researchsquare.com/files/rs-4333561/v1/f9784a5d5680d6ac5b4db724.jpg"},{"id":55814478,"identity":"8edc0467-d405-4bbf-8ead-dd7f8fa690ac","added_by":"auto","created_at":"2024-05-03 18:18:58","extension":"jpg","order_by":10,"title":"Figure 10","display":"","copyAsset":false,"role":"figure","size":506296,"visible":true,"origin":"","legend":"\u003cp\u003eMean number of adult \u003cem\u003eAleurocanthus spiniferus\u003c/em\u003e caught on sticky traps baited with seven synthetic blends at different concentrations (1, 10, and 100 \u003cem\u003eμ\u003c/em\u003eg/\u003cem\u003eμ\u003c/em\u003eL) when the traps were placed after 7 d in Tai’an in October 2023. (a) blend 1, (b) blend 2, (c) blend 3, (d) blend 4, (e) blend 5, (f) blend 6, and (g) blend 7. Bars indicate standard deviations. Different letters above bars indicate significant differences in the number of adult \u003cem\u003eAleurocanthus camellia\u003c/em\u003e at different treatments. \u003cem\u003eP \u003c/em\u003e\u0026lt; 0.05, one-way analysis of variance, Tukey’s-b multiple range test\u003c/p\u003e","description":"","filename":"Picture10.jpg","url":"https://assets-eu.researchsquare.com/files/rs-4333561/v1/b182cdee4a273b03961869fb.jpg"},{"id":55814480,"identity":"c629aafb-62e0-40b5-8f48-09aa7dcf2f65","added_by":"auto","created_at":"2024-05-03 18:18:58","extension":"jpg","order_by":11,"title":"Figure 11","display":"","copyAsset":false,"role":"figure","size":635089,"visible":true,"origin":"","legend":"\u003cp\u003eNumber of adult \u003cem\u003eAleurocanthus spiniferus\u003c/em\u003e in yellow sticky traps containing blend 4 at 100 \u003cem\u003eμ\u003c/em\u003eg/\u003cem\u003eμ\u003c/em\u003eL and blend 6 at 100 \u003cem\u003eμ\u003c/em\u003eg/\u003cem\u003eμ\u003c/em\u003eL at different times after application in 2022 (a and b) and 2023 (c and d). Bars indicate standard deviations. Different letters above bars indicate significant differences in the number of adult \u003cem\u003eAleurocanthus spiniferus \u003c/em\u003ein the different treatments. *\u003cem\u003eP\u003c/em\u003e\u0026lt;0.05; **\u003cem\u003eP\u003c/em\u003e\u0026lt;0.01; ***\u003cem\u003eP\u003c/em\u003e\u0026lt;0.001; ns indicate non-significant difference. All experiments were done in three replicates, two-tailed student’s \u003cem\u003et\u003c/em\u003e test\u003c/p\u003e","description":"","filename":"Picture11.jpg","url":"https://assets-eu.researchsquare.com/files/rs-4333561/v1/442163e5f816449032adb696.jpg"},{"id":60075756,"identity":"6d05f1a9-3b93-4238-b638-dc78ffe847ab","added_by":"auto","created_at":"2024-07-11 12:25:56","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":5781468,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4333561/v1/169af80a-a7e0-4ee0-9e40-ef701787f2dc.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Field application of tea volatiles mediating host selection of Aleurocanthus spiniferus","fulltext":[{"header":"Key Message","content":"\u003cp\u003e1.\u0026nbsp;\u003cem\u003eA. spiniferus\u003c/em\u003e occurrence varies significantly among different tea cultivars within the same plantation.\u003c/p\u003e\n\u003cp\u003e2.\u0026nbsp;It was speculated that adult \u003cem\u003eA. spiniferus\u003c/em\u003e may be differentially attracted by different cultivars.\u003c/p\u003e\n\u003cp\u003e3.\u0026nbsp;The odor components in tea volatiles play a crucial role in the host selection of adult\u0026nbsp;\u003cem\u003eA. spiniferus.\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e4.\u0026nbsp;Two attractants based on tea volatiles were found to be highly effective in trapping adult \u003cem\u003eA. spiniferus.\u003c/em\u003e\u003c/p\u003e"},{"header":"1. INTRODUCTION","content":"\u003cp\u003e \u003cem\u003eCamellia sinensis\u003c/em\u003e (L.) Kuntze (Theaceae), also known as tea, is an important economic crop, widely cultivated in tropical and subtropical regions such as China, Japan, and Kenya (Zhao et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). \u003cem\u003eAleurocanthus spiniferus\u003c/em\u003e (Quaintance) (Hemiptera: Aleyrodidae) is a major pest of tea plants (Tang et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Adults generally congregate to feed on tea shoots and lay eggs. In addition, first-instar nymphs can be active, while all other nymphal stages are stationary and feed on leaves, directly affecting tea yield and quality. Tea sooty mould, \u003cem\u003eNeocapnodium theae\u003c/em\u003e, is caused by the excrement of this pest, which reduces the photosynthetic capability of tea leaves and affects tea plant growth (Byrne and Bellows \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1991\u003c/span\u003e). \u003cem\u003eAleurocanthus spiniferus\u003c/em\u003e is widely distributed in China, Japan, and Indonesia (Uesugi et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Bragard et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Yang et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), which has become a common and major pest in Chinese tea plantations. Currently, chemical pesticides are often used to control \u003cem\u003eA. spiniferus\u003c/em\u003e (Hazarika et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Tian et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Therefore, safer and more effective prevention and control methods are urgently needed.\u003c/p\u003e \u003cp\u003eUsing insecticides or repellent components from plant volatiles can effectively prevent pests from selecting their hosts. Host plant location and selection is a crucial task for herbivorous insects in their search for resources, mates, and oviposition sites; and olfaction typically plays an important role in this process (Song et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Adults detect volatile organic compounds (VOCs) emitted by plants in a specific way, discriminating between different resistant plants or cultivars based on the types and proportions of volatile compounds (Ranger et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Huang et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Volatiles from non-host plants or resistant cultivars contain repellent components to prevent herbivorous insect infestation (Yuan et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). For example, \u003cem\u003eRosmarinus officinalis\u003c/em\u003e, a non-host plant with specifically bioactive compounds in its volatiles that repelling \u003cem\u003eEctropis obliqua\u003c/em\u003e adults (Zhang et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2015\u003c/span\u003e); ocimene, a dominant component of resistant wheat varieties repelled \u003cem\u003eSitodiplosis mosellana\u003c/em\u003e females (Huang et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Similarly, attractive components in host volatile blends are the crucial reason why host plants or susceptible cultivars are more susceptible to pest infestation (Birkett et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Yang et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Xiu et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). A key volatile, (\u003cem\u003eZ\u003c/em\u003e)-3-hexenyl-acetate, mediate the host and oviposition preference of \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e on maize (Wang et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Some specific compounds have been successfully used for population monitoring and integrated pest management (Li et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Chen et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). A binary repellent containing DMDS and 1,8-cineole, for instance, applied by a slow-release method against \u003cem\u003eEmpoasca onukii\u003c/em\u003e in a tea plantation (Cai et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn tea plantations, there are observable differences in the occurrence of \u003cem\u003eA. spiniferus\u003c/em\u003e on four tea cultivars. \u0026lsquo;Huangjinya\u0026rsquo; and \u0026lsquo;Fuding white tea\u0026rsquo; are more susceptible and suitable for \u003cem\u003eA. spiniferus\u003c/em\u003e (Tian et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Lu et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; He et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), while \u0026lsquo;Baiye No. 1\u0026rsquo; and \u0026lsquo;Longjing 43\u0026rsquo; are common resistant cultivars (Liang et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). To elucidate the potential role of volatiles in host selection, this study evaluated the selection preferences of adult \u003cem\u003eA. spiniferus\u003c/em\u003e towards four tea cultivars with different resistance properties. Differences in volatile components among the four cultivars with different resistances were analyzed using GC-MS. The odor components in tea volatiles, which play a crucial role in the host selection behavior of adult \u003cem\u003eA. spiniferus\u003c/em\u003e, were analyzed by electrophysiological (EAG) and behavioral measurements. Finally, these compounds and their mixtures were used as attractants in different tea plantations in two years to identify the most effective attractants. This study provides theoretical support for the development of monitoring and control technology for \u003cem\u003eA. spiniferus\u003c/em\u003e.\u003c/p\u003e"},{"header":"2. MATERIALS AND METHODS","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.1 Tea cultivars and insect collection\u003c/h2\u003e \u003cp\u003eFour tea cultivars, \u0026lsquo;Baiye No. 1\u0026rsquo;, \u0026lsquo;Longjing 43\u0026rsquo;, \u0026lsquo;Fuding white tea\u0026rsquo;, and \u0026lsquo;Huangjinya\u0026rsquo;, were used for headspace volatile analysis, assessment of behavioral preferences and population dynamics of adult \u003cem\u003eA. spiniferus\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eAdult \u003cem\u003eA. spiniferus\u003c/em\u003e were collected from the tea cultivars \u0026lsquo;Huangjinya\u0026rsquo; that were maintained in the tea plantations managed by Shandong Qianrun Ecological Agriculture Development Co., Ltd. in Tai\u0026rsquo;an, Shandong Province, China (32\u0026deg;08\u0026prime;N, 117\u0026deg;43\u0026prime;E). Before experiments, adults were acclimated for one day on four different tea cultivars in environmental incubators at 26\u0026thinsp;\u0026plusmn;\u0026thinsp;2\u0026deg;C with 70\u0026thinsp;\u0026plusmn;\u0026thinsp;5% RH and a 16L:8D photoperiod. Insect preference tests for two cultivars, the test insects were kept on the corresponding cultivars, 50/50 for each. Insects for the other laboratory tests were kept on \u0026lsquo;Huangjinya\u0026rsquo;. Due to the low sex ratio of males, we are unable to obtain enough males for subsequent experiments. Therefore, females and males were mixed for behavioral preference and EAG tests.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e2.2 Population dynamics of \u003cem\u003eA. spiniferus\u003c/em\u003e on four tea cultivars\u003c/h2\u003e \u003cp\u003eThe experiment was conducted in the same tea plantations as described above from May to November 2022. As typical of their habit of tending to aggregate towards tender leaves and buds, adult \u003cem\u003eA. spiniferus\u003c/em\u003e were mainly active in the upper branches of the tea plants. Therefore, the second and third leaves below the bud were selected for counting the density of adults and eggs in the four tea cultivars. The number of nymphs and pseudopupae on the middle and lower leaves of the tea plant was investigated. Specifically, one hundred leaves were randomly selected for counting the number of \u003cem\u003eA. spiniferus\u003c/em\u003e adults, eggs, nymphs, and pseudopupae as a biological replicate, respectively. There were four replicates for each tea cultivar.\u003c/p\u003e \u003cp\u003e2.3 Y-tube test for behavioral responses of adult \u003cem\u003eA. spiniferus\u003c/em\u003e to tea shoots and synthetic volatile compounds.\u003c/p\u003e \u003cp\u003eThe Y-tube olfactometer test has been refined with reference to relevant studies (Han and Chen \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2002\u003c/span\u003e), and the apparatus is shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e. Bioassays were conducted using a dual-choice glass Y-tube olfactometer, which consisted of a central tube and two lateral arms (15 cm long, 13 mm diameter) with an inside angle of 60\u0026deg;. For each paired bioassay, two different odor sources were placed in two odor bottles, which were connected to an activated carbon-filtered, humidified air source and the two arms of a Y-tube through Teflon\u0026reg; tubing. The airflow rate through each arm was 100 mL/min by the pre-experiment, measured using an LZB-3WB rotor meter.\u003c/p\u003e \u003cp\u003eTwo-year-old tea plants were used for tea plant testing. Two treatments were set up in the experiment: healthy plants and whitefly-infested plants. Pretreatment for whitefly-infested plants was as follows: eight healthy tea plants were placed in a nylon screen cage (50 cm \u0026times; 50 cm \u0026times; 50 cm) with 500 (\u0026plusmn;\u0026thinsp;10) \u003cem\u003eA. spiniferus\u003c/em\u003e adults per plant for each cultivar. Adults were removed after 4 hours of feeding, and tea plants were used when nymphs had developed to the 2nd-3rd instar. Eight healthy tea plants for each cultivar were placed in a nylon screen cage (50 cm \u0026times; 50 cm \u0026times; 50 cm) as a healthy plant treatment.\u003c/p\u003e \u003cp\u003eTo prepare the test chemicals, six standard synthetic chemicals were dissolved in hexane at three different concentrations (1, 10, and 100 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL). Odorants (10 \u003cem\u003e\u0026micro;\u003c/em\u003eL) were added dropwise to filter paper strips (2 \u0026times; 2 cm), then allowed to evaporate for 20 s and placed in an odor bottle. Filter paper strips (2 \u0026times; 2 cm) containing hexane were placed in another odor bottle.\u003c/p\u003e \u003cp\u003eExperimental adults were starved for 1 hour before the bioassay. A total of 80 insects were tested per bioassay, and each individual was only used once. In each test, a choice was recorded if the adult moved 5 cm past the Y-junction within ten minutes and remained there for at least one minute. Otherwise, it was recorded as no-choice. At the end of the odor source test, the Y-tube, odor bottles, and other glass components were cleaned with acetone and dried before reuse. The Y-tube olfactometer was placed in the laboratory with diffuse light at a temperature of 26\u0026thinsp;\u0026plusmn;\u0026thinsp;2\u0026deg;C and a relative humidity of 70% \u0026plusmn; 5% at about 09:30\u0026thinsp;\u0026minus;\u0026thinsp;15:30 hrs.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003e2.4 Volatile collection and analysis\u003c/h2\u003e \u003cp\u003eHeadspace volatiles from tea plants were collected using the dynamic headspace absorption technique (Sun et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Zhang et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). The Y-tube tests showed that the pre-infection of \u003cem\u003eA. spiniferus\u003c/em\u003e nymphs did not significantly affect the preference of the subsequent adult \u003cem\u003eA. spiniferus.\u003c/em\u003e Six 2-year-old healthy tea plants without nymphs were placed in the glass cylinder chamber. The inlet, outlet, and all interfaces of the glass cylinder chamber were sealed with sealing film (Bemis, Inc., Neenah, Wisconsin, USA). The airflow entered the glass cylinder chamber after passing through the silicone desiccant, activated carbon, and rotameter at 200 mL/min. The gas-carrying volatiles were extracted after collection onto the glass tube containing Super Q absorbent (35 mg; 80/100 mesh; Alltech Associates., Inc., Deerfield, IL, USA).\u003c/p\u003e \u003cp\u003eAll components of the volatile collection apparatus were connected by Teflon\u0026reg; tubing. After 1 hour of collection, the trapped tea shoot volatiles were extracted from the Super Q tube with chromatographically pure hexane (500 \u003cem\u003e\u0026micro;\u003c/em\u003eL). The eluting solution was stored in a glass vial to which decanoic acid ethyl ester (5 \u003cem\u003e\u0026micro;\u003c/em\u003eL) at a concentration of 10\u003csup\u003e\u0026minus;\u0026thinsp;4\u003c/sup\u003e g/mL was added as an internal standard. The sample was stored at -20\u0026deg;C for qualitative and quantitative analysis of volatile components.\u003c/p\u003e \u003cp\u003eThe GC was an Agilent 7890B model, and the MS was the 7000D model (Agilent, Inc., PaloAlto, CA, USA). Samples were analyzed on an HP-5ms (30 m \u0026times; 0.25 mm i.d., 0.25 \u003cem\u003e\u0026micro;\u003c/em\u003em film thickness). The solvent delay was 3 minutes. The inlet temperature was 230\u0026deg;C, and the GC/MS interface temperature was 280\u0026deg;C. The analysis program was as follows: the oven temperature was programmed at 40\u0026deg;C for 0 minutes, increased at 3\u0026deg;C/min to 190\u0026deg;C for 2 min, then increased at 10\u0026deg;C/min to 290\u0026deg;C and held for 3 minutes. Helium (99.999% purity) was used as the carrier gas at a flow rate of 1 ml per minute. The mass spectrometer was operated in the electron-impact (EI, at 70 eV) scan mode. The scan rate was twice per second. Qualitative analysis of the volatile components was processed by Agilent MassHunter Qualitative Analysis Navigator software. Compounds were identified by comparing the obtained spectra with the spectra of reference compounds from the National Institute of Standards and Technology (NIST17), as well as retention times in the relevant literature on tea shoot volatiles (Mu et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Sun et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Cai et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Cai et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Magsi et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The contents of each volatile compound were calculated by comparing their GC total ion current peak areas with the peak area of the internal standard.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003e2.5 Chemicals\u003c/h2\u003e \u003cp\u003eHexane (analytical pure) served as solvent and negative control was purchased from Tianjin Kaitong Chemical Reagent Co. Ltd. (Tianjin, China). Hexanol (99%, CAS: 111-27‐3), linalool (98%, CAS: 78‐70‐6), (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol (98%, CAS: 928‐96‐1), nonanal (96%, CAS: 124‐19‐6), (\u003cem\u003eE\u003c/em\u003e)-2-hexenal (98%, CAS: 6728‐26‐3) was purchased from Macklin Inc. (Shanghai, China), (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene (96%, CAS: 21499‐64‐9) was synthesized by the College of Chemistry, Shandong Agricultural University.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003e2.6 EAG experiments\u003c/h2\u003e \u003cp\u003eThe EAG test was based on the same method used for the determination of antennally-active volatiles for tea geometrids (Yuan et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). The EAG system (Syntech, Ltd., Hilversum, The Netherlands) consisted of an IDAC-4 interface box, a CS-55 air stimulus controller, and a desktop computer. To prepare the test chemicals, six standard chemical compounds were dissolved in hexane at five different concentrations (1, 10, 100, 250, and 500 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL). Exactly 10 \u003cem\u003e\u0026micro;\u003c/em\u003eL of the tested solution was placed onto a white filter paper strip (0.5 \u0026times; 6 cm), the solvent was allowed to evaporate for 30 seconds, then the strip was carefully transferred to a Pasteur pipette cartridge (14.5 cm long). Hexane (10 \u003cem\u003e\u0026micro;\u003c/em\u003eL) was used as the control using the same method. The adult \u003cem\u003eA. spiniferus\u003c/em\u003e was chilled in an ice box and removed immediately after the induction of torpor. An antenna was dissected under the stereoscopic microscope (Nikon, Ltd., Japan), and then the reference and recording electrodes were attached to the base and tip of the antenna using conductive adhesive, respectively. To perform the stimulation, the tip of a Pasteur pipette cartridge was inserted into a small hole in the wall of a steel tube (14 cm long, 8 mm diameter), approximately 3 mm diameter, located approximately 11 cm from the end of the tube. A regulated airflow (10 mL/s) entered the steel tube and was directed toward the antenna for chemical exposure. The antenna was located approximately 1 cm from the end of the steel tube. Each antenna was randomly stimulated for testing with three standard chemicals. The stimulation sequence consisted of a control stimulus, followed by three test chemical stimuli, and then another control stimulus. The duration of each stimulation was set to 0.5 s, with an interval of 60 s between two consecutive stimuli (Delorme and Payne \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1990\u003c/span\u003e). A total of 15 antennas were measured for each concentration level of volatiles. Only one antenna was randomly selected per adult. All EAG tests were conducted at a temperature of 26\u0026thinsp;\u0026plusmn;\u0026thinsp;2\u0026deg;C and a relative humidity of 70\u0026thinsp;\u0026plusmn;\u0026thinsp;5% from 09:30\u0026thinsp;\u0026minus;\u0026thinsp;15:30 hrs.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003e2.7 Field trial\u003c/h2\u003e \u003cp\u003eIn 1st October 2022, the field trial was conducted in the tea plantation in Gongjiazhuang Village, Jinan City, Shandong Province, China (36\u0026deg;31\u0026prime;N, 117\u0026deg;41\u0026prime;E) with low-density of \u003cem\u003eA. spiniferus\u003c/em\u003e adult. In 9th October 2023, it was conducted at the same tea plantation as described in population dynamics experiment, with high-density of \u003cem\u003eA. spiniferus\u003c/em\u003e adult, coincident with the third peak of \u003cem\u003eA. spiniferus\u003c/em\u003e adult occurrence in tea plantations. The average temperature and relative humidity during the field trials were 8\u0026ndash;15\u0026deg;C and 55\u0026ndash;70% RH, respectively, and the weather was sunny or cloudy with a gentle breeze. Attractants for tests were formulated as six single-component and seven different mixed-component blends (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) diluted with hexane to three concentrations (1, 10, and 100 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL) and adsorbed onto a rubber dispenser (Keyun, Co. Ltd., Henan, China), each containing 50 mg of the active ingredient (a.i.). The following treatments were included: (i) synergistic bait traps: attractants\u0026thinsp;+\u0026thinsp;yellow sticky traps (25\u0026times;20 cm, Lvpusen Technology, Ltd., Quanzhou, China); (ii) yellow sticky traps; (iii) single attractants: attractants\u0026thinsp;+\u0026thinsp;white sticky traps (25\u0026times;20 cm, Lvpusen Technology, Ltd., Quanzhou, China); and white sticky trap was used as a control. All sticky traps were attached to bamboo poles, and placed 5\u0026ndash;10 cm from the tea shoots; the spacing between each sticky trap was greater than 20 m to minimize the interference between compounds. The trial was conducted in three plots of approximately 1 ha each, with the above four treatments randomized between plots. The distance between the two plots was at least 15 m. The number of adult \u003cem\u003eA. spiniferus\u003c/em\u003e on sticky traps was counted on days 3, 7, 14, and 23 in 2022 and on days 7, 14, 23, and 30 in 2023 after the start of the experiment, and sticky traps were replaced after each survey.\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\u003eComposition of seven blends formulated as potential attractants for adult \u003cem\u003eAleurocanthus spiniferus\u003c/em\u003e\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"2\"\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 \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCode\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eComponent\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBlend 1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHexanol, nonanal, (\u003cem\u003eE\u003c/em\u003e)-2-hexenal, (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol, linalool, (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBlend 2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHexanol, (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol, linalool, (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBlend 3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNonanal, (\u003cem\u003eE\u003c/em\u003e)-2-hexenal\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBlend 4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eLinalool, (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBlend 5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e(\u003cem\u003eZ\u003c/em\u003e)-3-hexenol, (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBlend 6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e(\u003cem\u003eZ\u003c/em\u003e)-3-hexenol, linalool\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBlend 7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHexanol, (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003e2.8 Data analysis\u003c/h2\u003e \u003cp\u003eAll data were analyzed in Graphpad Prism 9 software (GraphPad Software, San Diego, CA, USA). Data on population dynamics, volatile quantities, relative EAG response, and field trials were analyzed by one-way analysis of variance (ANOVA). The normality of the residuals was tested for all data using the Shapiro-Wilk test. Means were compared using Tukey\u0026rsquo;s-b multiple range test or the Games-Howell test, depending on whether the treatment and control variances were equal (Shapiro-Wilks test, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.05) or unequal (Shapiro-Wilks test, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05). The Y-tube olfactometer behavioral test data were analyzed using a chi-squared test (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) to determine differences between pairs of treatments. Non-choice individuals were recorded but excluded from the statistical analysis. Data on lure longevity time was analyzed by two-tailed student\u0026rsquo;s \u003cem\u003et\u003c/em\u003e test.\u003c/p\u003e \u003c/div\u003e"},{"header":"3. RESULTS","content":"\u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003e3.1 Population dynamics of \u003cem\u003eA. spiniferus\u003c/em\u003e on four tea varieties\u003c/h2\u003e \u003cp\u003eThe peak flight season occurred in June. The number of \u003cem\u003eA. spiniferus\u003c/em\u003e adults and eggs peaked on 14 June 2022, with significant differences in occurrence among different cultivars (adults: \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;34.71, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001; eggs: \u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;63.34, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). The number of adults on \u0026lsquo;Huangjinya\u0026rsquo; reached 2597.20\u0026thinsp;\u0026plusmn;\u0026thinsp;386.99, which was significantly higher than that of \u0026lsquo;Fuding white tea\u0026rsquo; (807.60\u0026thinsp;\u0026plusmn;\u0026thinsp;57.86), \u0026lsquo;Baiye No.1\u0026rsquo; (156.20\u0026thinsp;\u0026plusmn;\u0026thinsp;14.21), and \u0026lsquo;Longjing 43\u0026rsquo; (149.60\u0026thinsp;\u0026plusmn;\u0026thinsp;20.25) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea). Egg abundance showed the same trend, with 3919.80\u0026thinsp;\u0026plusmn;\u0026thinsp;293.71 for \u0026lsquo;Huangjinya\u0026rsquo;, slightly lower for \u0026lsquo;Fuding white tea\u0026rsquo; at 2587.60\u0026thinsp;\u0026plusmn;\u0026thinsp;321.15, and only 504.80\u0026thinsp;\u0026plusmn;\u0026thinsp;45.04 and 275.00\u0026thinsp;\u0026plusmn;\u0026thinsp;21.47 for \u0026lsquo;Baiye No.1\u0026rsquo; and \u0026lsquo;Longjing 43\u0026rsquo; (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ec). On 14 June, nymph abundance showed significant differences among the different cultivars (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;102.16, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), with \u0026lsquo;Huangjinya\u0026rsquo; and \u0026lsquo;Fuding white tea\u0026rsquo; was 3111.80\u0026thinsp;\u0026plusmn;\u0026thinsp;225.70 and 1359.80\u0026thinsp;\u0026plusmn;\u0026thinsp;142.46, respectively; while \u0026lsquo;Baiye No.1\u0026rsquo; and \u0026lsquo;Longjing 43\u0026rsquo; were only 277.20\u0026thinsp;\u0026plusmn;\u0026thinsp;29.14 and 189.40\u0026thinsp;\u0026plusmn;\u0026thinsp;18.29, respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eb). Pseudopupae abundance on \u0026lsquo;Huangjinya\u0026rsquo; and \u0026lsquo;Fuding white tea\u0026rsquo; peaked on 31 May (\u0026lsquo;Huangjinya\u0026rsquo;: 2328.40\u0026thinsp;\u0026plusmn;\u0026thinsp;168.65; \u0026lsquo;Fuding white tea\u0026rsquo;: 1439.40\u0026thinsp;\u0026plusmn;\u0026thinsp;143.56) and 14 June (\u0026lsquo;Huangjinya\u0026rsquo;: 1960.00\u0026thinsp;\u0026plusmn;\u0026thinsp;272.04; \u0026lsquo;Fuding white tea\u0026rsquo;: 1697.20\u0026thinsp;\u0026plusmn;\u0026thinsp;206.26), at which time the number of pseudopupae on \u0026lsquo;Baiye No.1\u0026rsquo; (31 May: 250.40\u0026thinsp;\u0026plusmn;\u0026thinsp;19.15; 14 June: 343.80\u0026thinsp;\u0026plusmn;\u0026thinsp;57.64) and \u0026lsquo;Longjing 43\u0026rsquo; (31 May: 200.20\u0026thinsp;\u0026plusmn;\u0026thinsp;23.31; 14 June: 266.80\u0026thinsp;\u0026plusmn;\u0026thinsp;22.92) was relatively low (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ed). Based on almost one year of investigation, it was tentatively determined that the selectivity of \u003cem\u003eA. spiniferus\u003c/em\u003e on four tea cultivars under the same environment in a tea plantation was as follows: \u0026lsquo;Huangjinya\u0026rsquo; \u0026gt; \u0026lsquo;Fuding white tea\u0026rsquo; \u0026gt; \u0026lsquo;Baiye No.1\u0026rsquo; \u0026gt; \u0026lsquo;Longjing 43\u0026rsquo;.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003e3.2 Preference of \u003cem\u003eA. spiniferus\u003c/em\u003e among four tea cultivars.\u003c/h2\u003e \u003cp\u003eTo further clarify the preference of adult \u003cem\u003eA. spiniferus\u003c/em\u003e among the tested tea cultivars, we conducted Y-tube olfactometer experiments. The results showed that the pre-infection of \u003cem\u003eA. spiniferus\u003c/em\u003e nymphs did not significantly affect the preference of the subsequent adult \u003cem\u003eA. spiniferus\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea). The preference of adult \u003cem\u003eA. spiniferus\u003c/em\u003e for \u0026lsquo;Huangjinya\u0026rsquo; was significantly higher than that for the other three cultivars. When paired with \u0026lsquo;Baiye No.1\u0026rsquo; and \u0026lsquo;Longjing 43\u0026rsquo;, adult \u003cem\u003eA. spiniferus\u003c/em\u003e tended to choose \u0026lsquo;Fuding white tea\u0026rsquo;. There was no significant difference in the preference of adult \u003cem\u003eA. spiniferus\u003c/em\u003e between \u0026lsquo;Baiye No.1\u0026rsquo; and \u0026lsquo;Longjing 43\u0026rsquo; (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eb and \u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ec).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003e3.3 Volatile profiles of tested tea cultivars.\u003c/h2\u003e \u003cp\u003eA total of 14 compounds were identified, 11 of which were common to all four cultivars. The content of hexanol (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;10.86, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.003), (\u003cem\u003eE\u003c/em\u003e)-2-hexenal (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;4.49, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.04), and 7-methyl-heptadecane (\u003cem\u003eF\u003c/em\u003e\u0026thinsp;=\u0026thinsp;7.16, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.012) in the volatiles of the shoots from four cultivars showed significant differences. Hexanol and (\u003cem\u003eE\u003c/em\u003e)-2-hexenal were dominant in two resistant cultivars (\u0026lsquo;Baiye No. 1\u0026rsquo; and \u0026lsquo;Longjing 43\u0026rsquo;). There was no evidence that 7-methyl-heptadecane is a behaviorally active volatile organic compound in tea plants, thus this compound was not included in this study. Additionally, linalool was only present in the susceptible \u0026lsquo;Fuding white tea\u0026rsquo;. Nonanol was only absent in the resistant \u0026lsquo;Longjing 43\u0026rsquo;. (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene was only present in the susceptible \u0026lsquo;Huangjinya\u0026rsquo;, while (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol was only absent in the \u0026lsquo;Huangjinya\u0026rsquo; (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e and Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eIS, internal standard, decanoic acid ethyl ester\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\u003eIdentified volatile organic compounds (VOCs) emitted from four tea shoots (\u0026lsquo;Huangjinya\u0026rsquo;, \u0026lsquo;Fuding white tea\u0026rsquo;, \u0026lsquo;Baiye No. 1\u0026rsquo;, and \u0026lsquo;Longjing 43\u0026rsquo;) and mean emission amount \u003csup\u003ea\u003c/sup\u003e (ng plant\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e h\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" 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=\"left\" 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=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eVolatile compounds\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eRetention time (min)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"4\" nameend=\"c6\" namest=\"c3\"\u003e \u003cp\u003eConcentration (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SE) (ng plant\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e h\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eBaiye No. 1\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFuding white tea\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eHuangjinya\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eLongjing 43\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e1. Hexanol\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e3.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e6711.92\u0026thinsp;\u0026plusmn;\u0026thinsp;561.98 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3727.79\u0026thinsp;\u0026plusmn;\u0026thinsp;959.26 ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1667.67\u0026thinsp;\u0026plusmn;\u0026thinsp;322.83 b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e5633.52\u0026thinsp;\u0026plusmn;\u0026thinsp;688.38 a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e2. (\u003cem\u003eE\u003c/em\u003e)-2-Hexenal\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e7.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e235.80\u0026thinsp;\u0026plusmn;\u0026thinsp;49.13 ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e167.61\u0026thinsp;\u0026plusmn;\u0026thinsp;21.43 ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e90.83\u0026thinsp;\u0026plusmn;\u0026thinsp;21.33 b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e249.50\u0026thinsp;\u0026plusmn;\u0026thinsp;37.39 a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e3. (\u003cem\u003eZ\u003c/em\u003e)-3-Hexenol\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e7.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e48.28\u0026thinsp;\u0026plusmn;\u0026thinsp;4.37 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e29.11\u0026thinsp;\u0026plusmn;\u0026thinsp;4.10 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eND\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e32.96\u0026thinsp;\u0026plusmn;\u0026thinsp;4.83 a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e4. Benzaldehyde\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e8.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e567.62\u0026thinsp;\u0026plusmn;\u0026thinsp;124.79 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e406.04\u0026thinsp;\u0026plusmn;\u0026thinsp;84.73 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e172.94\u0026thinsp;\u0026plusmn;\u0026thinsp;88.29 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e235.60\u0026thinsp;\u0026plusmn;\u0026thinsp;35.35 a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e5. 2-Ethyl-1-hexanol\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e11.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e228.87\u0026thinsp;\u0026plusmn;\u0026thinsp;64.03 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e250.96\u0026thinsp;\u0026plusmn;\u0026thinsp;51.94 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e220.90\u0026thinsp;\u0026plusmn;\u0026thinsp;61.29 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e425.11\u0026thinsp;\u0026plusmn;\u0026thinsp;67.08 a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e6. (\u003cem\u003eE\u003c/em\u003e)-Ocimene\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e12.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1045.18\u0026thinsp;\u0026plusmn;\u0026thinsp;398.30 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e789.52\u0026thinsp;\u0026plusmn;\u0026thinsp;158.49 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e566.01\u0026thinsp;\u0026plusmn;\u0026thinsp;154.63 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e539.97\u0026thinsp;\u0026plusmn;\u0026thinsp;169.47 a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e7. Linalool\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e15.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eND\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e96.29\u0026thinsp;\u0026plusmn;\u0026thinsp;23.63\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eND\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eND\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e8. Nonanol\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e15.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e60.37\u0026thinsp;\u0026plusmn;\u0026thinsp;18.47 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e34.35\u0026thinsp;\u0026plusmn;\u0026thinsp;4.22 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e14.54\u0026thinsp;\u0026plusmn;\u0026thinsp;3.72 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eND\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e9. (\u003cem\u003eZ\u003c/em\u003e)-3-Hexenyl butanoate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e19.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1025.41\u0026thinsp;\u0026plusmn;\u0026thinsp;248.76 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e858.78\u0026thinsp;\u0026plusmn;\u0026thinsp;149.65 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e689.06\u0026thinsp;\u0026plusmn;\u0026thinsp;179.84 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e784.68\u0026thinsp;\u0026plusmn;\u0026thinsp;237.15 a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e10. 7-Methyl-heptadecane\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e26.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e681.58\u0026thinsp;\u0026plusmn;\u0026thinsp;202.02 b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e636.01\u0026thinsp;\u0026plusmn;\u0026thinsp;82.64 b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1815.75\u0026thinsp;\u0026plusmn;\u0026thinsp;137.89 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1355.68\u0026thinsp;\u0026plusmn;\u0026thinsp;336.37 ab\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e11. (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-Farnesene\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e30.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eND\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eND\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e34.25\u0026thinsp;\u0026plusmn;\u0026thinsp;2.60\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eND\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e12. Unknown\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e34.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e732.53\u0026thinsp;\u0026plusmn;\u0026thinsp;236.59 b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2160.95\u0026thinsp;\u0026plusmn;\u0026thinsp;319.25 ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3763.17\u0026thinsp;\u0026plusmn;\u0026thinsp;585.63 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3819.90\u0026thinsp;\u0026plusmn;\u0026thinsp;680.01 a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e13. Cadalene\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e40.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e814.17\u0026thinsp;\u0026plusmn;\u0026thinsp;256.77 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e963.19\u0026thinsp;\u0026plusmn;\u0026thinsp;174.40 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e766.72\u0026thinsp;\u0026plusmn;\u0026thinsp;159.95 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1713.63\u0026thinsp;\u0026plusmn;\u0026thinsp;303.49 a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e14. Caffeine\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e46.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e227.83\u0026thinsp;\u0026plusmn;\u0026thinsp;47.99 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e303.41\u0026thinsp;\u0026plusmn;\u0026thinsp;41.73 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e252.29\u0026thinsp;\u0026plusmn;\u0026thinsp;50.87 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e415.13\u0026thinsp;\u0026plusmn;\u0026thinsp;64.31 a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"6\"\u003ea Data in the table are presented as mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SE (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;3), and those followed by different letters in the same row indicate significant differences at 0.05 level by Tukey\u0026rsquo;s-b multiple range test. \u0026ldquo;ND\u0026rdquo; denotes that compounds were not detected\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003e3.4 Preference of \u003cem\u003eA. spiniferus\u003c/em\u003e for synthetic compounds of tea volatiles.\u003c/h2\u003e \u003cp\u003eAll six compounds were able to elicit an intense response in adult \u003cem\u003eA. spiniferus\u003c/em\u003e, and the response was concentration-dependent. Adults were significantly attracted to hexanol at 100 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL; to linalool at 100 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL; to (\u003cem\u003eE\u003c/em\u003e)-2-hexenal at 100 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL, 10 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL; to (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene at 100 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL, 10 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL, 1 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL. In contrast, they were repelled by both nonanal and (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol at all concentrations tested concentrations ranging from 1 to 100 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003e3.5 EAG responses elicited by synthetic compounds of tea volatiles.\u003c/h2\u003e \u003cp\u003eAmong six synthetic compounds of tea volatiles tested, four were able to elicit an obvious EAG response at 100 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL, including nonanol (relative EAG value\u0026thinsp;=\u0026thinsp;0.12 mV) (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003eb), (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol (relative EAG value\u0026thinsp;=\u0026thinsp;0.14 mV) (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ed), linalool (relative EAG value\u0026thinsp;=\u0026thinsp;0.13 mV) (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ee), and (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene (relative EAG value\u0026thinsp;=\u0026thinsp;0.11 mV) (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ef). The other two compounds, hexanol and (\u003cem\u003eE\u003c/em\u003e)-2-hexenal, elicited very weak or no response at all tested concentrations (relative EAG value\u0026thinsp;\u0026lt;\u0026thinsp;0.1 mV) (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ea and \u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ec).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003e3.6 Field trials of synthetic compounds and blends of tea volatiles.\u003c/h2\u003e \u003cp\u003eIn October 2022 and 2023, the field trials were conducted in two different tea gardens in Jinan and Tai\u0026rsquo;an, respectively. The result showed that the synergistic bait trap (attractant\u0026thinsp;+\u0026thinsp;yellow sticky trap) had the best attraction effect. The trapping effect of synergistic bait traps containing blend 3 at 100 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL, blend 4 at 100 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL, and blend 6 at 100 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL were significantly better than those of the other three treatments (CK, single attractant, and yellow sticky trap) in 2022 (Figs.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003e and \u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e9\u003c/span\u003e). The trapping effect of synergistic bait traps containing (\u003cem\u003eE\u003c/em\u003e)-2-hexenal at 1 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL, (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene at 1 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL, blend 1 at 1 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL, blend 3 at 10 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL, blend 4 at 100 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL, blend 5 at 100 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL, and blend 6 at 100 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL were significantly better than those of the other three treatments in 2023 (Figs.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003e and \u003cspan refid=\"Fig10\" class=\"InternalRef\"\u003e10\u003c/span\u003e). The lure longevity time of two attractants, blend 4 [the mixture of linalool and (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene] and blend 6 [the mixture of (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol and linalool] at a concentration of 100 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL, were assessed in 2022 and 2023. From 3 to 14 days after application in 2022 and from 7 to 23 days after application in 2023, compared to using yellow sticky traps alone, the number of adult \u003cem\u003eA. spiniferus\u003c/em\u003e was significantly higher in yellow sticky traps containing two types of attractants, blend 4 and blend 6 (Fig.\u0026nbsp;\u003cspan refid=\"Fig11\" class=\"InternalRef\"\u003e11\u003c/span\u003e). From 14 to 23 days after application in 2022 and from 23 to 30 days after application in 2023, there was no significant difference in the number of adults captured by each of the two treatments compared to the yellow sticky trap alone (Fig.\u0026nbsp;\u003cspan refid=\"Fig11\" class=\"InternalRef\"\u003e11\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"4. DISCUSSION","content":"\u003cp\u003eIt is well known that the type or variety of host plants affects the growth, development, and reproduction of herbivorous insects (Golizadeh et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Sarkar et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Therefore, the selection of optimal host plants is an important factor in the life of herbivorous insects. In this study, \u003cem\u003eA. spiniferus\u003c/em\u003e adults were more inclined to choose \u0026lsquo;Huangjinya\u0026rsquo; and \u0026lsquo;Fuding white tea\u0026rsquo; compared with \u0026lsquo;Baiye No.1\u0026rsquo; and \u0026lsquo;Longjing 43\u0026rsquo;, and laid more eggs, in the same environment of a tea plantation (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). \u0026lsquo;Huangjinya\u0026rsquo; is a novel chlorophyll-deficient albino cultivar that is susceptible to common pests of tea plants (e.g., \u003cem\u003eE. onukii\u003c/em\u003e and \u003cem\u003eToxoptera aurantii\u003c/em\u003e) (Tian et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; He et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). \u0026lsquo;Fuding white tea\u0026rsquo; is more suitable for herbivorous insects (e.g., \u003cem\u003eDendrothrips minowai\u003c/em\u003e, \u003cem\u003eT. aurantii\u003c/em\u003e, and \u003cem\u003eE. onukii\u003c/em\u003e) due to its high free amino acid content (Lu et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Although the nutritional level of plants often affects their suitability and resistance to herbivorous insects, insects search for and locate distant host plants using their sense of smell and vision in the early stages of pest infestation (Bruce et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2005\u003c/span\u003e, 2011). External morphological features of tea plants may be an important factor in the attraction of adult \u003cem\u003eA. spiniferus\u003c/em\u003e, such as the unique yellowish-green leaves of \u0026lsquo;Huangjinya\u0026rsquo;, which may be more attractive to insects with a preference for yellow (Li et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). However, the results of indoor Y-shaped olfactometer experiments showed that adult \u003cem\u003eA. spiniferus\u003c/em\u003e were still strongly attracted by \u0026lsquo;Huangjinya\u0026rsquo; and \u0026lsquo;Fuding white tea\u0026rsquo; even without visual cues. Therefore, olfaction plays a crucial role in the host-finding process of adult \u003cem\u003eA. spiniferus\u003c/em\u003e. When choosing between healthy plants and whitefly-infested plants of \u0026lsquo;Longjing 43\u0026rsquo;, 62.5 percent of adults did not make a choice (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea). Odor may be an major factor contributing to the low population density on \u0026lsquo;Longjing 43\u0026rsquo;.\u003c/p\u003e \u003cp\u003ePlants release varying amounts of volatile compounds from their leaves to regulate the behavior of various community members around them (Baldwin et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Soler et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). The comparison of the headspace volatiles of tea shoots from four cultivars showed that \u0026lsquo;Longjing 43\u0026rsquo; and \u0026lsquo;Baiye No. 1\u0026rsquo; released more attractive compounds, hexanol and (\u003cem\u003eE\u003c/em\u003e)-2-hexenal, compared with the other two cultivars. This is consistent with previously reported research; hexanol is a positive signal for attracting \u003cem\u003eNysomyia neivai\u003c/em\u003e (Machado et al. 2002); (\u003cem\u003eE\u003c/em\u003e)-2-hexenal has exhibited an attractive effect on herbivorous insects (e.g., \u003cem\u003ePlutella xylostella\u003c/em\u003e and \u003cem\u003eBemisia tabaci\u003c/em\u003e) (Li et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Yang et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Onnink et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), which can be used to lure \u003cem\u003eE. onukii\u003c/em\u003e into pheromone-baited traps (Mu et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). It is worth noting that hexanol and (\u003cem\u003eE\u003c/em\u003e)-2-hexenal did not elicit an obvious EAG response in adult \u003cem\u003eA. spiniferus\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ea and \u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ec). Therefore, the antennae may not serve as the primary sensory organ for their detection. Chemoreceptors are present in body hairs, appendages, mouthparts, and wings of insects (S\u0026aacute;nchez-Gracia et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). Odorant-binding proteins (OBPs) and chemosensory proteins (CSPs), which are involved in the binding and transport of odor molecules, are also expressed in tissues outside the antennae (Yi et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Lizana et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Tissue-specific transcriptome analysis revealed that CSPs are widely expressed in both the head (including the antennae) and body tissues of adult orange spiny whiteflies (Gao et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eLinalool have been only detected in \u0026lsquo;Fuding white tea\u0026rsquo;, and laboratory tests have confirmed that at a concentration of 100 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL, it can elicit obvious EAG responses and show attractive effects (Figs.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e and \u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ee). Natural linalool usually has two isomers: the levorotatory isomer (S)-(+)-linalool and the dextrorotatory isomer (R)-(\u0026minus;)-linalool (Aprotosoaie et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). (S)-(+)-linalool is mainly attracting pollinators, whereas (R)-(\u0026minus;)-linalool is mainly enhances resistance to herbivores and pathogens (Zhang et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Unfortunately, we have not been able to identify which isomer of linalool was released from \u0026lsquo;Fuding white tea\u0026rsquo; by GC-MS. Further work is needed to demonstrate the conformation as well as the specific roles of the linalool released from the non-flowering \u0026lsquo;Fuding white tea\u0026rsquo;. In this study, only trace amounts [34.25\u0026thinsp;\u0026plusmn;\u0026thinsp;2.60 (ng plant\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e h\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)] of (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene were detected in \u0026lsquo;Huangjinya\u0026rsquo;, and it showed significant attractive effects at the concentrations of 1, 10, and 100 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL. (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene is a common compound of herbivore-induced plant volatiles from many plants such as tea, soybean, pear, apple, and poplar, and it is a positive signal for attracting natural enemies (e.g., \u003cem\u003eAphidius ervi\u003c/em\u003e, \u003cem\u003eColeomegilla maculate\u003c/em\u003e, and \u003cem\u003eChrysoperla carnea\u003c/em\u003e) (Du et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Zhu et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Lin et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Consequently, we reasoned that the presence of (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene may have both positive and negative effects, as it attracts natural enemies while making the tea plant vulnerable to attacks from \u003cem\u003eA. spiniferus\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eExcept for the susceptible \u0026lsquo;Huangjinya\u0026rsquo;, (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol was detected in the headspace volatiles of the other three cultivars (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Laboratory tests confirmed that (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol had a repellent effect at concentrations of 1, 10, and 100 \u003cem\u003e\u0026micro;\u003c/em\u003eg/\u003cem\u003e\u0026micro;\u003c/em\u003eL (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). The accumulating evidence on the C6-volatile (\u003cem\u003eZ\u003c/em\u003e)‐3‐hexenol supports its crucial role in mediating indirect defense responses in plants. Tea plants release (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol as a defense response against herbivores (Dong et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Sun et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Cai et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). This compound contributes to the unique grassy aroma of green tea (Jing et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). We hypothesized that healthy tea plants emit trace amounts of (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol as an insect repellent; conversely, undamaged \u0026lsquo;Huangjinya\u0026rsquo; do not release the aforementioned repellent and are therefore favored by adult \u003cem\u003eA. spiniferus\u003c/em\u003e. Another compound with a repellent effect verified by behavioral experiments is nonanal, but it was not detected in the repellent variety \u0026lsquo;Longjing 43\u0026rsquo;. This could be attributed to the fact that most insects use mixtures of volatiles instead of single components for host identification (Delorme and Payne \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Bruce et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). For example, female \u003cem\u003eAulacphora foveicollis\u003c/em\u003e responded positively to the volatile mixture of \u003cem\u003eSolena amplexicaulis\u003c/em\u003e fruits due to the synergistic effect of nonanal and (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eZ\u003c/em\u003e)-2,6-nonadienals (Karmakar et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe application of plant-derived semiochemicals to control tea pests has been widely reported, such as the application of a mixture of dimethyl disulfide (DMDS) and 1,8-eucalyptin to repel \u003cem\u003eE. onukii\u003c/em\u003e in tea fields and the application of (\u003cem\u003eE\u003c/em\u003e)-2-hexenal, (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol, (\u003cem\u003eZ\u003c/em\u003e)-hexenyl acetate, (\u003cem\u003eE\u003c/em\u003e)-ocimene, linalool, and geraniol as \u003cem\u003eE. onukii\u003c/em\u003e attractants (Chen et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Cai et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Field trials had shown a significant improvement in trapping efficiency when combined with yellow sticky traps. Yellow sticky traps are commonly used for whitefly monitoring and control (Moreau and Isman \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Extensive control of adult \u003cem\u003eA. spiniferus\u003c/em\u003e using yellow sticky traps has been reported in Japan (Uesugi and Sato \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). The results of field trials showed that, the addition of attractants greatly increased the attractiveness of the yellow sticky traps.\u003c/p\u003e \u003cp\u003eIt is worth noting that more attractants had a significant luring effect in the tea plantation with high-density of adult \u003cem\u003eA. spiniferus\u003c/em\u003e. Previous studies have shown that when populations were low, enlarging the entry area can increased \u003cem\u003eDrosophila suzukii\u003c/em\u003e captures and improved selectivity for \u003cem\u003eD. suzukii\u003c/em\u003e (Renkema et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). When areas affected by extremely dense bark beetle populations, the use of anti-attractants did not prove effective (Jakuš et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Suggesting that population density must be taken into account in the testing of attractant effects.\u003c/p\u003e \u003cp\u003eIn the lure longevity time experiment in October 2023, the number of adult \u003cem\u003eA. spiniferus\u003c/em\u003e trapped by yellow sticky traps gradually declined, it is consistent with the results of population dynamics (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea). The field trial was conducted when the third peak of \u003cem\u003eA. spiniferus\u003c/em\u003e adult occurrence, subsequently, the number of adult \u003cem\u003eA. spiniferus\u003c/em\u003e gradually decreased. This is an indication that the lure longevity time may be affected by the season; the long-term monitoring tests will be required (Wang et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe field trials revealed two anomalies: (i) (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol and nonanal, which were repellent in the laboratory test, did not show repellent effects in the field experiments, but instead showed a positive response; (ii) in Tai\u0026rsquo;an (2023) but not Jinan (2022), the lowest dose of (\u003cem\u003eE\u003c/em\u003e)-2-hexenal, (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene and blend 1 increased the catches significantly, it was not consistent with laboratory tests. Background odors in the field can interfere with the attractiveness of plant volatile attractants to herbivorous insects (Xu et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Similarities were found in a screening study for \u003cem\u003eD. suzukii\u003c/em\u003e attractant volatiles, when the co-attractiveness of some of the individual compounds were tested under different background odor environments, they appears to change, suggesting that background odor can influence detection of potential attractants (Cha et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). In the other study, five traps were put at different sites in a cornfield that had record of emergence of \u003cem\u003eAnomala corpulenta\u003c/em\u003e, only the baits at site 1\u0026ndash;4 effectively lured and trapped (Qu et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Suggesting that for more accurate conclusions, field trials need to be carried out at different locations.\u003c/p\u003e"},{"header":"5. CONCLUSION","content":"\u003cp\u003eOur study demonstrates that plant volatiles mediate host selection of adult \u003cem\u003eA. spiniferus\u003c/em\u003e among tea cultivars. Blend 4 [the mixture of linalool and (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene] and blend 6 [the mixture of linalool and (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol], two combinations of attractants based on tea shoot volatiles, were found to be highly effective and long-lasting in trapping adult \u003cem\u003eA. spiniferus\u003c/em\u003e in two years. They have the potential to be developed as commercial attractants for the prevention and monitoring of adult \u003cem\u003eA. spiniferus\u003c/em\u003e.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAuthor\u0026rsquo;s contribution\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eYYX, ZZC and ZFJ conceived the idea and designed the research; ZFJ, LYH, YNB and KS conducted the experiments and analyzed the data; ZFJ wrote the manuscript; ZQL, YYX, ZZC and DDL edited and reviewed the manuscript. All authors commented and approved the manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgments\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe sincerely thank all the staff and students of the Laboratory of Insect Ecology and Physiology (Shandong Agricultural University), as well as Shandong Qianrun Ecological Agriculture Development Co., Ltd., for providing us with the experimental fields. We would like to thank DBMediting for professional english language editing services.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis research was funded by the National Key Research\u0026nbsp;\u0026amp;\u0026nbsp;Development Program of China (2023YFD1700405) and the Modern Tea Industry Technology System of Shandong Province (SDAIT-19-04).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors acknowledge no conflict of interest concerning the publication of this research article.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData availability\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe datasets generated and analysed during the current study are available from the corresponding author on reasonable request.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAprotosoaie AC, Hancianu M, Costache II, Miron A (2014) Linalool: A review on a key odorant molecule with valuable biological properties. Flavour Frag J 29:193\u0026minus;219. https://doi.org/10.1002/ffj.3197\u003c/li\u003e\n\u003cli\u003eBaldwin IT, Halitschke R, Paschold A, von Dahl CC, Preston CA (2006) Volatile signaling in plant\u0026minus;plant interactions:\u0026quot;Talking trees\u0026quot; in the genomics era. Science 311:812\u0026minus;815. https://science.org/doi/10.1126/science.1118446\u003c/li\u003e\n\u003cli\u003eBirkett MA, Bruce TJA, Martin JL, Smart LE, Oakley J, Wadhams LJ (2004) Responses of female orange wheat blossom midge, \u003cem\u003eSitodiplosis mosellana\u003c/em\u003e, to wheat panicle volatiles. J Chem Ecol 30:1319\u0026minus;1328. https://doi.org/10.1023/b:joec.0000037742.05022.9f\u003c/li\u003e\n\u003cli\u003eBragard C, Dehnen‐Schmutz K, Di Serio F et al (2018). Pest categorisation of \u003cem\u003eAleurocanthus spp.\u003c/em\u003e. EFSA J 16:5436. https://doi.org/10.2903/j.efsa.2018.5436\u003c/li\u003e\n\u003cli\u003eBruce TJA, Pickett JA (2011) Perception of plant volatile blends by herbivoro-us insects－Finding the right mix. Phytochemistry 72:1605\u0026minus;1611. https://doi.org/10.1016/j.phytochem.2011.04.011\u003c/li\u003e\n\u003cli\u003eBruce TJA, Wadhams LJ, Woodcock CM (2005) Insect host location: Avolatile situation. Trends Plant Sci 10:269\u0026minus;274. https://doi.org/10.1016/j.tplants.2005.04.003\u003c/li\u003e\n\u003cli\u003eByrne DN and Bellows TS (1991) Whitefly biology. Annu Rev Entomol 36:431\u0026ndash;457. https://doi.org/10.1146/annurev.en.36.010191.002243\u003c/li\u003e\n\u003cli\u003eCai XM, Sun XL, Dong WX, Wang GC, Chen ZM (2014) Herbivore species, infestation time, and herbivore density affect induced volatiles in tea plants. Chemoecology 24:1\u0026minus;14. https://doi.org/10.1007/s00049-013-0141-2\u003c/li\u003e\n\u003cli\u003eCai XM, Xu XX, Bian L, Luo ZX, Xin ZJ, Chen ZM (2015) Attractiveness of host volatiles combined with background visual cues to the tea leafhopper, \u003cem\u003eEmpoasca vitis\u003c/em\u003e. Entomol Exp Appl 157:291\u0026minus;299. https://doi.org/10.1111/eea.12364\u003c/li\u003e\n\u003cli\u003eCai XM, Luo ZX, Meng ZN et al (2020) Primary screening and application of repellent plant volatiles to control tea leafhopper, \u003cem\u003eEmpoasca onukii\u003c/em\u003e Matsuda. Pest Manag Sci 76:1304\u0026minus;1312. https://doi.org/10.1002/ps.5641\u003c/li\u003e\n\u003cli\u003eCha DH, Loeb GM, Linn CEJ, Hesler SP, Landolt PJ (2018) A multiple-choice bioassay approach for rapid screening of key attractant volatiles. Environ Entomol 47:946\u0026minus;950. https://doi.org/10.1093/ee/nvy054\u003c/li\u003e\n\u003cli\u003eChen K, Huang MX, Shi QC, Xie X, Jin LH, Xu WM, Li XY (2019) Screening of a potential leafhopper attractants and their applications in tea plantations. J Environ Sci Heal B 54:858\u0026minus;865.https://doi.org/10.1080/03601234.2019.1633856\u003c/li\u003e\n\u003cli\u003eDelorme JD, Payne TL (1990) Antennal olfactory responses of black turpentine beetle, \u003cem\u003eDendroctonus terebrans\u003c/em\u003e (Olivier), to bark beetle pheromones and host terpenes. J Chem Ecol 16:1321\u0026minus;1329. https://doi.org/10.1007/bf01021029\u003c/li\u003e\n\u003cli\u003eDong F, Yang Z, Baldermann S, Sato Y, Asai T, Watanabe N (2011) Herbivore-induced volatiles from tea (\u003cem\u003eCamellia sinensis\u003c/em\u003e) plants and their involveme-nt in intraplant communication and changes in endogenous nonvolatile me-tabolites. J Agric Food Chem 59:13131\u0026minus;13135. https://doi.org/10.1021/jf203396a\u003c/li\u003e\n\u003cli\u003eDu YJ, Poppy GM, Powell W, Pickett JA, Wadhams LJ, Woodcock CM (1998)Identification of semiochemicals released during aphid feeding that attract parasitoid \u003cem\u003eAphidius ervi\u003c/em\u003e. J Chem Ecol 24:1355\u0026minus;1368. https://doi.org/10.1023/a:1021278816970\u003c/li\u003e\n\u003cli\u003eGao YQ, Chen ZZ, Liu MY et al (2022) Characterization of antennal chemos-ensilla and associated chemosensory genes in the orange spiny whitefly, \u003cem\u003eA-leurocanthus spiniferus\u003c/em\u003e (Quaintanca). Front Physiol 13:847895. https://doi.org/10.3389/fphys.2022.847895\u003c/li\u003e\n\u003cli\u003eGolizadeh A, Abedi Z, Borzoui E, Golikhajeh N, Jafary M (2016) Susceptibilit-y of five sugar beet cultivars to the black bean aphid, \u003cem\u003eAphis fabae\u003c/em\u003e scopoli(Hemiptera: Aphididae). Neotrop Entomol 45:427\u0026ndash;432. https://doi.org/10.1007/s13744-016-0383-0\u003c/li\u003e\n\u003cli\u003eHan BY and Chen ZM (2002) Behavioral and electrophysiological responses ofnatural enemies to synomones from tea shoots and kairomones from tea a-phids, \u003cem\u003eToxoptera aurantii\u003c/em\u003e. J Chem Ecol 28:2203\u0026minus;2219. https://doi.org/10.1023/a:1021045231501\u003c/li\u003e\n\u003cli\u003eHazarika LK, Puzari KC, Wahab S (2001) Biological control of tea pests. In: Upadhyay RK, Mukerji KG and Chamola BP (ed) Biocontrol potential and its exploitation in sustainable agriculture. Springer, Boston, pp 159\u0026ndash;180.\u003c/li\u003e\n\u003cli\u003eHe Y, Lu C, Jiang W, Chen WL, Fan JJ, Niu SZ, Zhao DG (2023) Adaptability of \u003cem\u003eToxoptera aurantii\u003c/em\u003e (Hemiptera: Aphididae) to different tea cultivars. Agriculture 13:2039. https://doi.org/10.3390/agriculture13102039\u003c/li\u003e\n\u003cli\u003eHuang QT, Han XQ, Zhang GJ, Zhu-Salzman K, Cheng WN (2022) Plant vol-atiles mediate host selection of \u003cem\u003eSitodiplosis mosellana\u003c/em\u003e (Diptera: Cecidomyi-idae) among wheat varieties. J Agric Food Chem 70:10466\u0026minus;10475. https://doi.org/10.1021/acs.jafc.2c03244\u003c/li\u003e\n\u003cli\u003eJaku\u0026scaron; R, Modlinger R, Ka\u0026scaron;par J et al (2022) Testing the efficiency of the pus-h-and-pull strategy during severe \u003cem\u003eIps typographus\u003c/em\u003e outbreak and extreme dr-ought in Norway spruce stands. Forests 13:2175. https://doi.org/10.3390/f13122175\u003c/li\u003e\n\u003cli\u003eJing TT, Zhang N., Gao T et al (2019) Glucosylation of (Z)‐3‐hexenol informs intraspecies interactions in plants: A case study in \u003cem\u003eCamellia sinensis\u003c/em\u003e. Plant, Cell \u0026amp; Environment 42:1352\u0026ndash;1367. https://doi.org/10.1111/pce.13479 \u003c/li\u003e\n\u003cli\u003eKarmakar A, Mitra P, Koner A, Das S, Barik A (2020) Fruit volatiles of creeping cucumber (\u003cem\u003eSolena amplexicaulis\u003c/em\u003e) attract a generalist insect herbivore. J Chem Ecol 46:275\u0026minus;287. https://doi.org/10.1007/s10886-020-01154-w\u003c/li\u003e\n\u003cli\u003eLi CF, Ma JQ, Huang DJ, Ma CL, Jin JQ, Yao MZ, Chen L (2018) Comprehensive dissection of metabolic changes in albino and green tea cultivars. J Agric Food Chem 66:2040\u0026minus;2048. https://doi.org/10.1021/acs.jafc.7b05623\u003c/li\u003e\n\u003cli\u003eLi YF, Zhong ST, Qin YC, Zhang SQ, Gao ZL, Dang ZH, Pan WL (2014) Identification of plant chemicals attracting and repelling whiteflies. Arthropod-Plant Inte 8:183\u0026minus;190. https://doi.org/10.1007/s11829-014-9302-7\u003c/li\u003e\n\u003cli\u003eLiang LY, Liu LF, Yu XP, Han BY (2012) Evaluation of the resistance of different tea cultivars to tea aphids by EPG technique. J Integr Agr 11:2028\u0026minus;2034. https://doi.org/10.1016/S2095-3119(12)60460-2\u003c/li\u003e\n\u003cli\u003eLin J, Wang D, Chen X et al (2017) An (E,E)-\u0026alpha;-farnesene synthase gene of s-oybean has a role in defence against nematodes and is involved in synthe-sizing insect-induced volatiles. Plant Biotechnol J 15:510\u0026minus;519. https://doi.org/10.1111/pbi.12649\u003c/li\u003e\n\u003cli\u003eLizana P, Mutis A, Quiroz A, Venthur H (2022). Insights into chemosensory p-roteins from non-model insects: Advances and perspectives in the context of pest management. Front Physiol 13:924750. https://doi.org/10.3389/fphys.2022.924750\u003c/li\u003e\n\u003cli\u003eLu C, Shen N, Jiang W, et al (2023) Different tea germplasms distinctly influence the adaptability of \u003cem\u003eToxoptera aurantii\u003c/em\u003e (Hemiptera: Aphididae). Insects 14:695. https://doi.org/10.3390/insects14080695\u003c/li\u003e\n\u003cli\u003eMachado VE, Silva FBD, Goulart TM, Pinto MC (2022) Behavioral responses of sand fly \u003cem\u003eNyssomyia neivai\u003c/em\u003e (Psychodidae: Phlebotominae) to 1-hexanol and light. Acta Trop 236:106680. https://doi.org/10.1016/j.actatropica.2022.106680\u003c/li\u003e\n\u003cli\u003eMagsi FH, Luo ZX, Zhao YJ, Li ZQ, Cai XM, Bian L, Chen ZM (2021) Electrophysiological and behavioral responses of \u003cem\u003eDasychira baibarana\u003c/em\u003e (Lepidoptera: Lymantriidae) to tea plant volatiles. Environ Entomol 50:589\u0026minus;598. https://doi.org/10.1093/ee/nvab016\u003c/li\u003e\n\u003cli\u003eMoreau TL and Isman MB (2011) Trapping whiteflies? A comparison of greenhouse whitefly (\u003cem\u003eTrialeurodes vaporariorum\u003c/em\u003e) responses to trap crops and yellow sticky traps. Pest Manag Sci 67:408\u0026minus;413. https://doi.org/10.1002/ps.2078\u003c/li\u003e\n\u003cli\u003eMu D, Cui L, Ge J et al (2012) Behavioral responses for evaluating the attractiveness of specific tea shoot volatiles to the tea green leafhopper, \u003cem\u003eEmpoaca vitis\u003c/em\u003e. Insect Sci 19:229\u0026minus;238. https://doi.org/10.1111/j.1744-7917.2011.01476.x\u003c/li\u003e\n\u003cli\u003eOnnink CM, Williams AA, Williams JS (2017) Electrophysiological and behavioral evidence for (E)-2-hexenal as a female-attracting pheromone produced by disturbed \u003cem\u003eMegacopta cribraria\u003c/em\u003e (Hemiptera: Plataspidae). J Entomol Sci 52:39\u0026minus;51. https://doi.org/10.18474/JES16-18.1\u003c/li\u003e\n\u003cli\u003eQu CJ, Shi JT, Wang CZ, Guo H, Jiang XJ (2024) Identification of an adult attractant for \u003cem\u003eAnomala corpulenta\u003c/em\u003e by the reverse chemical ecology approach. J Pest Sci. https://doi.org/10.1007/s10340-023-01740-3\u003c/li\u003e\n\u003cli\u003eRanger CM, Winter REK, Backus EA, Rottinghaus GE, Ellersieck MR, Johnson DW (2005) Discrimination by the potato leafhopper (Hemiptera: Cicadellidae) of host volatiles from resistant and susceptible alfalfa, \u003cem\u003eMedicago sativa\u003c/em\u003e L. Environ Entomol 34:271\u0026minus;280. https://doi.org/10.1603/0046-225X-34.2.271\u003c/li\u003e\n\u003cli\u003eRenkema JM, Buitenhuis R, Hallett RH (2014) Optimizing trap design and trapping protocols for \u003cem\u003eDrosophila suzukii\u003c/em\u003e (Diptera: Drosophilidae). J Econ Entomol 107:2107\u0026minus;2118. https://doi.org/10.1603/ec14254\u003c/li\u003e\n\u003cli\u003eS\u0026aacute;nchez-Gracia A, Vieira FG, Rozas J (2009) Molecular evolution of the majorchemosensory gene families in insects. Heredity 103: 208\u0026minus;216. https://doi.org/10.1038/hdy.2009.55\u003c/li\u003e\n\u003cli\u003eSarkar S, Babu A, Chakraborty K, Deka B, Kashyap B, Adhikary B (2021) Study on varietal preference and nutritional indices of tea looper \u003cem\u003eHyposidra talaca\u003c/em\u003e (Lepidoptera: Geometridae) on ten selected tea varieties. Int J Trop Insect Sc 41:3087\u0026minus;3098. https://doi.org/10.1007/s42690-021-00502-x\u003c/li\u003e\n\u003cli\u003eSoler R, Harvey JA, Kamp AFD et al (2007) Root herbivores in fluence the behaviour of an aboveground parasitoid through changes in plant-volatile signals. Oikos 116:367\u0026minus;376. https://doi.org/10.1111/j.0030-1299.2007.15501.x\u003c/li\u003e\n\u003cli\u003eSong CF, Ma L, Zhao JY, Xue ZS, Yan XZ, Hao C (2022) Electrophysiological and behavioral responses of \u003cem\u003ePlutella xylostella\u003c/em\u003e (Lepidoptera:Plutellidae) to volatiles from a non-host plant, geranium, \u003cem\u003ePelargonium hortorum\u003c/em\u003e (Geraniaceae). J Agric Food Chem 70:5982\u0026minus;5992. https://doi.org/10.1021/acs.jafc.1c08165\u003c/li\u003e\n\u003cli\u003eSun XL, Wang GC, Cai XM, Jin S, Gao Y, Chen ZM (2010) The tea weevil, \u003cem\u003eMyllocerinus aurolineatus\u003c/em\u003e, is attracted to volatiles induced by conspecifics. J Chem Ecol 36:388\u0026minus;395. https://doi.org/10.1007/s10886-010-9771-9\u003c/li\u003e\n\u003cli\u003eSun XL, Wang GC, Gao Y, Zhang XZ, Xin ZJ, Chen ZM (2014) Volatiles emitted from tea plants infested by \u003cem\u003eEctropis obliqua\u003c/em\u003e larvae are attractive to conspecific moths. J Chem Ecol 40:1080\u0026minus;1089. https://doi.org/10.1007/s10886-014-0502-5\u003c/li\u003e\n\u003cli\u003eTang XT, Tao HH, Du YZ (2015) Microsatellite-based analysis of the genetic structure and diversity of \u003cem\u003eAleurocanthus spiniferus\u003c/em\u003e (Hemiptera: aleyrodidae)from tea plants in China. Gene 560:107\u0026ndash;113. https://doi.org/10.1016/j.gene.2015.01.050\u003c/li\u003e\n\u003cli\u003eTian Y, Zhao Y, Zhang L, Mu W, Zhang ZQ (2018) Morphological, physiolog-ical, and biochemical responses of two tea cultivars to \u003cem\u003eEmpoasca onukii\u003c/em\u003e (Hemiptera: Cicadellidae) infestation. J Econ Entomol 111:899\u0026minus;908. https://doi.org/10.1093/jee/toy011\u003c/li\u003e\n\u003cli\u003eTian YY, Chen ZJ, Huang XQ, Zhang LX, Zhang ZQ (2020) Evaluation of botanicals for management of piercing\u0026ndash;sucking pests and the effect on beneficial arthropod populations in tea trees \u003cem\u003eCamellia sinensis\u003c/em\u003e (L.) O. Kuntze (Theaceae). J Insect Sci 20: 27. https://doi.org/10.1093/jisesa/ieaa101\u003c/li\u003e\n\u003cli\u003eUesugi R and Sato Y (2013) Counting adults on a yellow sticky trap is useful for monitoring the tea spiny whitefly, \u003cem\u003eAleurocanthus camelliae\u003c/em\u003e (Hemiptera: Aleyrodidae), at the early invasion stage. JPN J Appl Entomol Z 57:35\u0026minus;41. https://doi.org/10.1303/jjaez.2013.35\u003c/li\u003e\n\u003cli\u003eUesugi R, Sato Y, Han BY, Huang ZD, Yara K, Furuhashi K (2016) Molecular evidence for multiple phylogenetic groups within two species of invasive spiny whiteflies and their parasitoid wasp. Bull Entomol Res 106:328\u0026ndash;340. https://doi.org/10.1017/S0007485315001030\u003c/li\u003e\n\u003cli\u003eWang HM, Bai PH, Zhang J, Zhang XM, Hui Q, Zheng HX, Zhang XH (2020) Attraction of bruchid beetles \u003cem\u003eCallosobruchus chinensis\u003c/em\u003e (L.)(Coleoptera: Bruchidae) to host plant volatiles. J Integr Agr 19:3035\u0026minus;3044. https://doi.org/10.1016/S2095-3119(20)63237-3\u003c/li\u003e\n\u003cli\u003eWang J, Wei J, Yi T, Li YY, Xu T, Chen L, Xu H (2023) A green leaf volat-ile, (Z)-3-hexenyl-acetate, mediates differential oviposition by \u003cem\u003eSpodoptera f-rugiperda \u003c/em\u003eon maize and rice. BMC Biol 21:140. https://doi.org/10.1186/s12915-023-01642-x\u003c/li\u003e\n\u003cli\u003eWang LY, Yang CX, Mei XD, Guo BB, Yang XL, Zhang T, Ning J (2024) Identification of sex pheromone in \u003cem\u003eMacdunnoughia crassisigna\u003c/em\u003e Warren (Lepidoptera: Noctuidae) and field optimization of the sex attractant. Pest Manage Sci 80:577\u0026minus;585. https://doi.org/10.1002/ps.7783\u003c/li\u003e\n\u003cli\u003eWang X, Zeng L, Liao Y, Li J, Tang J, Yang Z (2019) Formation of \u0026alpha;-farnes-ene in tea (\u003cem\u003eCamellia sinensis\u003c/em\u003e) leaves induced by herbivore-derived woundi-ng and its effect on neighboring tea plants. Int J Mol Sci 20:4151. https://doi.org/10.3390/ijms20174151\u003c/li\u003e\n\u003cli\u003eXiu CL, Xu B, Pan HS, Zhang W, Yang YZ, Lu YH (2019) Volatiles from \u003cem\u003eSophora japonica\u003c/em\u003e flowers attract \u003cem\u003eHarmonia axyridis\u003c/em\u003e adults (Coleoptera: Coccinellidae). J Integr Agr 18:873\u0026minus;883. https://doi.org/10.1016/S2095-3119(18)61927-6\u003c/li\u003e\n\u003cli\u003eXu XX, Cai XM, Bian L, Luo ZX, Li ZQ, Chen ZM (2017) Does backgroun-d odor in tea gardens mask attractants? Screening and application of attra-ctants for \u003cem\u003eEmpoasca onukii\u003c/em\u003e Matsuda. J Econ Entomol 110:2357\u0026minus;2363. https://doi.org/10.1093/jee/tox269\u003c/li\u003e\n\u003cli\u003eYang G, Zhang YN, Gurr GM, Vasseur L, You M (2016). Electroantennogram and behavioral responses of \u003cem\u003eCotesia plutellae\u003c/em\u003e to plant volatiles. Insect Sci 23:245\u0026minus;252. https://doi.org/10.1111/1744-7917.12308\u003c/li\u003e\n\u003cli\u003eYang TB, Song XH, Zhong Y, Wang B, Zhou CQ (2022) Field investigation‐ and dietary metabarcoding‐based screening of arthropods that prey on primary tea pests. Ecol Evol 12:e9060. https://doi.org/10.1002/ece3.9060\u003c/li\u003e\n\u003cli\u003eYi X, Zhang Y, Wang P, Qi JW, Hu MY, Zhong GH (2015) Ligands binding and molecular simulation: the potential investigation of a biosensor based on an insect odorant binding protein. Int J Biol Sci 11:75\u0026minus;87. https://www.ijbs.com/v11p0075.htm\u003c/li\u003e\n\u003cli\u003eYuan GG, Zhao LC, Du YW, Yu H, Shi XB, Chen WC,Chen G (2022) Repel-lence or attraction: secondary metabolites in pepper mediate attraction and defense against \u003cem\u003eSpodoptera litura\u003c/em\u003e. Pest Manage Sci 78:4859\u0026minus;4870. https://doi.org/10.1002/ps.7107\u003c/li\u003e\n\u003cli\u003eZhang L, Su QF, Wang LS, Lv MW, Hou YX, Li SS (2023) Linalool: A ubiquitous floral volatile mediating the communication between plants and insects. J Syst Evol 61:538\u0026minus;549. https://doi.org/10.1111/jse.12930\u003c/li\u003e\n\u003cli\u003eZhang ZQ, Sun XL, Xin ZJ, Luo ZX, Gao Y, Bian L, Chen ZM (2013) Ident-ification and field evaluation of non-host volatiles disturbing host location by the tea geometrid, \u003cem\u003eEctropis obliqua\u003c/em\u003e. J Chem Ecol 39:1284\u0026minus;1296. https://doi.org/10.1007/s10886-013-0344-6\u003c/li\u003e\n\u003cli\u003eZhang ZQ, Bian L, Sun XL, Luo ZX, Xin ZJ, Luo FJ, Chen ZM (2015) Ele-ctrophysiological and behavioural responses of the tea geometrid \u003cem\u003eEctropis obliqua\u003c/em\u003e (Lepidoptera: Geometridae) to volatiles from a non-host plant, ros-emary, \u003cem\u003eRosmarinus officinalis\u003c/em\u003e (Lamiaceae). Pest Manag Sci 71:96\u0026minus;104. https://doi.org/10.1002/ps.3771\u003c/li\u003e\n\u003cli\u003eZhao MY, Zhang N, Gao T et al (2020) Sesquiterpene glucosylation mediated by glucosyltransferase UGT91Q2 is involved in the modulation of cold str-ess tolerance in tea plants. New Phytol 226:362\u0026ndash;372. https://doi.org/10.1111/nph.16364\u003c/li\u003e\n\u003cli\u003eZhu JW, Cosse AA, Obrycki JJ, Boo KS, Baker TC (1999) Olfactory reactions of the twelve-spotted lady beetle, \u003cem\u003eColeomegilla maculata\u003c/em\u003e and the green lacewing, \u003cem\u003eChrysoperla carnea\u003c/em\u003e to semiochemicals released from their prey and host plant: electroantennogram and behavioral responses. J Chem Ecol 25:1163\u0026minus;1177. https://doi.org/10.1023/a:1020846212465\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Aleurocanthus spiniferus, tea cultivars, volatiles, behavioral assays, EAG, field trials","lastPublishedDoi":"10.21203/rs.3.rs-4333561/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4333561/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003e\u003cem\u003eAleurocanthus spiniferus \u003c/em\u003eis a major pest of tea trees; both nymphs and adults suck plant sap and induce tea sooty mould. Pest occurrence varies significantly among different tea cultivars, even within the same plantation environment. It was speculated that adult \u003cem\u003eA. spiniferus\u003c/em\u003e may be differentially attracted by different cultivars. This study aims to identify the bioactive components in tea volatiles that mediate host selection of \u003cem\u003eA. spiniferus\u003c/em\u003e and to utilize them for effective field trapping. Through field investigations and Y-tube olfactometer tests, we identifiedtwo highly preferred tea cultivars (‘Huangjinya’ and ‘Fuding white tea’) and two cultivars (‘Baiye No. 1’ and ‘Longjing 43’) that were not. Behavioral assays and gas chromatography-mass spectrometry (GC-MS) analysis identified four attractive compounds [hexanol, (\u003cem\u003eE\u003c/em\u003e)-2-hexenal, linalool, and (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene] and two repellent compounds [nonanal and (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol] from tea volatiles of the four. Among these compounds, four [nonanol, (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol, linalool and (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene] were able to elicit obvious electrophysiological (EAG) response at 100 \u003cem\u003eμ\u003c/em\u003eg/\u003cem\u003eμ\u003c/em\u003eL. In field trials, the synergistic bait trap equipped with two types of attractants, blend 4 [the mixture of linalool and (\u003cem\u003eE\u003c/em\u003e,\u003cem\u003eE\u003c/em\u003e)-\u003cem\u003eα\u003c/em\u003e-farnesene] at 100 \u003cem\u003eμ\u003c/em\u003eg/\u003cem\u003eμ\u003c/em\u003eL and blend 6 [the mixture of linalool and (\u003cem\u003eZ\u003c/em\u003e)-3-hexenol] at 100 \u003cem\u003eμ\u003c/em\u003eg/\u003cem\u003eμ\u003c/em\u003eL, had a significant attractant effect with a long effective period. Two synthetic mixtures of tea volatiles, blend 4 and blend 6, had the potential to be developed as commercial plant-based attractants for adult \u003cem\u003eA. spiniferus\u003c/em\u003e. This study contributes to the development of sustainable, environmentally friendly management strategies for a pest that is difficult to prevent and control.\u003c/p\u003e","manuscriptTitle":"Field application of tea volatiles mediating host selection of Aleurocanthus spiniferus","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-05-03 18:18:53","doi":"10.21203/rs.3.rs-4333561/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"e8070e20-2964-4eb1-9510-cd69500c61f4","owner":[],"postedDate":"May 3rd, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2024-07-11T12:17:48+00:00","versionOfRecord":[],"versionCreatedAt":"2024-05-03 18:18:53","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-4333561","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4333561","identity":"rs-4333561","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}