Season-Dependent Synergism between the Male- Attractive Plant Volatile Benzaldehyde and the Sex Pheromone of the Oriental Fruit Moth, Grapholita molesta

Season-Dependent Synergism between the Male- Attractive Plant Volatile Benzaldehyde and the Sex Pheromone of the Oriental Fruit Moth, Grapholita molesta | 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 Article Season-Dependent Synergism between the Male- Attractive Plant Volatile Benzaldehyde and the Sex Pheromone of the Oriental Fruit Moth, Grapholita molesta Ajay P. Giri, Brent D. Short, Jaime C. Pinero This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6637201/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 15 Oct, 2025 Read the published version in Scientific Reports → Version 1 posted 10 You are reading this latest preprint version Abstract Herbivorous insects possess highly developed olfactory systems that enable them to detect, process, and respond to volatile cues emitted by host and nonhost plants. This study evaluated the field response of male and female Oriental fruit moths. Grapholita molesta (Lepidoptera: Tortricidae) to the aromatic plant volatile benzaldehyde (BEN). We evaluated BEN alone, in combination with the G. molesta sex pheromone, or with a four-component kairomonal lure (PHEROCON MEGALURE CM 4K DUAL®). Our findings demonstrate that (1) BEN alone is a male G. molesta attractant with efficacy comparable to that of Megalure or the sex pheromone; (2) the addition of BEN at low or medium doses to either Megalure or pheromone lures significantly increases the number of male captures; and (3) BEN interacts with the G. molesta pheromone in a season-dependent manner, exhibiting synergistic effects during the middle-season and additive effects in the early and late seasons. These results highlight the potential of BEN to enhance pheromone-based lures without the need for additional host volatiles, providing a valuable tool for optimizing semiochemical-based monitoring and control strategies tailored to seasonal population dynamics. Biological sciences/Zoology/Entomology Biological sciences/Chemical biology/Chemical ecology Behavior monitoring semiochemical IPM synergism sensory ecology Figures Figure 1 Figure 2 Figure 3 Introduction Insects rely on highly specialized olfactory systems to detect, process, and respond to chemical cues in their environment [ 1 ]. Within Lepidoptera, semiochemicals, including host plant volatiles and sex pheromones, play pivotal roles in mediating insect behavior. This information is crucial to better understand the patterns of host plant use and for the development of more efficient monitoring and/or control tactics [ 2 – 4 ]. A major advancement in pest control is the implementation of mating disruption techniques, wherein synthetic sex pheromones are deployed at high densities to interfere with mate finding, thereby reducing reproductive success [ 5 – 9 ]. Additionally, semiochemical-based attractants that target both male and female moths are being explored for their potential in integrated pest management (IPM) strategies, including attract-and-kill techniques [ 10 ]. Because the success of these approaches relies on a comprehensive understanding of insect olfactory responses and the complex interactions between semiochemicals [ 11 ], unremitting research is essential to refine existing systems and develop more effective strategies for pest management. Research suggests that complex blends of host-plant-derived volatiles are often required to elicit significant attraction in tortricid moths [ 12 , 13 ]. This phenomenon is particularly relevant for the Oriental fruit moth (OFM), Grapholita molesta (Busck), a significant pest of stone and pome fruits worldwide [ 14 – 16 ]. Notably, a five-component blend of three green leaf volatiles and two aromatic compounds (benzaldehyde and benzonitrile) has been shown to be as attractive to mated female G. molesta [ 15 ] as the full suite of natural peach shoot volatiles, which contains more than 20 compounds [ 14 ]. However, individual plant volatiles often exhibit weak attractiveness when presented alone [ 12 , 14 , 17 ]. Understanding how some chemical compounds interact with sex pheromones may increase their utility in pest management programs. For example, combinations of plant volatiles and sex pheromones can be attractive to tortricid moths, as shown by Varela et al. [ 18 ], who reported increased responses of male G. molesta when suboptimal doses of sex pheromones were added to a 5-compound mixture. While significant strides have been made in devising attractants and improved trapping systems for G. molesta , refinements are still needed, particularly when potential behavioral differences among pest populations in response to lures in different regions of the world are considered [ 19 , 20 ]. Of particular interest is the development of synergistic lures that may provide high-quality attractant signals to both male and female moths, resulting in more reliable attraction under variable environmental conditions [ 21 ]. Examples of lures that act synergistically in tortricid moth species other than G. molesta include the work of Knight et al. [ 13 ] and Preti et al. [ 22 ], who reported the use of a 4-kairomone blend (PHEROCON MEGALURE CM 4K DUAL®, hereafter Megalure) consisting of pear ester (ethyl ( E,Z )-2,4-decadienoate), acetic acid, ( E )-4,8-dimethyl-1,3,7-nonatriene (DMNT) and pyranoid linalool oxide, which significantly increased catches of the codling moth Cydia pomonella (Linnaeus) (CM) (Lepidoptera: Tortricidae), males and females in the western USA compared with traps baited with only the CM sex pheromone. Benzaldehyde (BEN), an aromatic compound that is released by plants largely in the Rosaceae family, has been shown to increase the response of male but not female G. molesta when it is added to traps baited with Megalure [ 23 ]. In the same study, a 2-kairomone blend (linalool oxide and DMNT) attracted threelined leafrollers, Pandemis limitata (Robinson), only when BEN was added to the 2-kairomone blend. The observed differences in moth responses with and without BEN prompted us to further explore its interaction with semiochemical lures. In this study, we examined the behavioral responses of both male and female Grapholita molesta to BEN, both alone and in combination with sex pheromone and the commercially available lure Megalure. The primary objective of this study was to evaluate whether BEN can increase male attraction to existing semiochemical lures and to characterize its interaction with sex pheromone across distinct seasonal periods. We sought to elucidate the types of interactions (synergistic, additive) between BEN and the G. molesta sex pheromone to inform the development and refinement of semiochemical-based monitoring and management strategies for G. molesta . Results 2021 study Midseason captures (17 June to 22 July) . During this time period, the mixed-model analysis revealed a significant effect of treatment on the number of male G. molesta captured (F₇ˏ₂₁ = 5.1, P < 0.001). Megalure traps containing BEN at a low dose (BEN-L) caught significantly more male G. molesta than unbaited traps did (Fig. 1 A). Compared with the unbaited traps, all the other treatments did not attract male G. molesta . Only BEN at a low dose (BEN-L) and Megalure alone attracted female G. molesta during this period (Table 1 ). Only 14 C. pomonella males were captured by traps across all the treatments (S1 Table). Table 1 Mean number of female G. molesta captured according to the trapping period in 2021 and 2022. The release rates of benzaldehyde were 3, 6, 12, and 18 mg/day for the very low (VL), low (L), medium (M), and high (H) doses, respectively. Lures Average G. molesta female captures 2021 2022 Mid Late Early Mid BEN-VL - - 0 0 BEN-L 0.25 0 0.2 0 BEN-M 0 0 0 0 BEN-H 0 0 - - MEG 0.25 0.75 3.8 0.2 MEG + BEN-VL - - 3.4 0.6 MEG + BEN-L 0 0.25 4.6 0 MEG + BEN-M 0 0.75 3.2 0 MEG + BEN-H 0 0 - - Unbaited 0 0 0 0 Late season captures (23 July to 2 September). A significant effect of treatment was detected on male G. molesta captures (F₇ˏ₂₁ = 8.9, P < 0.001). During this period, traps baited with a low dose of BEN (BEN-L), Megalure with BEN-L, and Megalure with a medium dose of BEN (BEN-M) captured significantly more male G. molesta than nonbaited traps did (Fig. 1 B). All other treatments were statistically similar. In general, BEN at relatively high doses had a negative effect on G. molesta capture during this period (Fig. 1 B). A total of 10 female G. molesta were captured across all the treatments, with the highest average (1.0) recorded in Megalure + BEN-L (Table 1 ). With respect to C. pomonella , 10 male C. pomonella were captured during this period across all the treatments. 2022 study Early season captures (28 April to 8 June). The G. molesta populations peaked during this period, with 2,618 G. molesta (males and females) captured across all the treatments. The mixed-model analysis revealed a significant effect of treatment on the number of male G. molesta captured (F₇ˏ₂₈ = 24.3, P < 0.001). BEN alone and in combination with Megalure was attractive to male G. molesta . Compared with traps baited with BEN at very low doses (BEN-VL), traps baited with Megalure and BEN-L captured significantly more male G. molesta (Fig. 2 A). All other treatments resulted in intermediate levels of male G. molesta attraction. Numerically, Megalure + BEN-L captured the greatest number of female G. molesta (4.6 on average) during this period (Table 1 ). A total of 22 male C. pomonella were captured across all the treatments in this period (S1 Table), which was insufficient for statistical analysis. Midseason captures (9 June to 25 July) . The number of male G. molesta caught in traps differed significantly among the treatments (F₇ˏ₂₈ = 32.5, P < 0.001). Coinciding with the previous period, traps baited with Megalure + BEN-M captured significantly more male G. molesta adults than traps baited with Megalure + BEN-M, BEN-L and BEN-VL (Fig. 2 B). The level of attraction of BEN to G. molesta , regardless of the dose, was statistically similar to that of Megalure alone. Compared with the nonbaited control, all the treatments attracted significantly more male G. molesta . Only 4 female G. molesta were captured by traps during this period (Table 1 ). A total of 45 male C. pomonella were captured across all the treatments (S1 Table), which was very low for statistical analysis. 2024 study. Following the evaluation of BEN in combination with Megalure in 2021 and 2022, the 2024 study focused more specifically on characterizing the interaction between BEN and the G. molesta sex pheromone across different seasonal periods. Early season captures (30 April to 21 May). During this season, G. molesta populations peaked, with 67% of total male captures recorded across treatments. A mixed-model analysis indicated a significant effect of treatment (F₃,₁₆ = 34.7, P < 0.001), with the combination of the G. molesta sex pheromone and BEN capturing significantly more G. molesta than either BEN or pheromone alone did, whereas the captures in the BEN and pheromone treatments were statistically similar. ROI analysis provided evidence of an additive interaction between BEN and pheromone (Table 2 ). Table 2 Types of interactions observed between single attractants (Pheromone or BEN) and combined attractants (Pheromone + BEN) across early, middle, and late seasons (2024), as determined by ratios of interaction. Season (2024) Chemical Ratio of interaction (ROI) a Type of interaction Early Pheromone + BEN 0.87 ± 0.13 additive Mid Pheromone + BEN 1.55 ± 0.12 * Synergistic Late Pheromone + BEN 0.85 ± 0.03 additive Pheromone = OFM L2 ( G. molesta sex pheromone); BEN = Benzaldehyde. a Mean ± SE. * P < 0.05 for the null hypothesis that the ratio of interaction (ROI) = 1 (Hammack, 1996), according to a two-tailed t test (df = 4) Midseason captures (18 June to 7 July). During the middle of the season, the treatment effects remained significant (F₃,₁₆ = 20.2, P < 0.001), with the pheromone + BEN combination capturing more G. molesta than BEN, pheromone, or the control, although the captures among the BEN, pheromone, and control treatments were statistically similar. ROI analysis during this period revealed a synergistic interaction between BEN and pheromone (Table 2 ). Late season captures (5 August to 28 August). In the late season, the treatment effects were again significant (F₃,₁₆ = 33.7, P < 0.001), with the pheromone + BEN combination capturing the greatest number of G. molesta , whereas BEN and pheromone captures were statistically similar. As in the early season, the ROI analysis suggested an additive interaction between BEN and pheromone (Table 2 ). Discussion Herbivorous insects possess remarkable olfactory capabilities, allowing them to filter relevant odors from complex volatile blends emitted by host and nonhost plants [ 1 , 24 ]. While a few dominant compounds typically characterize plant volatile blends [ 25 ], minor components often play a critical role in shaping the behavioral response [ 15 , 26 ]. Our findings indicate that BEN is highly attractive to male G. molesta and synergistically interacts with sex pheromones during the middle of the season but has an additive effect in the early and late seasons. This finding aligns with previous research demonstrating that host-plant-derived volatiles can increase pheromone-mediated attraction in tortricid moths [ 12 – 16 ]. The combined results from 2021 and 2022 demonstrated that adding a low dose of BEN to Megalure (MEG + BEN-L) significantly increased its attractiveness to male G. molesta . Interestingly, in 2021, a higher BEN release rate (BEN-H) reduced G. molesta captures when used alone or in combination with Megalure, leading to its exclusion from the 2022 trials in favor of a lower dose. This pattern aligns with findings of Piñero & Dorn [ 15 ] and Piñero et al. [ 16 ], who reported that lower doses of BEN elicited stronger physiological and behavioral responses in G. molesta females. Notably, the 2022 study showed that BEN alone, at all tested release rates, was as attractive as Megalure to males. This is the second report demonstrating a strong male G. molesta response to BEN, following Giri et al. [ 23 ]. Previous studies have shown that insects exhibit seasonal variations in their responsiveness to semiochemicals, possibly due to shifts in mating behavior, host availability, or competitive pressures (Piñero & Dorn, 2007; Tasin et al., 2006) [ 12 , 15 , 16 ]. In the present study, trap captures varied seasonally across all study years, with significantly higher G. molesta catches in the early-season periods than in the mid- and late-season periods, regardless of the lure type. This seasonal variation in trap capture may be due to the synchronized emergence of adults of the overwintered generation [ 27 ]. As the previous season ends, more larvae enter diapause in response to decreasing day length, leading to a large population of moths emerging in spring and causing high early-season trap capture [ 27 ]. However, despite their large numbers, egg production is lower because of the fitness cost of diapause, such as reduced female fecundity [ 28 ]. As the season progresses, fewer moths emerge in later generations, resulting in lower trap captures in the middle- and late-season periods. Benzaldehyde, a naturally occurring aromatic aldehyde, is derived from the hydrolysis of amygdalin [ 29 , 30 ] and is a predominant volatile in Prunus spp. (Rosaceae) [ 14 ]. While BEN has been widely studied as a semiochemical agent in Curculionidae [ 31 – 36 ], it has also shown promising applications in Lepidoptera, including Spodoptera litura (Fabricius) [ 37 ], Helicoverpa armigera (Hübner) [ 38 ], Pieris rapae (Linnaeus) [ 39 ], and Manduca sexta (Linnaeus) [ 40 ]. The increased attraction observed when BEN is combined with sex pheromones suggests that this compound may play a role in mimicking host cues that amplify pheromone responses. From a practical perspective, our results suggest that BEN could be a valuable addition to existing lures, particularly when it is used at optimal dosages and for seasonal timings. Benzaldehyde alone was shown to be as attractive to male G. molesta as a sex pheromone across the entire growing season in 2024. However, its interaction with the pheromone varied seasonally, exhibiting additive effects in the early and late seasons but a synergistic effect in the middle of the season. The additive effect observed early in the season, despite high G. molesta populations, might be due to a ‘dilution’ effect of naturally emitted BEN from apple flowers, which peaks during petal fall [ 41 , 42 ]. As the season progresses, BEN emissions from fruit-bearing twigs steadily decline [ 42 ]. This may explain the synergistic interaction observed in the middle of the season when background levels in the orchard atmosphere are relatively low. In contrast, the late-season additive effect may be attributed to shifts in the volatile profile of the orchard as the fruit ripens. During this period, emissions of esters and alcohols increase, while aldehyde levels, including BEN, decrease [ 42 ]. This shift may result in reduced interference with the synthetic BEN in traps, maintaining an additive rather than a synergistic response. The incorporation of BEN into monitoring or control systems may enhance male responsiveness, particularly during the middle of the season, when synergistic effects are noted. While G. molesta sex pheromone lures that are specifically designed for use in mating disruption orchards already exist [ 43 , 44 ], BEN may be a promising candidate for additional trials aimed at improving detection sensitivity in these settings. Its demonstrated attractiveness, alone and in combination with the pheromone, suggests potential utility for improving trap performance under conditions where pheromone-based communication is disrupted. Furthermore, its integration into attract-and-kill strategies could contribute to population suppression by diverting males away from mating opportunities (El-Sayed et al., 2006) [ 10 ]. Female G. molesta were captured in low numbers across all study years, with a greater response to Megalure than to BEN alone. This aligns with findings by Piñero & Dorn [ 15 ], Piñero et al. [ 16 ], and Xiang et al. [ 45 ], which suggest that BEN primarily acts as a male attractant. Similarly, Luo et al. [ 46 ] demonstrated that while BEN could be detected in female Bactrocera dorsalis (Hendel) (Diptera: Tephritidae), it was insufficient to elicit a behavioral response unless it was presented in a multicomponent blend. Conclusions Our findings demonstrate that (1) BEN is a potent male attractant to a level that is comparable to the Megalure or OFM sex pheromone, (2) adding BEN at low or medium doses to these lures enhances male captures without affecting females, and (3) BEN interacts with the OFM pheromone in a season-dependent manner, showing synergism in the middle-season and additive effects in the early and late seasons. These results highlight benzaldehyde’s role in enhancing pheromone-based attraction without additional host volatiles, offering a valuable tool for improving pest monitoring and management. The incorporation of BEN into existing lures could optimize attractants on the basis of seasonal population dynamics. Future research should elucidate the neural and behavioral mechanisms underlying these effects and assess BEN–pheromone blends under varying field conditions. From an applied perspective, optimizing release rates and dispenser types could further refine their application in integrated pest management. Materials and methods Study site. Field-scale studies were conducted over a 3-year period (2021, 2022 and 2024) at the University of Massachusetts Cold Spring Orchard Research and Education Center in Belchertown, MA. The 2021 study was carried out from 17 June to 2 September, the 2022 study was carried out from 28 April to 25 July, and the 2024 study was carried out from 30 April to 28 August. This orchard received a standard insecticide spray program against key pests such as plum curculio (PC), Conotrachelus nenuphar (Herbst) (Coleoptera: Curculionidae), and apple maggot fly (AMF), Rhagoletis pomonella (Walsh) (Diptera: Tephritidae). No insecticides were sprayed against any other insect pests. Fungicides to control scab and other summer diseases were applied as necessary by the growers. The cultivars most commonly present in the test blocks were McIntosh, Empire, Red Delicious, Gala, Ginger Gold, and Cortland on the M.26 and M.7 rootstocks. Odor treatments and trap deployment. For the 2021 study, the following olfactory treatments were evaluated: (1) Benzaldehyde (BEN) low dose (BEN-L), (2) BEN medium dose (BEN-M), (3) BEN high dose (BEN-H), (4) Megalure CM 4K Dual® (hereafter referred to as Megalure), (5) Megalure + BEN-L, (6) Megalure + BEN-M, and (7) Megalure + BEN-H. The treatments evaluated in 2022 were (1) BEN very low dose (BEN-VL), (2) BEN-L, (3) BEN-M, (4) Megalure, (5) Megalure + BEN-VL, (6) Megalure + BEN-L, and (7) Megalure + BEN-M. The treatments evaluated in 2024 were (1) BEN-M, (2) Pherocon OFM L2 (pheromone), and (3) pheromone + BEN. The release rates of the various dosages of BEN were determined gravimetrically (at 25°C) to be 3, 6, 12, and 18 mg/day for very low, low, medium, and high release rates, respectively. For all the study years, unbaited traps served as negative controls. Each treatment was replicated 4 times in 2021 and 5 times in 2022 and 2024. All lures were placed inside orange-colored delta-shaped traps (Pherocon® VI, Trécé Inc., Adair, OK, USA) containing liners coated with cold-melt adhesive [ 47 ]. The experimental lures were formulated by Trécé, Inc., in a proprietary black polyvinyl chloride (PVC) matrix. The traps were spaced at 15-m intervals along the perimeter of the apple blocks and were suspended from the upper third of the tree canopy. During trap set-up, the relative position of each treatment was randomized within each experimental block. The traps were rotated weekly clockwise within a replication to minimize the effect of position. All lures were renewed at 6-week intervals, and the sticky liners were replaced every 4 weeks. Data collection For the 2021 study, traps were examined beginning on 24 June and then weekly until 2 September. For the 2022 study, traps were examined beginning on 5 May and then weekly until 25 July. For the 2024 study, traps were examined beginning on 30 April and then weekly until 28 August. All the captured adult moths were identified to species and placed in 25 ml glass vials containing 70% ethanol. The sex of each moth species was identified according to Fuková et al. [ 48 ] and Shang et al. [ 49 ] by examining the genitalia under a dissecting microscope (S1 Fig.). Statistical analysis : The number of male and female moths captured per trap per week was used as the dependent variable for the analyses. A preliminary analysis using repeated-measures ANOVA of trap-capture data from G. molesta (males only) revealed a significant interaction in all three years. This interaction indicated differential responses by the male moths to the treatments over time. Therefore, moth captures were divided into two seasonal time periods in 2021 (midseason: 17 June–22 July, and late season: 23 July --2 September), two seasonal time periods in 2022 (early season: 28 April–8 June, and midseason: 9 June–25 July) and three seasonal time periods in 2024 (early season: 30 April–21 May, midseason: 18 June–9 July, and late season: 5 August–28 August). For each year, the number of male G. molesta captured in each period was analyzed via generalized linear mixed models with a Poisson distribution, which assessed the effects of ‘treatment’ (fixed effect) and ‘replicate’ (random factor) and the 2-way interactions among them. Overdispersion was tested by examining the deviance goodness-of-fit test via a log link function. For all analyses, the data were transformed to (x + 0.5)^ 1/2 prior to analysis to stabilize variances. Where appropriate, treatment means were compared using Tukey’s protected HSD test (α = 0.05). In 2024, in addition to ANOVA, we performed comparisons of ratios of interaction (ROIs) [ 27 ] to examine the type of interaction (inhibitory, additive, or synergistic) among single- versus 2-component odor treatments. In our case, an ROI=[(pheromone + BEN) + Control]/[(Pheromone) + (BEN)], where (pheromone) represents G. molesta captures by traps baited with OFM L2 sex pheromone lure, (BEN) represents the capture of G. molesta by traps baited with BEN, (pheromone + BEN) denotes G. molesta captured by traps baited with OFM sex pheromone and BEN together, and the control represents the unbaited trps. ROI values significantly less than, equal to, or greater than 1 corresponded to inhibitory, additive, or synergistic interactions, respectively, between odor components [ 50 , 51 ]. Each ROI value was derived from one of the five replicates performed for each odor group. A two-tailed Student’s t test using such ROI values was subsequently performed to test the null hypothesis of ROI = 1 for each odor evaluated. All the statistical analyses were performed via STATISTICA v.13 (TIBCO Software Inc., Palo Alto, CA, USA) [ 52 ]. Female G. molesta and C. pomonella captures were not analyzed statistically because of insufficient numbers. Declarations Acknowledgments We thank Heriberto Godoy-Hernandez (University of Massachusetts, Amherst) for technical assistance. This material is based upon work supported by the National Institute of Food and Agriculture (NIFA), U.S. Department of Agriculture (USDA), the Center for Agriculture, Food and the Environment and the Stockbridge School of Agriculture at University of Massachusetts Amherst, under project number MAS00562. The contents are solely the responsibility of the authors and do not necessarily represent the official views of the USDA or NIFA. Author contributions statement J.C.P., A.G. and B.S. conceived the experiments; A.G. conducted the experiments; J.C.P. and A.G. analyzed the results. B.S. equally contributed to the syntheses of results and discussion. All authors reviewed the manuscript. Data Availability Declaration The datasets generated and analyzed during the current study are available from the corresponding author on reasonable request. References Suckling, D. M. Issues affecting the use of pheromones and other semiochemicals in orchards. Crop Prot. 19 , 677–683. https://doi.org/10.1016/S0261-2194(00)00090-9 (2000). Hern, A. & Dorn, S. Induction of volatile emissions from ripening apple fruits infested with Cydia pomonella and the attraction of adult females. Entomol. Exp. Appl. 102 , 145–151. https://doi.org/10.1046/j.1570-7458.2002.00934.x (2002). Cook, S. M., Khan, Z. R. & Pickett, J. A. The use of push-pull strategies in integrated pest management. Annu. Rev. 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Biol. 214 , 637–645. https://doi.org/10.1242/jeb.047316 (2011). Knight, A. L., Basoalto, E. & Stelinski, L. L. Variability in the efficacy of sex pheromone lures for monitoring oriental fruit moth ( Lepidoptera: Tortricidae ). J. Appl. Entomol. 140 , 261–267. https://doi.org/10.1111/jen.12253 (2016). Barros-Parada, W. et al. Captures of oriental fruit moth, Grapholita molesta (Lepidoptera: Tortricidae), in traps baited with host-plant volatiles in Chile. Appl. Entomol. Zool. 53 , 193–204. https://doi.org/10.1007/s13355-017-0543-7 (2018). Dorn, S. & Piñero, J. C. How do key tree-fruit pests detect and colonize their hosts: mechanisms and applications for IPM. In Biorational Tree-Fruit Pest Management (eds. Aluja, M., Leskey, T. C. & Vincent, C.) 85–109 (CABI, Wallingford), (2009). Preti, M. et al. Comparison of new kairomone-based lures for Cydia pomonella (Lepidoptera: Tortricidae) in Italy and USA. Insects 12, 72, (2021). https://doi.org/10.3390/insects12010072 Giri, A. P., Short, B. D. & Piñero, J. C. Male and female tortricid moth response to nonpheromonal semiochemicals. Insects 14 , 884. https://doi.org/10.3390/insects14110884 (2023). Bruce, T. J. A., Wadhams, L. J. & Woodcock, C. M. Insect host location: a volatile situation. Trends Plant. Sci. 10 , 269–274. https://doi.org/10.1016/j.tplants.2005.04.003 (2005). Hern, A. & Dorn, S. Monitoring seasonal variation in apple fruit volatile emissions in situ using solid-phase microextraction. Phytochem Anal. 14 , 232–240. https://doi.org/10.1002/pca.709 (2003). Masson, C. & Mustaparta, H. Chemical information processing in the olfactory system of insects. Physio Rev. 70 , 199–245. https://doi.org/10.1152/physrev.1990.70.1.199 (1990). Borchert, D. M., Stinner, R. E., Walgenbach, J. F. & Kennedy, G. G. Oriental fruit moth ( Lepidoptera: Tortricidae ) phenology and management with methoxyfenozide in North Carolina apples. J. Econ. Entomol. 97 , 1353–1364. https://doi.org/10.1093/jee/97.4.1353 (2004). Phillips, J. H. & Proctor, J. R. Studies of fecundity and behavior of the oriental fruit moth, Grapholita molesta (Lepidoptera: Tortricidae), on the Niagara Peninsula of Ontario. Can. Entomol. 101 , 1024–1033. https://doi.org/10.4039/Ent1011024-10 (1969). Fieser, L. F. & Fieser, M. Org. Chem 368–369 (D. C. Heath and Company, 1944). Opgrande, J. L. et al. Benzaldehyde. In Kirk-Othmer Enc. Chem. Tech. (John Wiley & Sons, Ltd.), https://doi.org/10.1002/0471238961.0205142615160718.a01 (2000). Leskey, T. C., Prokopy, R. J., Wright, S. E., Phelan, P. L. & Haynes, L. W. Evaluation of individual components of plum odor as potential attractants for adult plum curculios. J. Chem. Ecol. 27 , 1–17. https://doi.org/10.1023/A:1005667430877 (2001). Piñero, J. C., Wright, S. E. & Prokopy, R. J. Response of plum curculio ( Coleoptera: Curculionidae ) to odor-baited traps near woods. J. Econ. Entomol. 94 , 1386–1397. https://doi.org/10.1603/0022-0493-94.6.1386 (2001). Leskey, T. C. & Wright, S. E. Monitoring plum curculio, Conotrachelus nenuphar ( Coleoptera: Curculionidae ), populations in apple and peach orchards in the Mid-Atlantic. J. Econ. Entomol. 97 , 79–88. https://doi.org/10.1093/jee/97.1.79 (2004). Piñero, J. C. & Prokopy, R. J. Temporal dynamics of plum curculio, Conotrachelus nenuphar (Herbst.) ( Coleoptera: Curculionidae ), immigration into an apple orchard in Massachusetts. Environ. Entomol. 35 , 413–422. https://doi.org/10.1603/0046-225X-35.2.413 (2006). Lohonyai, Z. et al. Benzaldehyde: an alfalfa-related compound for the spring attraction of the pest weevil Sitona humeralis ( Coleoptera: Curculionidae ). Pest Manag Sci. 75 , 3153–3159. https://doi.org/10.1002/ps.5431 (2019). Ethington, M. W. et al. Chemically mediated colonization of black cherry by the peach bark beetle, Phloeotribus liminaris . J. Chem. Eco . 47 , 303–312. https://doi.org/10.1007/s10886-021-01256-z (2021). Fang, Y. et al. The synergistic attractiveness effect of plant volatiles to sex pheromones in a moth. J. Asia Pac. Entomol. 21 , 380–387. https://doi.org/10.1016/j.aspen.2018.01.009 (2018). Bruce, T. J. & Cork, A. Electrophysiological and behavioral responses of female Helicoverpa armigera to compounds identified in flowers of African marigold, Tagetes erecta . J. Chem. Eco . 27 , 1119–1131. https://doi.org/10.1023/A:1010359811418 (2001). Honda, K., Ômura, H. & Hayashi, N. Identification of floral volatiles from Ligustrum japonicum that stimulate flower-visiting by cabbage butterfly, Pieris rapae . J. Chem. Eco . 24 , 2167–2180. https://doi.org/10.1023/A:1020750029362 (1998). Riffell, J. A., Lei, H., Christensen, T. A. & Hildebrand, J. G. Characterization and coding of behaviorally significant odor mixtures. Curr. Biol. 19 , 335–340. https://doi.org/10.1016/j.cub.2009.01.041 (2009). Buchbauer, G., Jirovetz, L., Wasicky, M. & Nikiforov, A. Headspace and essential oil analysis of apple flowers. J. Agri Food Chem. 41 , 116–118 (1993). Vallat, A. & Dorn, S. Changes in volatile emissions from apple trees and associated response of adult female codling moths over the fruit-growing season. J. Agri Food Chem. 53 , 4083–4090. https://doi.org/10.1021/jf048499u (2005). Knight, A. et al. Monitoring oriental fruit moth ( Lepidoptera : Tortricidae ) with the Ajar bait trap in orchards under mating disruption. J. Appl. Entomol. 137 , 650–660. https://doi.org/10.1111/jen.12061 (2013). Walgenbach, J. F. et al. Comparison of sex pheromone and kairomone-enhanced pheromone lures for monitoring Oriental Fruit Moth (Lepidoptera: Tortricidae) in mating disruption and nondisruption tree fruit orchards. Environ. Entomol. 50 , 1063–1074. https://doi.org/10.1093/ee/nvab056 (2021). Xiang, H. M. et al. Peach-specific aldehyde nonanal attracts female oriental fruit moths, Grapholita molesta (Lepidoptera: Tortricidae). J. Asia Pac. Entomol. 20 , 1419–1424. https://doi.org/10.1016/j.aspen.2017.08.006 (2017). Luo, Z. et al. Benzaldehyde acts as a behaviorally active component in brewer’s yeast protein powder which attracts Bactrocera dorsalis through olfaction. J. Chem. Eco. 1–3 (2024). https://doi.org/10.1007/s10886-024-01500-2 (2024). Myers, C. T., Krawczyk, G. & Agnello, A. M. Response of tortricid moths and nontarget insects to pheromone trap color in commercial apple orchards. J. Entomol. Sci. 44 , 69–77. https://doi.org/10.18474/0749-8004-44.1.69 (2009). Fuková, I. et al. Rapid assessment of the sex of codling moth Cydia pomonella (Linnaeus) (Lepidoptera: Tortricidae) eggs and larvae. J. Appl. Entomol. 133 , 249–261. https://doi.org/10.1111/j.1439-0418.2008.01352.x (2009). Shang, S., Liu, N., Li, W. & Zhou, J. J. Morphological characteristics of reproductive system of the codling moth Cydia pomonella . ARTOAJ 25, 1–6, (2021). https://doi.org/10.19080/ARTOAJ.2021.25.556320 Hammack, L. Corn volatiles as attractants for northern and western corn rootworm beetles (Coleoptera: Chrysomelidae: Diabrotica spp). J. Chem. Eco . 22 , 1237–1253. https://doi.org/10.1007/bf02266963 (1996). Piñero, J. C. & Prokopy, R. J. Field evaluation of plant odor and pheromonal combinations for attracting plum curculios. J. Chem. Eco . 29 , 2735–2748. https://doi.org/10.1023/B:JOEC.0000008017.16911.aa (2003). TIBCO Software Inc. Statistica (data analysis software system), version 13, http://statistica.io. (2017). Additional Declarations No competing interests reported. Supplementary Files Girietal.2025ScientificReportsFINALsuppl.docx Cite Share Download PDF Status: Published Journal Publication published 15 Oct, 2025 Read the published version in Scientific Reports → Version 1 posted Editorial decision: Revision requested 30 Jun, 2025 Reviews received at journal 25 Jun, 2025 Reviews received at journal 20 Jun, 2025 Reviewers agreed at journal 05 Jun, 2025 Reviewers agreed at journal 03 Jun, 2025 Reviewers invited by journal 03 Jun, 2025 Editor assigned by journal 27 May, 2025 Editor invited by journal 27 May, 2025 Submission checks completed at journal 23 May, 2025 First submitted to journal 10 May, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. 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-6637201","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":466539699,"identity":"1e4fa0f2-35bf-4ea2-9da5-2e87c4b6a634","order_by":0,"name":"Ajay P. Giri","email":"","orcid":"","institution":"University of Massachusetts","correspondingAuthor":false,"prefix":"","firstName":"Ajay","middleName":"P.","lastName":"Giri","suffix":""},{"id":466539700,"identity":"20cd33b7-59df-4127-9ee0-9f8c6ec19dfb","order_by":1,"name":"Brent D. Short","email":"","orcid":"","institution":"Trécé Inc","correspondingAuthor":false,"prefix":"","firstName":"Brent","middleName":"D.","lastName":"Short","suffix":""},{"id":466539701,"identity":"2f08c0c8-dab2-4703-a8e8-ba6784fca20b","order_by":2,"name":"Jaime C. Pinero","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAwElEQVRIiWNgGAWjYBACxgbGBhAthxAgVosx8VpgIBGmkrAW5tnNbQ8+7rFJ33Dt8OOPPxhsZDccIOSwOQfbDWc8S8vdcDvNTJqHIc2YsJYZiW3SPAcOA7UkmDEzMBxOJE7LnwOH0w1up38GOuw/kVoYDhxOMLidYyDBw3CACC1zDrZJ9hxIM5x5O6dMmscg2XgmIS2Gs9ufSfw4YCPPdzt988cfFXayfQS1zEDhGhBQDgLyEkQoGgWjYBSMghEOAMhWSRL2MyT2AAAAAElFTkSuQmCC","orcid":"","institution":"University of Massachusetts","correspondingAuthor":true,"prefix":"","firstName":"Jaime","middleName":"C.","lastName":"Pinero","suffix":""}],"badges":[],"createdAt":"2025-05-11 02:08:11","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6637201/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6637201/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1038/s41598-025-19855-1","type":"published","date":"2025-10-15T15:58:14+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":84088900,"identity":"d90a3641-d5bb-456b-a6e3-8384d315e5a8","added_by":"auto","created_at":"2025-06-06 15:40:34","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":208841,"visible":true,"origin":"","legend":"\u003cp\u003eCaptures (mean ± SEM) of male \u003cem\u003eG. molesta\u003c/em\u003e in delta traps baited with seven olfactory treatments and nonbaited traps in (A) the middle season (17 June--22 July) and (B) late season (23 July --2 September) of2021. The release rates of benzaldehyde were determined to be 6, 12, and 18 mg/day for the low (L), medium (M), and high (H) dosages, respectively. Bars superscribed with the same letter do not differ significantly among treatments (Tukey’s protected HSD test, α = 0.05).\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-6637201/v1/e1219263bf3b0373ded7fcad.png"},{"id":84088906,"identity":"19522241-412c-42ea-8274-32d70942b4a5","added_by":"auto","created_at":"2025-06-06 15:40:34","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":293814,"visible":true,"origin":"","legend":"\u003cp\u003eCaptures (mean ± SEM) of male \u003cem\u003eG. molesta\u003c/em\u003e in delta traps baited with seven olfactory treatments and nonbaited traps in the (A) early-season (April 28 to June 8) and (B) midseason (July 9 to July 25) of 2022. The release rates of benzaldehyde were determined to be 3, 6, and 12 mg/day for the very low (VL), low (L) and medium (M) dosages, respectively. Bars superscribed with the same letter do not differ significantly among treatments (Tukey’s protected HSD test, α = 0.05).\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-6637201/v1/a000d7183a72f0b279e88dff.png"},{"id":84089308,"identity":"c2d7c98d-7305-40e4-950c-7c10f20eecfb","added_by":"auto","created_at":"2025-06-06 15:48:34","extension":"jpeg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":71425,"visible":true,"origin":"","legend":"\u003cp\u003eCaptures (mean ± SEM) of male \u003cem\u003eG. molesta\u003c/em\u003e in delta traps baited with three olfactory treatments and nonbaited traps in the early season (30 April--21 May), denoted by yellow lines inside the bar graph; the middle season (18 June--9 July), denoted by green dots inside the bar graph; and the late season (5 August--28 August), denoted by solid red bars, in 2024. Bars superscribed with the same letter do not differ significantly among treatments (Tukey’sprotected HSD test, α = 0.05).\u003c/p\u003e","description":"","filename":"floatimage3.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-6637201/v1/4da4d13048934a9f97155f7b.jpeg"},{"id":93956031,"identity":"f78f10b4-8c73-4ffd-b5ec-8de45f026cfc","added_by":"auto","created_at":"2025-10-20 16:09:30","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1262672,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6637201/v1/7706a5ce-4b49-46d2-85e3-4de188c029a1.pdf"},{"id":84089310,"identity":"48cb24a6-e0eb-4e42-b7f2-04805f82904a","added_by":"auto","created_at":"2025-06-06 15:48:34","extension":"docx","order_by":0,"title":"","display":"","copyAsset":false,"role":"supplement","size":373833,"visible":true,"origin":"","legend":"","description":"","filename":"Girietal.2025ScientificReportsFINALsuppl.docx","url":"https://assets-eu.researchsquare.com/files/rs-6637201/v1/46da3b85f9e01568088dd47c.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Season-Dependent Synergism between the Male- Attractive Plant Volatile Benzaldehyde and the Sex Pheromone of the Oriental Fruit Moth, Grapholita molesta","fulltext":[{"header":"Introduction","content":"\u003cp\u003eInsects rely on highly specialized olfactory systems to detect, process, and respond to chemical cues in their environment [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e]. Within Lepidoptera, semiochemicals, including host plant volatiles and sex pheromones, play pivotal roles in mediating insect behavior. This information is crucial to better understand the patterns of host plant use and for the development of more efficient monitoring and/or control tactics [\u003cspan additionalcitationids=\"CR3\" citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eA major advancement in pest control is the implementation of mating disruption techniques, wherein synthetic sex pheromones are deployed at high densities to interfere with mate finding, thereby reducing reproductive success [\u003cspan additionalcitationids=\"CR6 CR7 CR8\" citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e]. Additionally, semiochemical-based attractants that target both male and female moths are being explored for their potential in integrated pest management (IPM) strategies, including attract-and-kill techniques [\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e]. Because the success of these approaches relies on a comprehensive understanding of insect olfactory responses and the complex interactions between semiochemicals [\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e], unremitting research is essential to refine existing systems and develop more effective strategies for pest management.\u003c/p\u003e \u003cp\u003eResearch suggests that complex blends of host-plant-derived volatiles are often required to elicit significant attraction in tortricid moths [\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e, \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e]. This phenomenon is particularly relevant for the Oriental fruit moth (OFM), \u003cem\u003eGrapholita molesta\u003c/em\u003e (Busck), a significant pest of stone and pome fruits worldwide [\u003cspan additionalcitationids=\"CR15\" citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e]. Notably, a five-component blend of three green leaf volatiles and two aromatic compounds (benzaldehyde and benzonitrile) has been shown to be as attractive to mated female \u003cem\u003eG. molesta\u003c/em\u003e [\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e] as the full suite of natural peach shoot volatiles, which contains more than 20 compounds [\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e]. However, individual plant volatiles often exhibit weak attractiveness when presented alone [\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e]. Understanding how some chemical compounds interact with sex pheromones may increase their utility in pest management programs. For example, combinations of plant volatiles and sex pheromones can be attractive to tortricid moths, as shown by Varela et al. [\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e], who reported increased responses of male \u003cem\u003eG. molesta\u003c/em\u003e when suboptimal doses of sex pheromones were added to a 5-compound mixture.\u003c/p\u003e \u003cp\u003eWhile significant strides have been made in devising attractants and improved trapping systems for \u003cem\u003eG. molesta\u003c/em\u003e, refinements are still needed, particularly when potential behavioral differences among pest populations in response to lures in different regions of the world are considered [\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e]. Of particular interest is the development of synergistic lures that may provide high-quality attractant signals to both male and female moths, resulting in more reliable attraction under variable environmental conditions [\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e]. Examples of lures that act synergistically in tortricid moth species other than \u003cem\u003eG. molesta\u003c/em\u003e include the work of Knight et al. [\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e] and Preti et al. [\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e], who reported the use of a 4-kairomone blend (PHEROCON MEGALURE CM 4K DUAL\u0026reg;, hereafter Megalure) consisting of pear ester (ethyl (\u003cem\u003eE,Z\u003c/em\u003e)-2,4-decadienoate), acetic acid, (\u003cem\u003eE\u003c/em\u003e)-4,8-dimethyl-1,3,7-nonatriene (DMNT) and pyranoid linalool oxide, which significantly increased catches of the codling moth \u003cem\u003eCydia pomonella\u003c/em\u003e (Linnaeus) (CM) (Lepidoptera: Tortricidae), males and females in the western USA compared with traps baited with only the CM sex pheromone.\u003c/p\u003e \u003cp\u003eBenzaldehyde (BEN), an aromatic compound that is released by plants largely in the Rosaceae family, has been shown to increase the response of male but not female \u003cem\u003eG. molesta\u003c/em\u003e when it is added to traps baited with Megalure [\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e]. In the same study, a 2-kairomone blend (linalool oxide and DMNT) attracted threelined leafrollers, \u003cem\u003ePandemis limitata\u003c/em\u003e (Robinson), only when BEN was added to the 2-kairomone blend. The observed differences in moth responses with and without BEN prompted us to further explore its interaction with semiochemical lures.\u003c/p\u003e \u003cp\u003eIn this study, we examined the behavioral responses of both male and female \u003cem\u003eGrapholita molesta\u003c/em\u003e to BEN, both alone and in combination with sex pheromone and the commercially available lure Megalure. The primary objective of this study was to evaluate whether BEN can increase male attraction to existing semiochemical lures and to characterize its interaction with sex pheromone across distinct seasonal periods. We sought to elucidate the types of interactions (synergistic, additive) between BEN and the \u003cem\u003eG. molesta\u003c/em\u003e sex pheromone to inform the development and refinement of semiochemical-based monitoring and management strategies for \u003cem\u003eG. molesta\u003c/em\u003e.\u003c/p\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2021 study\u003c/h2\u003e \u003cp\u003e \u003cem\u003eMidseason captures (17 June to 22 July)\u003c/em\u003e. During this time period, the mixed-model analysis revealed a significant effect of treatment on the number of male \u003cem\u003eG. molesta\u003c/em\u003e captured (F₇ˏ₂₁ = 5.1, P\u0026thinsp;\u0026lt;\u0026thinsp;0.001). Megalure traps containing BEN at a low dose (BEN-L) caught significantly more male \u003cem\u003eG. molesta\u003c/em\u003e than unbaited traps did (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eA). Compared with the unbaited traps, all the other treatments did not attract male \u003cem\u003eG. molesta\u003c/em\u003e. Only BEN at a low dose (BEN-L) and Megalure alone attracted female \u003cem\u003eG. molesta\u003c/em\u003e during this period (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Only 14 \u003cem\u003eC. pomonella\u003c/em\u003e males were captured by traps across all the treatments (S1 Table).\u003c/p\u003e \u003cp\u003e \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\u003e\u003cb\u003eMean number of female\u003c/b\u003e \u003cb\u003eG. molesta\u003c/b\u003e \u003cb\u003ecaptured according to the trapping period in 2021 and 2022.\u003c/b\u003e The release rates of benzaldehyde were 3, 6, 12, and 18 mg/day for the very low (VL), low (L), medium (M), and high (H) doses, respectively.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eLures\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"4\" nameend=\"c5\" namest=\"c2\"\u003e \u003cp\u003eAverage \u003cem\u003eG. molesta\u003c/em\u003e female captures\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e \u003cp\u003e2021\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003e2022\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMid\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLate\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eEarly\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMid\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBEN-VL\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBEN-L\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBEN-M\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBEN-H\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMEG\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMEG\u0026thinsp;+\u0026thinsp;BEN-VL\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.6\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMEG\u0026thinsp;+\u0026thinsp;BEN-L\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e4.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMEG\u0026thinsp;+\u0026thinsp;BEN-M\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMEG\u0026thinsp;+\u0026thinsp;BEN-H\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eUnbaited\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cem\u003eLate season captures (23 July to 2 September).\u003c/em\u003e A significant effect of treatment was detected on male \u003cem\u003eG. molesta\u003c/em\u003e captures (F₇ˏ₂₁ = 8.9, P\u0026thinsp;\u0026lt;\u0026thinsp;0.001). During this period, traps baited with a low dose of BEN (BEN-L), Megalure with BEN-L, and Megalure with a medium dose of BEN (BEN-M) captured significantly more male \u003cem\u003eG. molesta\u003c/em\u003e than nonbaited traps did (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eB). All other treatments were statistically similar. In general, BEN at relatively high doses had a negative effect on \u003cem\u003eG. molesta\u003c/em\u003e capture during this period (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eB). A total of 10 female \u003cem\u003eG. molesta\u003c/em\u003e were captured across all the treatments, with the highest average (1.0) recorded in Megalure\u0026thinsp;+\u0026thinsp;BEN-L (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). With respect to \u003cem\u003eC. pomonella\u003c/em\u003e, 10 male \u003cem\u003eC. pomonella\u003c/em\u003e were captured during this period across all the treatments.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003e2022 study\u003c/h3\u003e\n\u003cp\u003e \u003cem\u003eEarly season captures (28 April to 8 June).\u003c/em\u003e The \u003cem\u003eG. molesta\u003c/em\u003e populations peaked during this period, with 2,618 \u003cem\u003eG. molesta\u003c/em\u003e (males and females) captured across all the treatments. The mixed-model analysis revealed a significant effect of treatment on the number of male \u003cem\u003eG. molesta\u003c/em\u003e captured (F₇ˏ₂₈ = 24.3, P\u0026thinsp;\u0026lt;\u0026thinsp;0.001). BEN alone and in combination with Megalure was attractive to male \u003cem\u003eG. molesta\u003c/em\u003e. Compared with traps baited with BEN at very low doses (BEN-VL), traps baited with Megalure and BEN-L captured significantly more male \u003cem\u003eG. molesta\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eA). All other treatments resulted in intermediate levels of male \u003cem\u003eG. molesta\u003c/em\u003e attraction. Numerically, Megalure\u0026thinsp;+\u0026thinsp;BEN-L captured the greatest number of female \u003cem\u003eG. molesta\u003c/em\u003e (4.6 on average) during this period (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). A total of 22 male \u003cem\u003eC. pomonella\u003c/em\u003e were captured across all the treatments in this period (S1 Table), which was insufficient for statistical analysis.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cem\u003eMidseason captures (9 June to 25 July)\u003c/em\u003e. The number of male \u003cem\u003eG. molesta\u003c/em\u003e caught in traps differed significantly among the treatments (F₇ˏ₂₈ = 32.5, P\u0026thinsp;\u0026lt;\u0026thinsp;0.001). Coinciding with the previous period, traps baited with Megalure\u0026thinsp;+\u0026thinsp;BEN-M captured significantly more male \u003cem\u003eG. molesta\u003c/em\u003e adults than traps baited with Megalure\u0026thinsp;+\u0026thinsp;BEN-M, BEN-L and BEN-VL (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eB). The level of attraction of BEN to \u003cem\u003eG. molesta\u003c/em\u003e, regardless of the dose, was statistically similar to that of Megalure alone. Compared with the nonbaited control, all the treatments attracted significantly more male \u003cem\u003eG. molesta\u003c/em\u003e. Only 4 female \u003cem\u003eG. molesta\u003c/em\u003e were captured by traps during this period (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). A total of 45 male \u003cem\u003eC. pomonella\u003c/em\u003e were captured across all the treatments (S1 Table), which was very low for statistical analysis.\u003c/p\u003e \u003cp\u003e \u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e2024 study.\u003c/span\u003e \u003c/p\u003e \u003cp\u003eFollowing the evaluation of BEN in combination with Megalure in 2021 and 2022, the 2024 study focused more specifically on characterizing the interaction between BEN and the \u003cem\u003eG. molesta\u003c/em\u003e sex pheromone across different seasonal periods.\u003c/p\u003e \u003cp\u003e \u003cem\u003eEarly season captures (30 April to 21 May).\u003c/em\u003e During this season, \u003cem\u003eG. molesta\u003c/em\u003e populations peaked, with 67% of total male captures recorded across treatments. A mixed-model analysis indicated a significant effect of treatment (F₃,₁₆ = 34.7, P\u0026thinsp;\u0026lt;\u0026thinsp;0.001), with the combination of the \u003cem\u003eG. molesta\u003c/em\u003e sex pheromone and BEN capturing significantly more \u003cem\u003eG. molesta\u003c/em\u003e than either BEN or pheromone alone did, whereas the captures in the BEN and pheromone treatments were statistically similar. ROI analysis provided evidence of an additive interaction between BEN and pheromone (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\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\u003eTypes of interactions observed between single attractants (Pheromone or BEN) and combined attractants (Pheromone\u0026thinsp;+\u0026thinsp;BEN) across early, middle, and late seasons (2024), as determined by ratios of interaction.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"4\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSeason\u003c/p\u003e \u003cp\u003e(2024)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eChemical\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eRatio of interaction (ROI)\u003csup\u003e\u003cem\u003ea\u003c/em\u003e\u003c/sup\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eType of interaction\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEarly\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePheromone\u0026thinsp;+\u0026thinsp;BEN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.87 \u0026plusmn; 0.13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eadditive\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMid\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePheromone\u0026thinsp;+\u0026thinsp;BEN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e1.55 \u0026plusmn; 0.12 \u003cem\u003e*\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSynergistic\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePheromone\u0026thinsp;+\u0026thinsp;BEN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.85 \u0026plusmn; 0.03\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eadditive\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"4\"\u003ePheromone\u0026thinsp;=\u0026thinsp;OFM L2 (\u003cem\u003eG. molesta\u003c/em\u003e sex pheromone); BEN\u0026thinsp;=\u0026thinsp;Benzaldehyde.\u003c/td\u003e\u003c/tr\u003e \u003ctr\u003e\u003ctd colspan=\"4\"\u003e\u003csup\u003e\u003cem\u003ea\u003c/em\u003e\u003c/sup\u003e Mean \u0026plusmn; SE.\u003c/td\u003e\u003c/tr\u003e \u003ctr\u003e\u003ctd colspan=\"4\"\u003e\u003cem\u003e*\u003c/em\u003e P\u0026thinsp;\u0026lt;\u0026thinsp;0.05 for the null hypothesis that the ratio of interaction (ROI)\u0026thinsp;=\u0026thinsp;1 (Hammack, 1996), according to a two-tailed \u003cem\u003et\u003c/em\u003e test (df\u0026thinsp;=\u0026thinsp;4)\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cem\u003eMidseason captures (18 June to 7 July).\u003c/em\u003e During the middle of the season, the treatment effects remained significant (F₃,₁₆ = 20.2, P\u0026thinsp;\u0026lt;\u0026thinsp;0.001), with the pheromone\u0026thinsp;+\u0026thinsp;BEN combination capturing more \u003cem\u003eG. molesta\u003c/em\u003e than BEN, pheromone, or the control, although the captures among the BEN, pheromone, and control treatments were statistically similar. ROI analysis during this period revealed a synergistic interaction between BEN and pheromone (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cem\u003eLate season captures (5 August to 28 August).\u003c/em\u003e In the late season, the treatment effects were again significant (F₃,₁₆ = 33.7, P\u0026thinsp;\u0026lt;\u0026thinsp;0.001), with the pheromone\u0026thinsp;+\u0026thinsp;BEN combination capturing the greatest number of \u003cem\u003eG. molesta\u003c/em\u003e, whereas BEN and pheromone captures were statistically similar. As in the early season, the ROI analysis suggested an additive interaction between BEN and pheromone (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eHerbivorous insects possess remarkable olfactory capabilities, allowing them to filter relevant odors from complex volatile blends emitted by host and nonhost plants [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e, \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e]. While a few dominant compounds typically characterize plant volatile blends [\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e], minor components often play a critical role in shaping the behavioral response [\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e, \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e]. Our findings indicate that BEN is highly attractive to male \u003cem\u003eG. molesta\u003c/em\u003e and synergistically interacts with sex pheromones during the middle of the season but has an additive effect in the early and late seasons. This finding aligns with previous research demonstrating that host-plant-derived volatiles can increase pheromone-mediated attraction in tortricid moths [\u003cspan additionalcitationids=\"CR13 CR14 CR15\" citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eThe combined results from 2021 and 2022 demonstrated that adding a low dose of BEN to Megalure (MEG\u0026thinsp;+\u0026thinsp;BEN-L) significantly increased its attractiveness to male \u003cem\u003eG. molesta\u003c/em\u003e. Interestingly, in 2021, a higher BEN release rate (BEN-H) reduced \u003cem\u003eG. molesta\u003c/em\u003e captures when used alone or in combination with Megalure, leading to its exclusion from the 2022 trials in favor of a lower dose. This pattern aligns with findings of Pi\u0026ntilde;ero \u0026amp; Dorn [\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e] and Pi\u0026ntilde;ero et al. [\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e], who reported that lower doses of BEN elicited stronger physiological and behavioral responses in \u003cem\u003eG. molesta\u003c/em\u003e females. Notably, the 2022 study showed that BEN alone, at all tested release rates, was as attractive as Megalure to males. This is the second report demonstrating a strong male \u003cem\u003eG. molesta\u003c/em\u003e response to BEN, following Giri et al. [\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e].\u003c/p\u003e \u003cp\u003ePrevious studies have shown that insects exhibit seasonal variations in their responsiveness to semiochemicals, possibly due to shifts in mating behavior, host availability, or competitive pressures (Pi\u0026ntilde;ero \u0026amp; Dorn, 2007; Tasin et al., 2006) [\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e, \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e]. In the present study, trap captures varied seasonally across all study years, with significantly higher \u003cem\u003eG. molesta\u003c/em\u003e catches in the early-season periods than in the mid- and late-season periods, regardless of the lure type. This seasonal variation in trap capture may be due to the synchronized emergence of adults of the overwintered generation [\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e]. As the previous season ends, more larvae enter diapause in response to decreasing day length, leading to a large population of moths emerging in spring and causing high early-season trap capture [\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e]. However, despite their large numbers, egg production is lower because of the fitness cost of diapause, such as reduced female fecundity [\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e]. As the season progresses, fewer moths emerge in later generations, resulting in lower trap captures in the middle- and late-season periods.\u003c/p\u003e \u003cp\u003eBenzaldehyde, a naturally occurring aromatic aldehyde, is derived from the hydrolysis of amygdalin [\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e] and is a predominant volatile in \u003cem\u003ePrunus\u003c/em\u003e spp. (Rosaceae) [\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e]. While BEN has been widely studied as a semiochemical agent in Curculionidae [\u003cspan additionalcitationids=\"CR32 CR33 CR34 CR35\" citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e], it has also shown promising applications in Lepidoptera, including \u003cem\u003eSpodoptera litura\u003c/em\u003e (Fabricius) [\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e], \u003cem\u003eHelicoverpa armigera\u003c/em\u003e (H\u0026uuml;bner) [\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e], \u003cem\u003ePieris rapae\u003c/em\u003e (Linnaeus) [\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e39\u003c/span\u003e], and \u003cem\u003eManduca sexta\u003c/em\u003e (Linnaeus) [\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e]. The increased attraction observed when BEN is combined with sex pheromones suggests that this compound may play a role in mimicking host cues that amplify pheromone responses. From a practical perspective, our results suggest that BEN could be a valuable addition to existing lures, particularly when it is used at optimal dosages and for seasonal timings.\u003c/p\u003e \u003cp\u003eBenzaldehyde alone was shown to be as attractive to male \u003cem\u003eG. molesta\u003c/em\u003e as a sex pheromone across the entire growing season in 2024. However, its interaction with the pheromone varied seasonally, exhibiting additive effects in the early and late seasons but a synergistic effect in the middle of the season. The additive effect observed early in the season, despite high \u003cem\u003eG. molesta\u003c/em\u003e populations, might be due to a \u0026lsquo;dilution\u0026rsquo; effect of naturally emitted BEN from apple flowers, which peaks during petal fall [\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e, \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e]. As the season progresses, BEN emissions from fruit-bearing twigs steadily decline [\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e]. This may explain the synergistic interaction observed in the middle of the season when background levels in the orchard atmosphere are relatively low. In contrast, the late-season additive effect may be attributed to shifts in the volatile profile of the orchard as the fruit ripens. During this period, emissions of esters and alcohols increase, while aldehyde levels, including BEN, decrease [\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e]. This shift may result in reduced interference with the synthetic BEN in traps, maintaining an additive rather than a synergistic response.\u003c/p\u003e \u003cp\u003eThe incorporation of BEN into monitoring or control systems may enhance male responsiveness, particularly during the middle of the season, when synergistic effects are noted. While \u003cem\u003eG. molesta\u003c/em\u003e sex pheromone lures that are specifically designed for use in mating disruption orchards already exist [\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e, \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e], BEN may be a promising candidate for additional trials aimed at improving detection sensitivity in these settings. Its demonstrated attractiveness, alone and in combination with the pheromone, suggests potential utility for improving trap performance under conditions where pheromone-based communication is disrupted. Furthermore, its integration into attract-and-kill strategies could contribute to population suppression by diverting males away from mating opportunities (El-Sayed et al., 2006) [\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eFemale \u003cem\u003eG. molesta\u003c/em\u003e were captured in low numbers across all study years, with a greater response to Megalure than to BEN alone. This aligns with findings by Pi\u0026ntilde;ero \u0026amp; Dorn [\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e], Pi\u0026ntilde;ero et al. [\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e], and Xiang et al. [\u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e45\u003c/span\u003e], which suggest that BEN primarily acts as a male attractant. Similarly, Luo et al. [\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e46\u003c/span\u003e] demonstrated that while BEN could be detected in female \u003cem\u003eBactrocera dorsalis\u003c/em\u003e (Hendel) (Diptera: Tephritidae), it was insufficient to elicit a behavioral response unless it was presented in a multicomponent blend.\u003c/p\u003e"},{"header":"Conclusions","content":"\u003cp\u003eOur findings demonstrate that (1) BEN is a potent male attractant to a level that is comparable to the Megalure or OFM sex pheromone, (2) adding BEN at low or medium doses to these lures enhances male captures without affecting females, and (3) BEN interacts with the OFM pheromone in a season-dependent manner, showing synergism in the middle-season and additive effects in the early and late seasons. These results highlight benzaldehyde\u0026rsquo;s role in enhancing pheromone-based attraction without additional host volatiles, offering a valuable tool for improving pest monitoring and management. The incorporation of BEN into existing lures could optimize attractants on the basis of seasonal population dynamics.\u003c/p\u003e \u003cp\u003eFuture research should elucidate the neural and behavioral mechanisms underlying these effects and assess BEN\u0026ndash;pheromone blends under varying field conditions. From an applied perspective, optimizing release rates and dispenser types could further refine their application in integrated pest management.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cp\u003e \u003cb\u003eStudy site.\u003c/b\u003e Field-scale studies were conducted over a 3-year period (2021, 2022 and 2024) at the University of Massachusetts Cold Spring Orchard Research and Education Center in Belchertown, MA. The 2021 study was carried out from 17 June to 2 September, the 2022 study was carried out from 28 April to 25 July, and the 2024 study was carried out from 30 April to 28 August. This orchard received a standard insecticide spray program against key pests such as plum curculio (PC), \u003cem\u003eConotrachelus nenuphar\u003c/em\u003e (Herbst) (Coleoptera: Curculionidae), and apple maggot fly (AMF), \u003cem\u003eRhagoletis pomonella\u003c/em\u003e (Walsh) (Diptera: Tephritidae). No insecticides were sprayed against any other insect pests. Fungicides to control scab and other summer diseases were applied as necessary by the growers. The cultivars most commonly present in the test blocks were McIntosh, Empire, Red Delicious, Gala, Ginger Gold, and Cortland on the M.26 and M.7 rootstocks.\u003cdiv class=\"BlockQuote\"\u003e\u003cp\u003e \u003cb\u003eOdor treatments and trap deployment.\u003c/b\u003e For the 2021 study, the following olfactory treatments were evaluated: (1) Benzaldehyde (BEN) low dose (BEN-L), (2) BEN medium dose (BEN-M), (3) BEN high dose (BEN-H), (4) Megalure CM 4K Dual\u0026reg; (hereafter referred to as Megalure), (5) Megalure\u0026thinsp;+\u0026thinsp;BEN-L, (6) Megalure\u0026thinsp;+\u0026thinsp;BEN-M, and (7) Megalure\u0026thinsp;+\u0026thinsp;BEN-H. The treatments evaluated in 2022 were (1) BEN very low dose (BEN-VL), (2) BEN-L, (3) BEN-M, (4) Megalure, (5) Megalure\u0026thinsp;+\u0026thinsp;BEN-VL, (6) Megalure\u0026thinsp;+\u0026thinsp;BEN-L, and (7) Megalure\u0026thinsp;+\u0026thinsp;BEN-M. The treatments evaluated in 2024 were (1) BEN-M, (2) Pherocon OFM L2 (pheromone), and (3) pheromone\u0026thinsp;+\u0026thinsp;BEN. The release rates of the various dosages of BEN were determined gravimetrically (at 25\u0026deg;C) to be 3, 6, 12, and 18 mg/day for very low, low, medium, and high release rates, respectively. For all the study years, unbaited traps served as negative controls. Each treatment was replicated 4 times in 2021 and 5 times in 2022 and 2024.\u003c/p\u003e\u003c/div\u003e\u003c/p\u003e \u003cp\u003eAll lures were placed inside orange-colored delta-shaped traps (Pherocon\u0026reg; VI, Tr\u0026eacute;c\u0026eacute; Inc., Adair, OK, USA) containing liners coated with cold-melt adhesive [\u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e47\u003c/span\u003e]. The experimental lures were formulated by Tr\u0026eacute;c\u0026eacute;, Inc., in a proprietary black polyvinyl chloride (PVC) matrix. The traps were spaced at 15-m intervals along the perimeter of the apple blocks and were suspended from the upper third of the tree canopy. During trap set-up, the relative position of each treatment was randomized within each experimental block. The traps were rotated weekly clockwise within a replication to minimize the effect of position. All lures were renewed at 6-week intervals, and the sticky liners were replaced every 4 weeks.\u003c/p\u003e \u003cp\u003e \u003cstrong\u003eData collection\u003c/strong\u003e \u003cp\u003eFor the 2021 study, traps were examined beginning on 24 June and then weekly until 2 September. For the 2022 study, traps were examined beginning on 5 May and then weekly until 25 July. For the 2024 study, traps were examined beginning on 30 April and then weekly until 28 August. All the captured adult moths were identified to species and placed in 25 ml glass vials containing 70% ethanol. The sex of each moth species was identified according to Fukov\u0026aacute; et al. [\u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e48\u003c/span\u003e] and Shang et al. [\u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e49\u003c/span\u003e] by examining the genitalia under a dissecting microscope (S1 Fig.).\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eStatistical analysis\u003c/b\u003e: The number of male and female moths captured per trap per week was used as the dependent variable for the analyses. A preliminary analysis using repeated-measures ANOVA of trap-capture data from \u003cem\u003eG. molesta\u003c/em\u003e (males only) revealed a significant interaction in all three years. This interaction indicated differential responses by the male moths to the treatments over time. Therefore, moth captures were divided into two seasonal time periods in 2021 (midseason: 17 June\u0026ndash;22 July, and late season: 23 July --2 September), two seasonal time periods in 2022 (early season: 28 April\u0026ndash;8 June, and midseason: 9 June\u0026ndash;25 July) and three seasonal time periods in 2024 (early season: 30 April\u0026ndash;21 May, midseason: 18 June\u0026ndash;9 July, and late season: 5 August\u0026ndash;28 August). For each year, the number of male \u003cem\u003eG. molesta\u003c/em\u003e captured in each period was analyzed via generalized linear mixed models with a Poisson distribution, which assessed the effects of \u0026lsquo;treatment\u0026rsquo; (fixed effect) and \u0026lsquo;replicate\u0026rsquo; (random factor) and the 2-way interactions among them. Overdispersion was tested by examining the deviance goodness-of-fit test via a log link function. For all analyses, the data were transformed to (x\u0026thinsp;+\u0026thinsp;0.5)^\u003csup\u003e1/2\u003c/sup\u003e prior to analysis to stabilize variances. Where appropriate, treatment means were compared using Tukey\u0026rsquo;s protected HSD test (α\u0026thinsp;=\u0026thinsp;0.05).\u003c/p\u003e \u003cp\u003eIn 2024, in addition to ANOVA, we performed comparisons of ratios of interaction (ROIs) [\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e] to examine the type of interaction (inhibitory, additive, or synergistic) among single- versus 2-component odor treatments. In our case, an ROI=[(pheromone\u0026thinsp;+\u0026thinsp;BEN)\u0026thinsp;+\u0026thinsp;Control]/[(Pheromone) + (BEN)], where (pheromone) represents \u003cem\u003eG. molesta\u003c/em\u003e captures by traps baited with OFM L2 sex pheromone lure, (BEN) represents the capture of \u003cem\u003eG. molesta\u003c/em\u003e by traps baited with BEN, (pheromone\u0026thinsp;+\u0026thinsp;BEN) denotes \u003cem\u003eG. molesta\u003c/em\u003e captured by traps baited with OFM sex pheromone and BEN together, and the control represents the unbaited trps. ROI values significantly less than, equal to, or greater than 1 corresponded to inhibitory, additive, or synergistic interactions, respectively, between odor components [\u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e50\u003c/span\u003e, \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e51\u003c/span\u003e]. Each ROI value was derived from one of the five replicates performed for each odor group. A two-tailed Student\u0026rsquo;s t test using such ROI values was subsequently performed to test the null hypothesis of ROI\u0026thinsp;=\u0026thinsp;1 for each odor evaluated.\u003c/p\u003e \u003cp\u003eAll the statistical analyses were performed via STATISTICA v.13 (TIBCO Software Inc., Palo Alto, CA, USA) [\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e52\u003c/span\u003e]. Female \u003cem\u003eG. molesta\u003c/em\u003e and \u003cem\u003eC. pomonella\u003c/em\u003e captures were not analyzed statistically because of insufficient numbers.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgments\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe thank Heriberto Godoy-Hernandez (University of Massachusetts, Amherst) for technical assistance. This material is based upon work supported by the National Institute of Food and Agriculture (NIFA), U.S. Department of Agriculture (USDA), the Center for Agriculture, Food and the Environment and the Stockbridge School of Agriculture at University of Massachusetts Amherst, under project number MAS00562. The contents are solely the responsibility of the authors and do not necessarily represent the official views of the USDA or NIFA.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions statement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eJ.C.P., A.G. and B.S. conceived the experiments; A.G. conducted the experiments; J.C.P. and A.G. analyzed the results. B.S. equally contributed to the syntheses of results and discussion. All authors reviewed the manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData Availability Declaration\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe datasets generated and analyzed during the current study are available from the corresponding author on reasonable request.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eSuckling, D. M. Issues affecting the use of pheromones and other semiochemicals in orchards. \u003cem\u003eCrop Prot.\u003c/em\u003e \u003cb\u003e19\u003c/b\u003e, 677\u0026ndash;683. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/S0261-2194(00)00090-9\u003c/span\u003e\u003cspan address=\"10.1016/S0261-2194(00)00090-9\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2000).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHern, A. \u0026amp; Dorn, S. 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(2017).\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"Behavior, monitoring, semiochemical, IPM, synergism, sensory ecology","lastPublishedDoi":"10.21203/rs.3.rs-6637201/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6637201/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eHerbivorous insects possess highly developed olfactory systems that enable them to detect, process, and respond to volatile cues emitted by host and nonhost plants. This study evaluated the field response of male and female Oriental fruit moths. \u003cem\u003eGrapholita molesta\u003c/em\u003e (Lepidoptera: Tortricidae) to the aromatic plant volatile benzaldehyde (BEN). We evaluated BEN alone, in combination with the \u003cem\u003eG. molesta\u003c/em\u003e sex pheromone, or with a four-component kairomonal lure (PHEROCON MEGALURE CM 4K DUAL\u0026reg;). Our findings demonstrate that (1) BEN alone is a male \u003cem\u003eG. molesta\u003c/em\u003e attractant with efficacy comparable to that of Megalure or the sex pheromone; (2) the addition of BEN at low or medium doses to either Megalure or pheromone lures significantly increases the number of male captures; and (3) BEN interacts with the \u003cem\u003eG. molesta\u003c/em\u003e pheromone in a season-dependent manner, exhibiting synergistic effects during the middle-season and additive effects in the early and late seasons. These results highlight the potential of BEN to enhance pheromone-based lures without the need for additional host volatiles, providing a valuable tool for optimizing semiochemical-based monitoring and control strategies tailored to seasonal population dynamics.\u003c/p\u003e","manuscriptTitle":"Season-Dependent Synergism between the Male- Attractive Plant Volatile Benzaldehyde and the Sex Pheromone of the Oriental Fruit Moth, Grapholita molesta","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-06-06 15:40:30","doi":"10.21203/rs.3.rs-6637201/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-06-30T21:28:53+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-06-25T23:13:38+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-06-20T12:08:02+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"71011495751631274578710217464737945205","date":"2025-06-05T20:55:45+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"6215352722935291203754043797848829289","date":"2025-06-03T18:02:17+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-06-03T16:40:13+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-05-27T14:16:49+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"","date":"2025-05-27T04:46:22+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-05-23T14:44:08+00:00","index":"","fulltext":""},{"type":"submitted","content":"Scientific Reports","date":"2025-05-11T01:59:00+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"8a47c46f-107b-47bf-ab59-4295cf19728e","owner":[],"postedDate":"June 6th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[{"id":49534317,"name":"Biological sciences/Zoology/Entomology"},{"id":49534318,"name":"Biological sciences/Chemical biology/Chemical ecology"}],"tags":[],"updatedAt":"2025-10-20T16:03:47+00:00","versionOfRecord":{"articleIdentity":"rs-6637201","link":"https://doi.org/10.1038/s41598-025-19855-1","journal":{"identity":"scientific-reports","isVorOnly":false,"title":"Scientific Reports"},"publishedOn":"2025-10-15 15:58:14","publishedOnDateReadable":"October 15th, 2025"},"versionCreatedAt":"2025-06-06 15:40:30","video":"","vorDoi":"10.1038/s41598-025-19855-1","vorDoiUrl":"https://doi.org/10.1038/s41598-025-19855-1","workflowStages":[]},"version":"v1","identity":"rs-6637201","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6637201","identity":"rs-6637201","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}