-
被称为第3代绿色农药的“昆虫性信息素”在害虫绿色防控中起着非常重要的作用,由于其生物活性高、专一性强、不产生耐药性、对天敌无害、无污染、有利于食品安全、使用简便等特点,人们已经逐渐意识到昆虫性信息素防治害虫的优势及重要性,使得昆虫性信息素技术得到迅速发展[1-3]。目前已鉴定出来的2 000多种昆虫性信息素组分中,以鳞翅目昆虫的最详细,大多数为不饱和脂肪族化合物,碳链长度一般为10~20碳,双键的构型为顺式(Z)、反式(E)、顺反式(Z/E)等,末端功能基多为乙酸酯、醇、醛类或者酮类,少数为丙酸酯或环氧化物,按其分子结构可分为末端有功能基TypeⅠ(夜蛾科、螟蛾科、卷蛾科、巢蛾科等22个科)、末端无功能基TypeⅡ(尺蛾科、毒蛾科、灯蛾科等5个科)和具有侧链的化合物[4-5]。
自1959年,第一个天然性信息素成分—蚕蛾醇(E10Z12-16:OH)被推测出其化学结构后,随之,《Science》和《Nature》便报道了“性信息素迷向干扰法防控害虫”的想法及概念,昆虫通过性信息素相互联络求偶交配,如果能干扰破坏雌雄间化学通讯交流系统,昆虫便不能交配和繁殖后代[6-9]。利用性信息素干扰防控害虫,即使用最佳的缓释装置,均匀速率将其释放至空气中,使空气中到处弥漫性信息素的气味,故而雄虫丧失对雌虫的定向行为能力,或使雄虫的触角长时间接触高浓度的性信息素而处于麻痹状态,失去对雌虫的性召唤反应能力,雌雄成虫无法正常交配,这种交配活动的减少将导致下一代的虫口密度降低,可作为一种有效的害虫防治措施。
昆虫性信息素的野外应用主要表现为:监测(Monitor)、大量诱捕(Mass trapping)和迷向干扰(Mating disruption),而“监测”和“大量诱捕”要求性信息素具有特定的比例和剂量,尤其对比例的要求更严格,然而“迷向干扰法防控害虫”对性信息素的比例并非严格,即按照雌虫本身释放性信息素的比例、或单一主要组分、或增大次要组分剂量(某些信息素主要组分价格昂贵),并结合合适的缓释装置,便可干扰雄虫对雌虫的交配定位[10-16],因此,寻找合适的信息素组分(比例)、较好的缓释载体及缓释剂型,改善性信息素的应用方式和缓释效果,是迷向干扰防控害虫的重要一环。目前,性信息素迷向干扰防控技术已在许多昆虫(鳞翅目为主)上取得重大突破,连续大面积使用3~5年,可达到最佳效果。本文陈述了国内外迷向技术的应用进展及影响迷向技术开展的生态因素,进而探讨不同缓释载体的剂型,旨在为利用昆虫性信息素迷向干扰害虫提供技术借鉴和参考。
-
近些年,昆虫性信息素迷向技术已在农业、林业、果树、蔬菜等多种害虫的防控中取得较大成功,自1970年开始,关于迷向干扰法防控害虫的文章、专利等其他科研成果也是逐年递增(图 1),其中,以鳞翅目(Lepidoptera)害虫为主,广泛应用于卷蛾科(Tortricidae)、螟蛾科(Pyralidae)、毒蛾科(Lymantriidae)等多种害虫的绿色防控中(表 1)[17];鞘翅目(Coleoptera)害虫也初步应用了迷向干扰技术进行防控[18-22];此外,同翅目(Homoptera)、半翅目(Hemiptera)、膜翅目(Hymenoptera)害虫也有了相关的性信息素迷向干扰技术报道[23-27]。目前,采用昆虫性信息素迷向法防控害虫应用面积最广的主要为舞毒蛾(Lymantria dispar)、苹果蠹蛾(Cydia pomonella)、葡萄花翅小卷蛾(Lobesia botrana)。自2002至2012年,性信息素迷向防控害虫的应用面积增长率极快,但也有部分害虫的应用情况呈下降趋势,如番茄麦茎蛾Keiferia lycopersicella的应用防控面积下降率达到了80%,目前全球采用性信息素迷向防控害虫的应用面积达到了756 000 hm2,较2002年的433 000 hm2提高了323 000 hm2(表 2)。
图 1 迷向干扰技术论文发表数量(左:基于www.pherobase.com;右:基于Google scholar)
Figure 1. The Number of publications about mating disruption. (Left: Based on the www.pherobase.com; Right: Based on Google scholar)
表 1 性信息素迷向法防控害虫的应用实例(鳞翅目)
Table 1. Application examples of pheromone mating disruption for controlling insects (Lepidoptera)
科Family 种类Species 学名Scientific name 透翅蛾科Sesiidae 3 Ichneumonoptera chrysophanes、Synanthedon tipuliformis、Vitacea polistiformis 麦蛾科Gelechiidae 5 Tuta absoluta、Scrobipalpopsis solanivora、Sitotroga cerealella、Pectinophora gossypiella、Keiferia lycopersicella 夜蛾科Noctuidae 5 Spodoptera exigua、Sesamia nonagrioides、Mamestra brassicae、Helicoverpa armigera、Trichoplusia ni 细蛾科Gracillariidae 3 Cameraria ohridella、Phyllocnistis citrella、Phyllonorycter ringoniella 卷蛾科Tortricidae 27 Cydia pomonella、Grapholita molesta、et al 螟蛾科Pyralidae 10 Chilo suppressalis、Plodia interpunctella、et al 菜蛾科Plutellidae 1 Plutella xylostella 巢蛾科Yponomeutidae 1 Prays oleae 潜蛾科Lyonetiidae 1 Leucoptera coffeella 毒蛾科Lymantriidae 3 Lymantria dispar、Orgyia pseudotsugata、Euproctis pseudoconspersa 草螟科Crambidae 1 Ostrinia nubilalis 尺蛾科Geometridae 1 Ascotis selenaria cretacea 表 2 性信息素迷向防控害虫的全球应用
Table 2. Worldwide use of mating disruption
昆虫种类Species 主要寄主Main host 区域Region 面积Aera/hm2 2002A 2012B 舞毒蛾 Lymantria dispar 森林 美国 60 000 200 000 苹果蠹蛾 Cydia pomonella 苹果、梨 全球 120 000 220 000 梨小食心虫 Grapholita molesta 梨、苹果、桃 全球 50 000 60 000 葡萄花翅小卷蛾Lobesia botrana 葡萄 欧洲、智利 41 000 150 000 环针单纹卷蛾 Eupoecilia ambiguella 葡萄 欧洲、智利 32 000 60 000 二化螟 Chilo suppressalis 水稻 西班牙 4 000 8 000 卷叶蛾 Leafroller moths 梨、苹果、桃、茶 美国、日本、澳大利亚、欧洲 24 000 15 000 棉红铃虫 Pectinophora gossypiella 棉花 美国、以色列、南美洲、欧洲 55 000 19 000 番茄麦茎蛾Keiferia lycopersicella 番茄 美国 10 000 2 000 小菜蛾 Plutella xylostella 甘蓝 美国 2 000 2 000 Synanthedon spp. 桃、杏、黑加仑 全球 5 000 6 000 Zeuzerina pyrina 梨、橄榄 全球 2 000 3 000 Endopiza viteana 葡萄 欧洲 1 000 1 000 其它种类Other species 水果、蔬菜 全球 27 000 10 000 注:A:基于[39];B:基于Shin-Etsu Corporation 自1869年,舞毒蛾被偶然在美国发现后,迅速在全美地区蔓延,到1970年,超过300多种的树木已被严重取食,传播速度极快,危害面积达7 500万英亩,因此,美国农业部林务局启动STS(Slow the Spread)项目,主要基于信息素迷向及诱捕技术,大大降低了舞毒蛾的传播速度,从每年13英里降低至3英里,约有1.5亿英亩的树木受到保护,具有显著的经济效益和生态效益[28-32];而在1991年,关于苹果蠹蛾信息素迷向干扰装置在美国登记后,100 g·hm-2浓度的信息素可将苹果蠹蛾的种群数量及发生动态控制在不足危害的水平,目前其性信息素年产量可达25 000 kg,防治面积达220 000 hm2,迷向干扰法防治苹果蠹蛾受到全球认可,成为苹果蠹蛾绿色防控技术的关键部分[33-35];葡萄花翅小卷蛾严重制约着欧洲葡萄产业的发展,1977年,法国开始使用迷向干扰技术防治葡萄花翅小卷蛾,但效果较差,主要由于其性信息素主要组分E7Z9-12:AC的化学合成技术遇到困难,化学纯度较低,完善并改进性信息素的化学合成技术后,E7Z9-12:AC按≈50~60 μg·h-1的释放速率可控制葡萄花翅小卷蛾的发生危害,仅在智利,迷向干扰法已应用于40 000 hm2的葡萄园,根除了新发生的种群数量[36-38]。
由于昆虫信息素的专一性,一种迷向剂只对靶标害虫有干扰效果,若对几种害虫同时危害的生态系统,制备每一害虫迷向剂的成本过高,因此,若在害虫发生区放置几种害虫信息素的复合迷向剂,进而对多种害虫产生迷向效果,具有显著的经济效益。如:针对在澳大利亚果园内经常混合发生的苹果蠹蛾和梨小食心虫(Grapholita molesta)的危害,一种载有苹果蠹蛾信息素E8E10-12:OH(215 mg)、12:OH(120 mg)、14:OH(27.5 mg)和梨小食心虫信息素Z8-12:Ac(62.44 mg)、E8-12:Ac(4.6 mg)、Z8-12:OH(0.7 mg)的混合迷向散发器(Isomate® CM/OFM TT,Shin-Etsu Chemical Co. Ltd,Japan),防治效果高于单种害虫独自使用,每公顷500个迷向装置,可取得显著的经济效益[40-41];装有Z11-14:Ac和E11-14:Ac(98:2)的迷向释放器可成功的干扰Choristoneura rosaceana和Pandemis limitata的交配[42];Judd等[43]也报道了装有E8E10-12:OH和Z11-14:Ac的散发器,每公顷布置500个,可有效的干扰苹果蠹蛾和4种卷叶蛾[43];此外, 复合迷向技术也在Argyrotaenia velutinana、Platynota flavedana、P. ideausalis等卷叶蛾上均取得了突破性的成功[44]; 但是,在野外诱捕试验(Trapping)中,复合诱芯有时却起到相反的作用,像梨小食心虫和苹果蠹蛾,载有2种昆虫信息素组分的诱芯,显著增加了梨小食心虫的诱捕数量,降低了苹果蠹蛾的诱捕量,这可能是2种昆虫均属卷蛾科,二者在种间信息素组分系统上存在相似之处(12C的共轭二烯醇、单烯醇、单烯乙酸酯),导致苹果蠹蛾的信息素组分影响梨小食心虫的诱捕量[45-46];而桃小食心虫(Carposina niponensis)(果蛀蛾科)和金纹细蛾(Lithocolletis ringoniella) (细蛾科)的单一诱芯均显著高于复合诱芯,相互之间不产生影响[47];因此,不同昆虫之间应根据自身进化、生理行为、寄主选择等,进而选用合适的野外诱捕技术。
如今,国外迷向技术已在应用面积及害虫种类上取得突破,而国内运用性信息素迷向技术仅仅发生在梨小食心虫、苹果蠹蛾、小菜蛾、桃小食心虫、茶毛虫(Euproctis pseudoconspersa)、甘蔗条螟(Chilo sacchariphagus)、亚洲玉米螟(Ostrinia furnacalis)、白杨透翅蛾(Parathrene tabaniformis)、棉红铃虫(Pectinophora gossypiella)、桃蛀螟(Dichocrocis punctiferalis)等[48-57],其中,迷向干扰防治技术稍加成熟的主要为梨小食心虫和苹果蠹蛾的防控;但自2008—2013年,采用迷向干扰法防控梨小食心虫的应用面积不到200 hm2 [58],尚未形成规范化的迷向防治,其应用面积较少。国内有关化学生态学方面的企业或公司也未见形成大批量生产的信息素迷向干扰产品,其迷向干扰产品主要以毛细管迷向丝为主,尚未报道其他较好的缓释载体。
当然,迷向干扰防控害虫的失败应用实例也存在,对于一些飞翔力较强的昆虫,迷向防控时效果往往较差,像Synanthedon pictipes、小菜蛾、Anomala orientalis等害虫的迷向干扰工作并未取得理想的效果,防治区的害虫种群密度或作物危害情况并未降低,这种现象主要由于交配雌虫比较方便的可以从周边其他栖息地飞至防治区[19, 59-60],接而进行生长繁殖,故迷向干扰技术的开展理应注意野外的各种生态因素,合理的开展害虫防控。
性信息素迷向干扰防控害虫的研究进展及应用前景
Research Progress and Application Prospect of Insect Sex Pheromone Mating Disruption
-
摘要:
目的 本文陈述国内外迷向技术的应用进展及影响迷向技术开展的生态因素,进而探讨不同缓释载体的剂型,旨在为利用昆虫性信息素迷向干扰害虫提供技术借鉴和参考。 方法 通过文献检索及个人研究方向,重点介绍国内外迷向干扰技术的应用情况及环境影响因素,探索常见缓释载体的类型(毛细管迷向丝、微胶囊、Puffer®、SPLAT®、蜡滴、空气纤维、静电纺丝/纳米纤维),指出不同缓释装置的优缺点。 结果 昆虫性信息素迷向干扰技术作为一种高效专一、环境友好的新型绿色防控害虫技术,已经成为害虫综合防控体系IPM(Integrated Pest Management)的重要组成部分,在全球范围内广泛应用于鳞翅目害虫的防治,鞘翅目、同翅目、半翅目、膜翅目等也有少量报道,尤以舞毒蛾、苹果蠹蛾、葡萄花翅小卷蛾的应用最为成功;明确了昆虫性信息素缓释载体及释放速率控制是有效实施性信息素迷向干扰技术的前提,揭示昆虫性信息素迷向干扰技术的应用机理,得出限制昆虫性信息素迷向干扰技术的经济及政策因素,旨在为我国更好的开展性信息素迷向干扰技术防控害虫提供参考。 结论 昆虫性信息素迷向干扰防控害虫的应用前景广泛,在农林业害虫的绿色防控中至关重要。 Abstract:Objective Application progress, ecological factor and controlling-release carrier types of insect sex pheromone mating disruption are discussed in depth, it could provide technical references for controlling insect pests with the mating disruption technology. Method The application, environmental factors, controlling-release carrier types (capillary, microcapsule, Puffer®, SPLAT®, paraffin, fiber, electrospun/nanofibers) and the mechanism of sex pheromone mating disruption at home and abroad were presented, in addition, the merits and demerits of different controlling-release carrier were indicated. Result Mating disruption of insect sex pheromone is a novel and green technology due to its high efficiency, specificity, environment-friendly and as an important part of Integrated Pest Management. And it has been widely used in Lepidoptera pests control, as well as Coleoptera, Homoptera, Hemiptera, Hymenoptera. Controlling-release carrier and rates are the precondition of sex pheromone mating disruption technology, and the application mechanism was revealed. The economic and policy factor restricting the application of mating disruption is further clarified, so as to provide references for controlling insect pests with sex pheromone mating disruption in China. Conclusion Mating disruption of insect sex pheromone is playing an important role in the pest control, and has a broad prospect. -
Key words:
- sex pheromone
- / mating disruption
- / controlling-release carrier
- / application
-
表 1 性信息素迷向法防控害虫的应用实例(鳞翅目)
Table 1. Application examples of pheromone mating disruption for controlling insects (Lepidoptera)
科Family 种类Species 学名Scientific name 透翅蛾科Sesiidae 3 Ichneumonoptera chrysophanes、Synanthedon tipuliformis、Vitacea polistiformis 麦蛾科Gelechiidae 5 Tuta absoluta、Scrobipalpopsis solanivora、Sitotroga cerealella、Pectinophora gossypiella、Keiferia lycopersicella 夜蛾科Noctuidae 5 Spodoptera exigua、Sesamia nonagrioides、Mamestra brassicae、Helicoverpa armigera、Trichoplusia ni 细蛾科Gracillariidae 3 Cameraria ohridella、Phyllocnistis citrella、Phyllonorycter ringoniella 卷蛾科Tortricidae 27 Cydia pomonella、Grapholita molesta、et al 螟蛾科Pyralidae 10 Chilo suppressalis、Plodia interpunctella、et al 菜蛾科Plutellidae 1 Plutella xylostella 巢蛾科Yponomeutidae 1 Prays oleae 潜蛾科Lyonetiidae 1 Leucoptera coffeella 毒蛾科Lymantriidae 3 Lymantria dispar、Orgyia pseudotsugata、Euproctis pseudoconspersa 草螟科Crambidae 1 Ostrinia nubilalis 尺蛾科Geometridae 1 Ascotis selenaria cretacea 表 2 性信息素迷向防控害虫的全球应用
Table 2. Worldwide use of mating disruption
昆虫种类Species 主要寄主Main host 区域Region 面积Aera/hm2 2002A 2012B 舞毒蛾 Lymantria dispar 森林 美国 60 000 200 000 苹果蠹蛾 Cydia pomonella 苹果、梨 全球 120 000 220 000 梨小食心虫 Grapholita molesta 梨、苹果、桃 全球 50 000 60 000 葡萄花翅小卷蛾Lobesia botrana 葡萄 欧洲、智利 41 000 150 000 环针单纹卷蛾 Eupoecilia ambiguella 葡萄 欧洲、智利 32 000 60 000 二化螟 Chilo suppressalis 水稻 西班牙 4 000 8 000 卷叶蛾 Leafroller moths 梨、苹果、桃、茶 美国、日本、澳大利亚、欧洲 24 000 15 000 棉红铃虫 Pectinophora gossypiella 棉花 美国、以色列、南美洲、欧洲 55 000 19 000 番茄麦茎蛾Keiferia lycopersicella 番茄 美国 10 000 2 000 小菜蛾 Plutella xylostella 甘蓝 美国 2 000 2 000 Synanthedon spp. 桃、杏、黑加仑 全球 5 000 6 000 Zeuzerina pyrina 梨、橄榄 全球 2 000 3 000 Endopiza viteana 葡萄 欧洲 1 000 1 000 其它种类Other species 水果、蔬菜 全球 27 000 10 000 注:A:基于[39];B:基于Shin-Etsu Corporation -
[1] 杜家伟.昆虫信息素及其应用[M].北京:中国林业出版社, 1988. [2] 孟宪佐.我国昆虫信息素研究与应用进展[J].昆虫知识, 2000, 37(2):75-84. doi: 10.3969/j.issn.0452-8255.2000.02.002 [3] 马涛, 温秀军, 李兴文.昆虫性信息素人工合成技术研究进展[J].世界林业研究, 2012, 25(6):46-50. [4] Ando T, Yamakawa R. Analyses of lepidopteran sex pheromones by mass spectrometry[J]. Trends in Analytical Chemistry, 2011, 30(7):990-1002. doi: 10.1016/j.trac.2011.03.010 [5] Ma T, Li Y Z, Sun Z H, et al. (Z, E)-9, 12-Tetradecadien-1-ol:a major sex pheromone component from Euzophera pyriella (Lepidoptera:Pyralididae) in Xinjiang, China[J]. Florida Entomologist, 2014, 97(2):496-503. doi: 10.1653/024.097.0221 [6] Babson A L. Eradicating the gypsy moth[J]. Science, 1963, 142(3591):447-448. [7] Burgess E D. Gypsy moth control[J]. Science, 1964, 143(3606):526. [8] Wright R H. After pesticides-what?[J]. Nature, 1964, 204(4954):121-125. doi: 10.1038/204121a0 [9] Gaston L K, Shorey H H, Saario C A. Insect population control by the use of sex pheromones to inhibit orientation between the sexes[J]. Nature, 1967, 213(5081):1155. [10] Suckling D M, Clearwater J R. Small scale trials of mating disruption of Epiphyas postvittana (Lepidoptera:Tortricidae)[J]. Environmental Entomology, 1990, 19(6):1702-1709. doi: 10.1093/ee/19.6.1702 [11] Trimble R M, Tyndall C A. Disruption of mating in the spotted tentiform leafminer (Lepidoptera:Gracillariidae) using synthetic sex pheromone[J]. Canadian Entomologist, 2000, 132(1):107-117. doi: 10.4039/Ent132107-1 [12] Mo J, Glover M, Munro S, et al. Evaluation of mating disruption for control of light brown apple moth (Lepidoptera:Tortricidae) in citrus[J]. Journal of Economic Entomology, 2006, 99(2):421-426. doi: 10.1093/jee/99.2.421 [13] Alfaro C, Navarro-Llopis V, Primo J. Optimization of pheromone dispenser density for managing the rice striped stem borer, Chilo suppressalis (Walker), by mating disruption[J]. Crop Protection, 2009, 28(7):547-628. doi: 10.1016/j.cropro.2009.03.011 [14] Witzgall P, Kirsch P, Cork A. Sex pheromone and their impact on pest management[J]. Journal of Chemical Ecology, 2010, 36(1):80-100. doi: 10.1007/s10886-009-9737-y [15] Samietz J, Baur R, Hillbur Y. Potential of synthetic sex pheromone blend for mating disruption of the swede midge, Contarinia nasturtii[J]. Journal of Chemical Ecology, 2012, 38(9):1171-1177. doi: 10.1007/s10886-012-0180-0 [16] Mori B A, Evenden M L. Challenges of mating disruption using aerosol-emitting pheromone puffers in red clover seed production fields to control Coleophora deauratella (Lepidoptera:Coleophoridae)[J]. Environmental Entomology, 2015, 44(1):34-43. doi: 10.1093/ee/nvu001 [17] Reddy G V, Guerrero A. New pheromones and insect control strategies[J]. Vitamins and Hormones, 2010, 83(3):493-519. [18] Koppenhöfer A K, Behle R W, Dunlap C, et al. Pellet Formulations of sex pheromone components for mating disruption of Oriental Beetle (Coleoptera:Scarabaeidae) in Turfgrass[J]. Environmental Entomology, 2008, 37(5):1126-1135. doi: 10.1603/0046-225X(2008)37[1126:PFOSPC]2.0.CO;2 [19] Behle R W, Cossé A A, Dunlap C, et al. Developing wax-based granule formulations for mating disruption of oriental beetles (Coleoptera:Scarabaeidae) in Turfgrass[J]. Journal of Economic Entomology, 2009, 101(6):1856-1863. [20] Rodriguez-Saona C, Polk D, Holdcraft R, et al. Splat-OrB Reveals competitive attraction as a mechanism of mating disruption in oriental beetle (Coleoptera:Scarabaeidae)[J]. Environmental Entomology, 2010, 39(6):1980-1989. doi: 10.1603/EN10062 [21] Arakaki N, Hokama Y, Nagayama A, et al. Mating disruption for control of the white grub beetle Dasylepida ishigakiensis (Coleoptera:Scarabaeidae) with synthetic sex pheromone in sugarcane fields[J]. Applied Entomology and Zoology, 2013, 48(4):441-446. doi: 10.1007/s13355-013-0202-6 [22] Mahroof R M, Phillips T W. Mating disruption of Lasioderma serricorne (Coleoptera:Anobiidae) in stored product habitats using the synthetic pheromone serricornin[J]. Journal of Applied Entomology, 2014, 138(5):378-386. doi: 10.1111/jen.2014.138.issue-5 [23] Bar Zakay I, Peleg B A, Hefetz A. Mating disruption of the California red scale Aonidiella aurantii[J]. Hassadeh, 1987, 69:1228-1231. [24] Moawad G M, El Hamaky M A, Gergis M F, et al. The impact of sex pheromones and insecticides on cotton piercing sucking pests in middle Egypt[J]. Bulletin of Entomological Society of Egypt, 1991, 19:231-236. [25] Martini A, Baldassari N, Baronio P, et al. Mating disruption of the pine sawfly Neodiprion sertifer (Hymenoptera:Diprionidae) in isolated pine stands[J]. Agricultural and Forest Entomology, 2002, 4:195-201. doi: 10.1046/j.1461-9563.2002.00143.x [26] Kakizaki M. The sex pheromone components for mating disruption of the rice leaf bug, Trigonotylus caelestialium (Heteroptera:Miridae)[J]. Applied Entomology and Zoology, 2004, 39(2):221-228. doi: 10.1303/aez.2004.221 [27] Fernandez D E, Beers E H, Brunner J F, et al. Horticultural mineral oil applications for apple powdery mildew and codling moth, Cydia pomonella (L.)[J]. Crop Protection, 2006, 25(6):585. doi: 10.1016/j.cropro.2005.08.014 [28] Sharov A A, Roberts E A, Liebhold A M, et al. Gypsy moth (Lepidoptera:Lymantriidae) spread in the central Appalachians:three methods for species boundary estimation[J]. Environmental Entomology, 1995, 24(6):1529-1538. doi: 10.1093/ee/24.6.1529 [29] Mayo J H, Straka T J, Leonard D S. The cost of slowing the spread of the gypsy moth (Lepidoptera:Lymantriidae)[J]. Journal of Economic Entomology, 2003, 96(5):1448-1454. doi: 10.1093/jee/96.5.1448 [30] Tcheslavskaia K S, Thorpe K, Brewster C, et al. Optimization of pheromone dosage for gypsy moth mating disruption[J]. Entomologia Experimentalis et Applicata, 2005, 115(3):355-361. doi: 10.1111/eea.2005.115.issue-3 [31] Thorpe K W, Tcheslavskaia K S, Tobin P C, et al. Persistent effects of aerial applications of disparlure on gypsy moth trap catch and mating success[J]. Entomologia Experimentalis et Applicata, 2007, 125(3):223-229. doi: 10.1111/eea.2007.125.issue-3 [32] Bigsby K M, Ambrose M J, Tobin P C, et al. The cost of gypsy moth sex in the city[J]. Urban Forestry and Urban Greening, 2014, 13(3):459-468. doi: 10.1016/j.ufug.2014.05.003 [33] Femenia-Ferreri B, Bosch D, Moya P, et al. Field assays of a new biodegradable controlled-release pheromone dispensers for mating disruption of Cydia pomonella (L.)[J]. IOBC WPRS Bulletins, 2007, 30:107-114. [34] Welter S, Cave F. Pheromone mating disruption of Cydia pomonella (L.) in California pears:Balancing dispenser emission rates and program performance[J]. IOBC WPRS Bulletins, 2007, 30:123-124. [35] Witzgall P, Stelinski L, Gut L, et al. Codling moth management and chemical ecology[J]. Annual Review of Entomology, 2008, 53(1):503-522. doi: 10.1146/annurev.ento.53.103106.093323 [36] Roehrich R, Carles J P, Tresor C. Essai pré-liminaire de protection du vignoble contre Lobesia botrana Schiff. au moyen de la phéromone sexuelle de synthèse (méthode de la confusion)[J]. Revue de Zoologie Agricole et de Pathologie Vegetale, 1977, 76:25-36. [37] Pasquier D, Charmillot PJ. Survey of pheromone emission from different kinds of dispensers used for mating disruption in orchards and vineyards[J]. IOBC WPRS Bulletins, 2005, 28:335-340. [38] Ioriatti C, Anfora G, Tasin M, et al. Chemical ecology and management of Lobesia botrana (Lepidoptera:Tortricidae)[J]. Journal of Economic Entomology, 2011, 104(4):1125-1137. doi: 10.1603/EC10443 [39] Cardé R T. Using pheromones to disrupt mating of moth pests, perspectives in ecological theory and integrated pest management[M].Cambridge:Cambridge University Press, 2007:122-169. [40] Il'ichev A L, Williams D G, Gut L J. Dual pheromone dispenser for combined control of codling moth Cydia pomonella L. and oriental fruit moth Grapholita molesta (Busck) (Lep., Tortricidae) in pears[J]. Journal of Applied Entomology, 2010, 131(5):368-376. [41] Bohnenblust E, Hull L A, Krawczyk G. A comparison of various mating disruption technologies for control of two internally feeding Lepidoptera in apples[J]. Entomologia Experimentalis et Applicata, 2011, 138(3):202-211. doi: 10.1111/eea.2011.138.issue-3 [42] Evenden M L, Judd G J R, Borden J H. Mating disruption of two sympatric, orchard-inhabiting tortricids, Choristoneura rosaceana and Pandemis limitata (Lepidoptera:Tortricidae) with pheromone components of both species' blends[J]. Journal of Economic Entomology, 1999, 92(2):380-390. doi: 10.1093/jee/92.2.380 [43] Judd G J R, Gardiner M G T. Simultaneous disruption of pheromone communication and mating in Cydia pomonella, Choristoneura rosaceana and Pandemis limitata (Lepidoptera:Tortricidae) using Isomate-CM/LR in apple orchards[J]. Journal of the Entomological Society of British Columbia, 2004, 101:3-13. [44] Pfeiffer D G, Kaakeh W, Killian J C, et al. Mating disruption to control damage by leafrollers in Virginia apple orchards[J]. Entomologia Experimentalis et Applicata, 2011, 67(1):47-56. [45] Knight A, Cichon L, Lago J, et al. Monitoring oriental fruit moth and codling moth (Lepidoptera:Tortricidae) with combinations of pheromones and kairomones[J]. Journal of Applied Entomology, 2015, 138(10):783-794. [46] 朱虹昱, 徐婧, 张润志.苹果蠹蛾性信息素对梨小食心虫的诱集和迷向作用[J].生物安全通报, 2015, 24(4):320-326. [47] 于海利, 张林林, 张国辉, 等.桃小食心虫与金纹细蛾不同性诱芯的诱蛾效果[J].西北农林科技大学学报, 2010, 38(10):121-125. [48] 涂洪涛, 张金勇, 陈汉杰, 等.应用性信息素缓释剂迷向防治桃树梨小食心虫研究[J].果树学报, 2012, 29(2):286-290. [49] 朱虹昱, 刘伟, 崔艮中, 等.苹果蠹蛾迷向防治技术效果初报[J].应用昆虫学报, 2012, 49(1):121-129. [50] 王香萍, 张钟宁.性诱剂迷向法防治高山甘蓝田小菜蛾研究[J].植物保护, 2008, 34(5):110-113. doi: 10.3969/j.issn.0529-1542.2008.05.025 [51] 李晓龙, 夏国宁, 何建川, 等.复合式膏体迷向剂对梨小、桃小食心虫的防控效果[J].植物保护, 2013, 39(6):147-152. doi: 10.3969/j.issn.0529-1542.2013.06.028 [52] 王永模, 戈峰, 刘向辉, 等.应用性信息素迷向法防治茶毛虫的田间试验[J].昆虫知识, 2006, 43(1):60-63. doi: 10.3969/j.issn.0452-8255.2006.01.014 [53] 张日火, 黄其军, 周行飞, 等.性诱剂迷向防治甘蔗条螟的应用[J].甘蔗糖业, 2002(1):13-14. doi: 10.3969/j.issn.1005-9695.2002.01.004 [54] 田畴, 金桂兰, 贺达汉, 等.亚洲玉米螟的迷向防治研究[J].宁夏农学院学报, 1995, 16(4):19-22. [55] 王华志, 余国和, 张万斌, 等.性信息素迷向防治白杨透翅蛾的研究[J].山东林业科技, 1993(4):56-59. [56] 蔡述宏.红铃虫性激素迷向技术之改进与设想[J].中国棉花, 1988(5):45-48. [57] 杨振亚, 宋其星, 吕金武, 等.板栗桃蛀螟性信息素迷向防治初探[J].落叶果树, 1986(2):45. [58] Cui G Z, Zhu J W. Pheromone-based pest management in China:past, present and future prospects[J]. Journal of Chemical Ecology, 2016, 42(7):557-570. doi: 10.1007/s10886-016-0731-x [59] Pfeiffer D G, Killian J C, Rajotte E G, et al. Mating disruption for reduction of damage by lesser peach tree borer (Lepidoptera:Sesiidae) in Virginia and Pennsylvania peach orchards[J]. Journal of Economic Entomology, 1991, 84(1):218-223. doi: 10.1093/jee/84.1.218 [60] Schroeder P C, Shelton A M, Ferguson C S, et al. Application of synthetic sex pheromone for management of diamondback moth, Plutella xylostella, in cabbage[J]. Entomologia Experimentalis et Applicata, 2010, 94(3):243-248. [61] Gut L J, Stelinski L L, Thomson D R, et al. Behavior modifying chemicals:prospects and constraints in IPM[M]. New York:CABI, 2004:73-121. [62] Nakano R, Takanashi T, Surlykke A. Moth hearing and sound communication[J]. Journal of Comparative Physiology A, 2015, 201(1):111-121. doi: 10.1007/s00359-014-0945-8 [63] Polajnar J, Eriksson A, Virant-Doberlet M, et al. Mating disruption of a grapevine pest using mechanical vibrations:from laboratory to the field[J]. Journal of Pest Science, 2016, 89(4):909-921. doi: 10.1007/s10340-015-0726-3 [64] Borchert D M, Walgenbach J F. Comparison of pheromone-mediated mating disruption and conventional insecticides for management of tufted apple bud moth (Lepidoptera:Tortricidae)[J]. Journal of Economic Entomology, 2000, 93(3):769-776. doi: 10.1603/0022-0493-93.3.769 [65] Brunner J, Welter S, Calkins C, et al. Mating disruption of codling moth:a perspective from the Western United States[J]. IOBC WPRS Bulletins, 2002, 25:1-11. [66] Cardé R T, Mafra-Neto A, Staten R T, et al. Evaluation of communication disruption in the pink bollworm in field wind tunnels[J]. Bulletin OILB/SROP, 1993, 16(10):23-28. [67] Cardé R T, Minks A K. Control of moth pests by mating disruption:successes and constraints[J]. Annual Review of Entomology, 1995, 40(1):559-585. doi: 10.1146/annurev.en.40.010195.003015 [68] Onufrieva K S, Thorpe K W, Hickman A D, et al. Gypsy moth mating disruption in open landscapes[J]. Agricultural and Forest Entomology, 2010, 10(3):175-179. [69] Cork A, De Souza K, Hall D R, et al. Development of PVC-resin-controlled release formulation for pheromones and use in mating disruption of yellow rice stem borer, Scirpophaga incertulas[J]. Crop Protection, 2008, 27(2):248-255. doi: 10.1016/j.cropro.2007.05.011 [70] Sartwell C, Daterman G E, Sower L L, et al. Mating disruption with synthetic sex attractant controls damage by Eucosma sonomana (Lepidoptera:Tortricidae:Oletreutinae) in Pinus ponderosa plantations. I. Manually applied polyvinyl chloride formulations[J]. Canadian Entomologist, 1980, 112(2):159-162. doi: 10.4039/Ent112159-2 [71] Felland C M, Hull L A, Barrett B A, et al. Small plot mating disruption trials for tufted apple bud moth, Platynota idaeusalis, in Pennsylvania apple orchards[J]. Entomologia Experimentalis et Applicata, 2011, 74(2):105-114. [72] Chamberlain D J, Brown N J, Jones O T, et al. Field evaluation of a slow release pheromone formulation to control the American bollworm, Helicoverpa armigera (Lepidoptera:Noctuidae) in Pakistan[J]. Bulletin of Entomological Research, 2000, 90(3):183-190. doi: 10.1017/S0007485300000304 [73] Grassi A, Zini M, Forno F, et al. Mating disruption field trials to control the currant clearwing moth, Synanthedon tipuliformis Clerck:a three year study[J]. Bulletin-OILB/SROP, 2002, 25:69-76. [74] Doane C C. Controlled-release devices for pheromones[M].New York:Marcel Dekker, 1999:295-317. [75] Trimble R M, Pree D J, Barszcz E S, et al. Comparison of a sprayable pheromone formulation and two hand-applied pheromone dispensers for use in the integrated control of Oriental fruit moth (Lepidoptera:Tortricidae)[J]. Journal of Economic Entomology, 2004, 97(2):482-489. doi: 10.1093/jee/97.2.482 [76] Il'ichev A L, Stelinski L L, Williams D G, et al. Sprayable microencapsulated sex pheromone formulation for mating disruption of oriental fruit moth (Lepidoptera:Tortricidae) in Australian peach and pear orchards[J]. Journal of Economic Entomology, 2006, 99(6):2048-2054. doi: 10.1093/jee/99.6.2048 [77] Waldstein D W, Gut L J. Effects of rain and sunlight on oriental fruit moth (Lepidoptera:Tortricidae) microcapsules applied to apple foliage[J]. Journal of Agricultural and Urban Entomology, 2004, 21(2):117-128. [78] Shorey H H, Gerber R G. Disruption of Pheromone Communication through the Use of Puffers for Control of Beet Armyworm (Lepidoptera:Noctuidae) in Tomatoes[J]. Environmental Entomology, 1996a, 25(6):1401-1405. doi: 10.1093/ee/25.6.1401 [79] Shorey H H, Gerber R G. Use of puffers for disruption of sex pheromone communication among navel orange worm moths (Lepidoptera:Pyralidae) in almonds, pistachios, and walnuts[J]. Environmental Entomology, 1996b, 25(5):1154-1157. doi: 10.1093/ee/25.5.1154 [80] Shorey H H, Sisk C B, Gerber R G. Widely separated pheromone release sites for disruption of sex pheromone communication in two species of Lepidoptera[J]. Environmental Entomology, 1996, 25(2):446-451. doi: 10.1093/ee/25.2.446 [81] Knight A L. Managing codling moth (Lepidoptera:Tortricidae) with an internal grid of either aerosol Puffers or dispenser clusters plus border applications of individual dispensers[J]. Journal of the Entomological Society of British Columbia, 2004, 101:69-77. [82] Shorey H H, Gerber R G. Use of puffers for disruption of sex pheromone communication of codling moths (Lepidoptera:Tortricidae) in walnut orchards[J]. Environmental Entomology, 1996c, 25(6):1398-1400. doi: 10.1093/ee/25.6.1398 [83] Stelinski L L, Gut L J, Haas M, et al. Evaluation of aerosol devices for simultaneous disruption of sex pheromone communication in Cydia pomonella and Grapholita molesta (Lepidoptera:Tortricidae)[J]. Journal of Pest Science, 2007, 80(4):225-233. doi: 10.1007/s10340-007-0176-7 [84] Steinmann K P, Zhang M H, Grant J A, et al. Pheromone-based pest management can be cost-effective for walnut growers[J]. California Agriculture, 2008, 62(3):105-110. doi: 10.3733/ca.v062n03p105 [85] Welter S C, Pickel C, Millar J, et al. Pheromone mating disruption offers selective management options for key pests[J]. California Agriculture, 2005, 59(1):16-22. doi: 10.3733/ca.v059n01p16 [86] Baldessari M, Rizzi C, Tolotti G, et al. Evaluation of an aerosol emitter for mating disruption of Cydia pomonella in Italy[J]. Communications in Agricultural and Applied Biological Science, 2013, 78(2):267-271. [87] Onufrieva K S, Thorpe K W, Hickman A D, et al. Effects of SPLAT® GM sprayable pheromone formulation on gypsy moth mating success[J]. Entomologia Experimentalis et Applicata, 2010, 136(2):109-115. doi: 10.1111/eea.2010.136.issue-2 [88] Onufrieva K S, Hickman A D, Leonard D S, et al. Efficacies and second-year effects of SPLAT GMTM and SPLAT GMTM organic formulations[J]. Insects, 2015, 6(1):1-12. [89] Baker T C, Francke W, Löfstedt C, et al. Isolation, identification and synthesis of sex pheromone components of the carob moth, Ectomyelois ceratoniae[J]. Tetrahedron Letters, 1989, 30(22):2901-2902. doi: 10.1016/S0040-4039(00)99153-6 [90] Todd J L, Millar J G, Vetter R S, et al. Behavioral and electrophysiological activity of (Z, E)-7, 9, 11-dodecatrienyl formate, a mimic of the major sex pheromone component of carob moth, Ectomyelois ceratoniae[J]. Journal of Chemical Ecology, 1992, 18(12):2331-2352. doi: 10.1007/BF00984953 [91] Atterholt C A, Delwiche M J, Rice R E, et al. Study of biopolymers and paraffin as potential controlled-release carriers for insect pheromones[J]. Journal of Agricultural and Food Chemistry, 1998, 46(10):4429-4434. doi: 10.1021/jf980642u [92] Stelinski L L, Miller J R, Ledebuhr R, et al. Mechanized applicator for large-scale field deployment of paraffin-wax dispensers of pheromone for mating disruption in tree fruit[J]. Journal of Economic Entomology, 2006, 99(5):1705-1710. doi: 10.1093/jee/99.5.1705 [93] Atterholt C A, Delwiche M J, Rice R E, et al. Controlled release of insect sex pheromones from paraffin wax and emulsions[J]. Journal of Control Release, 1999, 57(3):233-247. doi: 10.1016/S0168-3659(98)00119-9 [94] Weatherston I, Miller D, Lavoie-Dornik J. Commercial hollow-fiber pheromone formulations:The degrading effect of sunlight on celcon fibers causing increased release rates of the active ingredient[J]. Journal of Chemical Ecology, 1985, 11(12):1631-1644. doi: 10.1007/BF01012117 [95] Chronakis I S. Micro-/nano-fibers by electrospinning technology:processing, properties and applications[M]. Elsevier, 2015:513-548. [96] Chakraborty S, Liao I C, Adler A, et al. A facile technique to fabricate drug delivery systems[J]. Advanced Drug Delivery Reviews, 2009, 61(12):1043-1054. doi: 10.1016/j.addr.2009.07.013 [97] Kikionis S, Ioannou E, Konstantopoulou M, et al. Electrospun micro/nanofibers as controlled release systems for pheromones of Bactrocera oleae and Prays oleae[J]. Journal of Chemical Ecology, 2017, 43(3):254-262. doi: 10.1007/s10886-017-0831-2 [98] Ridgway R L, Silverstein R M, Inscoe M N. Behavior-modifying chemicals for insect management:applications of pheromones and other attractants[M].New York:Marcel Dekker, 1990. [99] Jones O T. Practical applications of pheromones and other semiochemicals[M]. Chapman and Hall, London, 1998, 261-355. [100] Yamanaka T. Mating disruption or mass trapping? Numerical simulation analysis of a control strategy for lepidopteran pests[J]. Population Ecology, 2007, 49(1):75-86. doi: 10.1007/s10144-006-0018-0 [101] Miller J R, Gut L J. Mating disruption for the 21st century matching technology with mechanism[J]. Environmental Entomology, 2015, 44(3):427-453. doi: 10.1093/ee/nvv052 [102] Bento J M S, Parra J R P, de Miranda S H G, et al. How much is a pheromone worth?[J]. F1000Research, 2016, 5:1763. doi: 10.12688/f1000research [103] Thomson D R, Gut L J, Jenkins J W. Pheromones for insect control[M].New Jersey:Humana Press Inc, 1998, 385-412. [104] Hathaway D O, Tamaki G, Moffitt H R, et al. Impact of removal of males with sex-pheromone-baited traps on suppression of the peach-twig borer, Anarsia lineatella (Zeller)[J]. Canadian Entomologist, 1985, 117(5):643-645. doi: 10.4039/Ent117643-5