昆虫嗅觉受体功能的研究进展

武韩,  易杰群,  刘键柏,  毛永凯,  成印洁,  胡文浩,  李继虎,  唐睿

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武韩, 易杰群, 刘键柏, 等. 昆虫嗅觉受体功能的研究进展 [J]. 环境昆虫学报, 2025, 47(1): 159-173. doi: 10.3969/j.issn.1674-0858.2025.01.17
引用本文: 武韩, 易杰群, 刘键柏, 等. 昆虫嗅觉受体功能的研究进展 [J]. 环境昆虫学报, 2025, 47(1): 159-173. doi: 10.3969/j.issn.1674-0858.2025.01.17
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WU Han, YI Jie-Qun, LIU Jian-Bai, et al. Recent advances in the study of insect olfactory receptor function [J]. Journal of Environmental Entomology, 2025, 47(1): 159-173. doi: 10.3969/j.issn.1674-0858.2025.01.17
Citation: WU Han, YI Jie-Qun, LIU Jian-Bai, et al. Recent advances in the study of insect olfactory receptor function [J]. Journal of Environmental Entomology, 2025, 47(1): 159-173. doi: 10.3969/j.issn.1674-0858.2025.01.17
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昆虫嗅觉受体功能的研究进展

doi: 10.3969/j.issn.1674-0858.2025.01.17
  • 1.

    广东省科学院南繁种业研究所, 广州 510316;

  • 2.

    广东省动物保护与资源利用重点实验室, 广东省野生动物保护与利用公共实验室, 广东省科学院动物研究所, 广州 510260

基金项目: 

国家自然科学基金 32172471;

广州市科技计划重点研发计划 2024B03J1272;

广东省植物保护新技术重点实验室开放基金 植重2021-05

详细信息
    作者简介:

    武韩，博士，副研究员，研究方向为昆虫的化学感受及生物防治，E-mail：antenna1217@163.com

    共同通讯作者 Author for correspondence:

    李继虎，高级农艺师，研究方向为害虫绿色防控，E-mail：ljhhz@126.com;

    唐睿，博士，副研究员，研究方向为昆虫神经行为学，E-mail：tangr@giz.gd.cn

  • 中图分类号: Q968.1

    文献标识码: A

    文章编号: 1674-0858(2025)01-0159-15

Recent advances in the study of insect olfactory receptor function

  • 1.

    Institute of Nanfan & Seed Industry, Guangdong Academy of Sciences, Guangzhou 510316, China;

  • 2.

    Guangdong Key Laboratory of Animal Conservation and Resource Utilization, Guangdong Public Laboratory of Wild Animal Conservation and Utilization, Institute of Zoology, Guangdong Academy of Sciences, Guangzhou 510260, China

    摘要
  • 摘要:

    嗅觉对昆虫的生存和繁殖至关重要。近年来对昆虫嗅觉识别的分子机制研究表明，嗅觉受体主要包括气味受体和离子型受体，它们在气味分子的识别过程中发挥关键和核心作用。为系统了解昆虫嗅觉受体功能的研究现状，本文综述了气味受体和离子型受体的发现历程、结构特征、表达定位，重点描述了气味受体和离子型受体的功能研究及其研究方法，力求为嗅觉受体研究和昆虫行为调控研究提供基础和参考。

     

    Abstract:

    Olfaction plays an important role in the survival and reproduction of insects. In recent years, extensive studies have been conducted to reveal the molecular mechanism of olfactory sensation. Two major types of olfactory receptors, including odorant receptors and ionotropic receptors, play a vital role in the process of odor reception. To get a comprehensive view of the function of olfactory receptors, we summarized the discovery process, structural features, expression, and localization of odorant receptors and ionotropic receptors in this review, with an emphasis on the function and research methods. This review aims to provide implications for future functional studies of olfactory receptors and ethological dissection of insect behavior.

     

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  • 嗅觉在昆虫求偶、寻找寄主和发现合适的产卵地点等行为上发挥举足轻重的作用。昆虫能够利用嗅觉对空气中不同的气味分子进行质和量的编码，进而产生求偶、定位寄主、选择产卵地点和躲避天敌的行为。昆虫对气味信息的识别是一个有序且复杂的过程。一般情况下，昆虫的外周嗅觉器官触角和下颚须上的嗅觉感受神经元（Olfactory sensory neurons，OSNs）首先对气味分子进行质和量的编码，此时气味信息由化学信号转变为电信号，以神经冲动的方式传递至昆虫的嗅觉初级中枢触角叶（Antennal lobe，AL），气味信息经触角叶整合后传递到嗅觉高级中枢蕈体（Mushroom body，MB）和侧角（Lateral horn，LH），经解码后产生行为输出（Wilson and Mainen，2006）。

    在昆虫嗅觉信号转导过程中，众多化学感受蛋白参与了气味信息的识别和感受过程。已知昆虫参与嗅觉认知的化学感受相关蛋白主要包括气味结合蛋白（Odorant binding protein，OBP），化学感受蛋白（Chemosensory protein，CSP），气味受体（Odorant receptor，OR），感觉神经元膜蛋白（Sensory neuron membrane protein，SNMP），离子型受体（Ionotropic receptor，IR）和气味降解酶（Odorant degrading enzyme，ODE）（Leal，2013）。

    OBP/CSP选择性结合气味分子，是昆虫感知外界气味信息的第一步，其主要作用是运输脂溶性的气味分子到达OSNs树突膜上的OR结合位点，同时与昆虫生长、发育、繁殖等生理功能及昆虫对杀虫剂的抗性相关（巩雪燕等，2023）。OR特异地识别一种或多种气味分子，直接触发OSN树突膜去极化产生动作电位，将化学信号转变成电信号，并经级联放大传导至神经系统，再经神经系统进一步加工、整合，最终引起昆虫的行为响应，故OR在昆虫对气味分子的识别过程中发挥关键和核心作用。另外，有一部分IR参与了嗅觉识别，而IR属于配体门控离子通道，在气味分子以配体形式结合IR后离子通道直接打开，引发动作电位和后续的神经传导，所以IR和OR一样在气味分子的识别过程中发挥关键作用（Fleischer et al.，2018；Wicher and Miazzi，2021）（图 1）。OR和IR统称为嗅觉受体，它们是昆虫感知外部环境气味及气味信号转导途径中的核心元件，在昆虫的生存和适应环境中扮演着至关重要的角色。近年来，随着生物信息学和结构生物学的飞速发展，昆虫嗅觉受体的功能研究方法也日益丰富，伴随着嗅觉受体研究的不断深入，我们对昆虫嗅觉受体的功能和机制有了更加深入的理解。

    图  1  昆虫的嗅觉受体
    注：A，气味受体（Odorant receptor，OR）；B，离子型受体（Ionotropic receptor，IR）（仿Fleischer et al.，2018；Wicher and Miazzi，2021）。
    Fig.  1  Olfactory receptor of insects
    下载: 全尺寸图片

    1.   嗅觉受体功能的研究方法

    嗅觉受体蛋白（包括OR和IR）的功能的研究方法分为体外（In vitro）和体内（In vivo）两类。体外研究主要借助于细胞系表达法，目前主要有HEK293细胞、昆虫Sf9细胞系和非洲爪蟾Xenopus oocytes卵母细胞表达系统，昆虫学研究中其中最常用且成熟的方法是爪蟾卵母细胞表达系统（Liu et al.，2013a；Jiang et al.，2014）。体内研究主要是借助于GAL4-UAS转基因果蝇空神经元系统、CRISPR-Cas9或者RNAi系统，其中利用转基因果蝇空神经元系统研究嗅觉受体功能较为普遍（Kurtovic et al.，2007；Guo et al.，2022a）。

    爪蟾卵母细胞表达系统是把靶标嗅觉受体基因表达到卵母细胞中，再利用双电极电压钳技术检测化合物引起的细胞膜内外的电流变化，从而阐明目标嗅觉受体-配体的结合模式，已在包括双翅目、鞘翅目、鳞翅目和膜翅目昆虫的嗅觉受体功能研究中得到了广泛应用（Wang et al.，2010；Jiang et al.，2014；Shan et al.，2019；Xiao et al.，2020）。爪蟾卵母细胞表达系统操作方便、效率高，适合大规模的初筛嗅觉受体的配体化合物，但此系统和昆虫真实环境下的生理系统偏离，在操作时配体化合物以溶液形式给予细胞刺激，因此研究结果可能在一定程度上存在假阳性（Fleischer et al.，2018）。

    转基因果蝇的空神经元系统是利用GAL4-UAS系统将靶标嗅觉受体基因表达在果蝇的特定感器内的神经元中，然后利用单感受器记录技术研究神经元的感受谱，从而明确靶标嗅觉受体基因的功能（Kurtovic et al.，2007；Wang et al.，2016a；Fleischer et al.，2018）。该系统异源表达靶标嗅觉受体于果蝇嗅觉感器内，且果蝇嗅觉感器内含有嗅觉信号转导所需要的其它辅助因子，因此一定程度上更接近于真实情况，后期还可以和果蝇的行为学实验结合，能够直观清晰地明确嗅觉受体的功能。

    2.   气味受体的功能研究进展

    1999年，在黑腹果蝇Drosophila melanogaster基因组全序列的基础上，根据受体蛋白的特征搜索所有的G蛋白偶联的受体家族，果蝇的OR基因率先被鉴定（Clyne et al.，1999；Gao and Chess，1999；Vosshall et al.，1999）。目前果蝇已鉴定的OR基因有60个，编码62个蛋白，形成了一个高度特化的受体家族，与线虫和脊椎动物或其它G蛋白偶联受体家族没有同源性。昆虫气味受体具有G蛋白经典的7次跨膜结构，但它所呈现的膜拓扑结构与哺乳动物气味受体刚好相反，即昆虫气味受体蛋白的N末端在胞内，C末端在胞外（图 1-A）（Benton et al.，2009）。昆虫的OR又可归为两类：一类是序列和功能高度演化的传统气味受体（Conventional odorant receptor），另一类是在不同种间高度保守的非典型性气味受体（Atypical odorant receptor）Or83b，即已知的昆虫气味受体共受体Orco。Orco在绝大多数昆虫物种的OSN中广泛表达，它不直接参与气味识别，而是与传统气味受体共表达，形成OR∶Orco = 1∶3的异聚体配体门控通道（Wang et al.，2024a；Zhao et al.，2024），进而辅助气味识别或者协助表达相同OR的OSN的轴突投射到同一神经纤维球。

    随着生物信息学的发展和转录组、基因组测序技术的进步，越来越多非模式昆虫物种的气味受体数量和基因序列得到了鉴定，不同昆虫的气味受体数量从无到几百个不等（Peñalva-Arana et al.，2009），这类数据积累为从进化上解析昆虫关键嗅觉表型的演化历程提供了宝贵参考（详细信息见网络版增强出版附表 1）。研究昆虫气味受体的功能，同样为发展害虫防治新思路、新方法和新途径提供理论基础（详细信息见网络版增强出版附表 2）。本世纪初，昆虫气味受体的功能研究主要集中于昆虫的性信息素受体。在模式昆虫黑腹果蝇中发现，雄性果蝇产生挥发性信息素cVA（cis-vacceny acetate），研究证实在雄性果蝇中cVA能通过激活表达OR67d的嗅觉感受神经元来促进雄性对雄性的攻击性（Kurtovic et al.，2007），OR67d也在雌性果蝇中的嗅觉感受神经元表达，但在雌性果蝇中OR67d的激活促进它们更易接受其它雄性，进一步研究表明cVA通过OR67d在雌雄果蝇中激活一个性别双态性的神经回路（Datta et al.，2008；Wang and Anderson，2010）。由性信息素介导的昆虫性别角色转变在原始鳞翅目小金蝠蛾Thitarodes xiaojinensis中发现了中间型，即雌雄成虫均积极找寻配偶的独特行为（Tang et al.，2024），推测此后通过气味受体的平行进化形成了经典的鳞翅目性信息素嗅觉调控模式。

    国内外的昆虫性信息素受体功能研究主要集中在鳞翅目昆虫中（Wanner et al.，2010；Fleischer and Krieger，2018；Bastin-Heline et al.，2019；Tian et al.，2021）。在昆虫中，触角是重要的嗅觉和味觉器官，果蝇和蛾类触角上的主要感受器主要包括三类，分别是毛型感器、锥形感器和腔锥感器（Zacharuk，1985；Steinbrecht，1996）。在棉铃虫Helicoverpa armigera、烟青虫Helicoverpa assulta和烟芽夜蛾Heliothis virescens的雄蛾中发现，毛型感器中高表达的OR13用来感受性信息素组分Z11-16:Ald，而感受其他组分Z9-14:Ald和Z9-16:Ald的气味受体在这3种物种中又有所不同，棉铃虫中OR14b却调谐其性腺微量组分Z9-14:Ald，而烟青虫中OR14b是调谐其主要性信息素组分Z9-16: ld，烟芽夜蛾中OR6用来调谐其主要性信息素组分Z9-14:Ald（Wang et al.，2011；Liu et al.，2013a；Jiang et al.，2014；Yang et al.，2017）。在甜菜夜蛾Spodoptera exigua中发现，气味受体OR13用来感受其主要性信息素组分Z9，E12-14:OAc，OR16用来感受第二信息素组分Z9-14: OH（Liu et al.，2013b）。近年入侵中国的外来物种草地贪夜蛾Spodoptera frugiperda的性信息素受体功能也得到了鉴定，发现该物种的OR13用来感受主要性性信息素组分Z9-14:OAc（Guo et al.，2022a）。

    性信息素受体在蛾类化学通讯中不仅介导同种识别，而且还介导异种间的生殖隔离。4种铃夜蛾属近缘种的比较研究表明，两对直系同源的性信息素受体OR14b和OR16在功能上发生了分化，在OR14b受体中，164和232位点直接参与配体结合，其突变导致了棉铃虫和美洲棉铃虫Helicoverpa zea OR14b的功能转变；而在OR16受体中，尽管66位点不直接参与配体结合，但其突变可能通过位阻效应调控底物识别，是导致4个近缘物种OR16功能分化的主要原因（Cao et al.，2023）。在铃夜蛾属物种中，OR11和OR13共同表达在雄蛾的A型感器，此现象非常高度保守。多年来众多研究已证实OR13属于性信息素受体用来感受雌蛾性腺分泌的Ⅰ类性信息素，近期的研究表明棉铃虫OR11用来感受雄蛾腹部分泌的Ⅱ类性信息素3，6，9-二十一碳三烯（3Z，6Z，9Z-21:H）（Wang et al.，2024b）。

    此外，在鞘翅目暗黑鳃金龟Holotrichia parallela中发现，雄性感知雌性释放信息素的能力以48 h为周期，而且发现雄性气味受体OR14用来感受L-异亮氨酸甲酯（L-isoleucine methyl），这在揭示鞘翅目性信息素受体功能上尚属首次（Wang et al.，2024c）。行为学实验证明十四醛（Tetradecanal，14:Ald）和2-十七烷酮（2-heptadecanone，2-Hep）是膜翅目昆虫棉铃虫齿唇姬蜂Campoletis chlorideae的两个性信息素组分，通过果蝇T1神经元表达和单感器记录、RNAi试验表明雄蜂触角中高表达的气味受体OR18和OR47分别用来感受14:Ald和2-Hep（Guo et al.，2022b）。

    近年来，昆虫感受植物挥发物的受体功能研究取得了很大进展。研究发现，苜蓿盲蝽Adelphocoris lineolatus中气味受体OR59参与感受水杨酸甲酯（Xiao et al.，2020），大灰优食蚜蝇Eupeodes corollae中气味受体OR25参与感受芳香物质丁香酚（Eugenol），对甲酚（p-cresol），和甲基丁香酚（Methyl eugenol）（Li et al.，2020b）。甜菜夜蛾的OR3和烟青虫的OR23用来感受法尼烯类化合物（Liu et al.，2014；Wu et al.，2019b），斜纹夜蛾Spodoptera litura的气味受体OR12专门用来感受顺-3-己烯乙酸酯（Zhang et al.，2013a）。在鳞翅目夜蛾中发现棉铃虫和海灰翅夜蛾Spodoptera littoralis触角上锥形感器表达的气味受体OR42专一性的感受花的常见挥发物苯乙醛，而且进化分析表明在鳞翅目昆虫中聚类在一支或相邻分支上的ORs功能具有相似性，大多数感受芳香族化合物的ORs较感受萜烯类和脂肪族化合物的ORs更早分化，且功能较为保守（Guo et al.，2021）。蔬菜害虫小菜蛾Plutella xylostella能够利用十字花科植物产生的次级代谢物异硫氰酸酯（Isothiocyanate）为嗅觉信号对寄主植物进行定位和产卵，通过电生理、行为学研究手段表明小菜蛾触角上的气味受体OR35和OR49介导了小菜蛾对十字花科的识别（Liu et al.，2020c），为小菜蛾的防治提供了新的视角。

    反-β-法尼烯（E-β-farnesene，EBF）被鉴定为绝大多数蚜虫的报警信息素组分，通过比较基因组学，结合嗅觉受体基因体外功能以及转基因果蝇的方法证实豌豆蚜Acyrthosiphon pisum气味受体OR5特异性地用于感受EBF（Zhang et al.，2017b），进一步研究表明豌豆蚜虫的天敌大灰优食蚜蝇借助于气味受体OR3感受EBF从而成功定位蚜虫（Wang et al.，2022b）。

    雌虫选择产卵地点时往往会避开同种雌虫已经产卵的地方，从而减少其后代之间的剧烈竞争。研究表明，长链脂肪酸甲酯（C16:0ME、C18:0ME和C18:1ME）是棉铃虫的卵表挥发物，在棉铃虫的产卵驱避中发挥重要作用，功能试验证实雌性棉铃虫的气味受体OR56介导了对这3种化合物的识别（Zhang et al.，2024）。一般认为，植食性昆虫利用触角内的嗅觉受体去感受植物信息从而找到寄主，然后对寄主进行定位和产卵。但研究发现烟草天蛾Manduca sexta的喙中也有气味受体表达，而且在定位烟草中发挥重要作用（Haverkamp et al.，2016）。在烟青虫中也发现，产卵器具有嗅觉感知功能，产卵器中高表达的OR31用来感受植物挥发物顺-3-己烯丁酸酯，行为实验揭示烟青虫偏好在含有顺-3-己烯丁酸酯的介质上产卵（Li et al.，2020c）。

    3.   离子型受体的功能研究进展

    Benton等（2009）在黑腹果蝇中发现了一类全新的化学感受受体，命名为离子型受体。通过对离子型受体的结构分析发现，IR属于谷氨酸受体家族，均包括胞外N端（N terminus）、配体结合域（Ligand-binding domain，LBD）、离子通道区域和胞内C端（C terminus）（图 1-B）；但和经典的离子型谷氨酸受体（Ionotropic glutamate receptors，iGluRs）不同，IRs存在着不同的配体结合域，缺少典型的谷氨酸作用残基，而且IR表达部位是在感受神经元的树突而不是在神经元的突触。借助于果蝇基因组和基因定位技术发现锥型感器和毛型感器内都表达有嗅觉感受必须的共受体Orco，说明锥型感器和毛型感器内的神经元参与了嗅觉感受，但在腔锥感器内除了有一类神经元表达OR35a/Orco外，其它的神经元表达大量IR受体，电生理实验表明腔锥感器内除表达OR35a/Orco的嗅觉神经元外，其它表达IR的神经元能够感受多种挥发性的胺类、酸类挥发物及苯乙醛，表明IR可作为一类新的嗅觉受体参与嗅觉信号的识别（Benton，2009）。在果蝇中的后续研究发现，IR还介导了对味觉、听觉、温度和湿度的识别功能（Giesen and Garrity，2017）（详见网络版增强出版附表 2）。

    与OR相比，IR的表达模式相对复杂。果蝇的腔锥感器中含有1~4个OSNs，其中单个OSN中能表达2~3个IR基因，果蝇的触角芒（Arista）和感受囊（Sacculus）也表达IR基因（Benton et al.，2009；Rytz et al.，2013）。另外，IR也广泛表达于果蝇的味觉感受器官，例如唇瓣、咽和足等部位（Hussain et al.，2016；Chen and Amrein，2017）。基因组分析表明果蝇含有66个IR基因，其中16个在触角上表达；在这16个IR基因中，10个在腔锥感器中的OSN中表达，4个在触角芒和感受囊中表达。除了在触角高表达外，IR在昆虫喙、下唇须、跗节等器官中也表达（Du et al.，2018；Tang et al.，2020；Liu et al.，2021b），暗含着IR可参与昆虫的多种感知功能。在果蝇中，Benton等根据氨基酸序列分析和基因表达模式的研究，可以将IRs分为3个亚家族：触角IRs（Antennal IRs），分化IRs（Divergent IRs）和共受体IRs（Co-receptor IRs）。

    在果蝇中，IR共受体的功能研究比较系统和深入。IR25a和IR8a基因在不同物种中相对保守，其在腔锥感器内、触角芒和感受囊内均广泛表达，研究证实其主要作用是作为共受体与其它IRs共同表达，从而行使各种功能（Benton et al.，2009；Abuin et al.，2011；Ai et al.，2013；Rytz et al.，2013；Tang et al.，2020）。例如IR8a和IR25a在果蝇感受酸类和多氨类化合物中是必须的；果蝇通过IR8a和IR84a感知苯乙醛和苯乙酸从而促进求偶行为（Grosjean et al.，2011）；IR8a和IR64a以及IR8a和IR75a/c/d参与了酸类化合物的嗅觉感受（Abuin et al.，2011；Ai et al.，2013）。IR76b和IR41a介导了果蝇长距离感受挥发性的多胺化合物（Hussain et al.，2016）。

    此外，IR共受体还介导了果蝇的味觉感受功能。例如共受体IR76b和IR25a介导了果蝇跗足味觉神经元对酸的感受，诱导果蝇产生产卵行为，光遗传学实验进一步证明激活IR76b和IR25a是该味觉神经元感受酸类物质的必要条件，此外IR76b还介导了果蝇对盐离子和钙离子的感受（Zhang et al.，2013b；Lee et al.，2018）；共受体IR20a介导了果蝇对糖和信息素的感受（Koh et al.，2014）。

    除了果蝇外，其它昆虫IR共受体的功能研究也在逐步开展。在中红侧沟茧蜂Microplitis mediator中发现，IR64a1和IR8a、IR64a2和IR8a表达在同一个嗅觉感器内但分属不同的嗅觉感受神经元，表达IR64a1和IR8a的神经元感受谱较广，对含有6~8个碳的短链醛、酸、醇和酯都有反应；表达IR64a2和IR8a的神经元则感受谱则较窄，仅对挥发性较弱的长链化合物的反应；另外，Z9-14:Ald是许多鳞翅目夜蛾科昆虫的性腺腺体组分（Arn et al.，1992），在中红侧沟茧蜂中发现，IR64a2和IR8a能够介导对Z9-14:Ald的感受（Shan et al.，2019），推测Z9-14:Ald是中红侧沟茧蜂寻找寄主的嗅觉信号。在烟草天蛾中发现，基因敲除IR8a后雌蛾对同种幼虫粪便挥发物中的抑卵信息素3-甲基戊酸和己酸的反应降低，行为实验进一步证实雌蛾产卵不再避开被幼虫取食过的叶片（Zhang et al.，2019b）。富含乙酸的糖醋液是许多夜蛾科昆虫的食诱剂，在东方粘虫Mythimna separata中发现，IR8a与IR64a，IR75q1和IR75q2在触角嗅觉感受神经元中具有共定位现象，在果蝇中IR8是感受酸类物质的必要受体，结合序列比及行为学实验推测东方粘虫的IR8a很可能是介导乙酸嗅觉系统的共受体之一（Tang et al.，2020）。在黄地老虎Agrotis segetum中发现，IR75p和IR75q基因家族发生了扩张，它们与共受体IR8a在毛型感器和锥型感器而非腔锥型感器中表达，功能研究表明IR75p1和IR75q1主要分别感受己酸和辛酸（Hou et al.，2022）。

    嗅觉受体除了OR和IR外，瞬时受体电位离子通道（Transient receptor potential，TRP）也可发挥嗅觉受体的作用来探测气味分子。TRP是一个位于细胞膜上的离子通道大家族，此类蛋白属于保守的痛觉受体，TRP已被证实在无脊椎动物和脊椎动物中广泛存在，包括扁虫、果蝇到人类中都发现有TRP的表达（Venkatachalam and Montell，2007）。TRP在生物体内扮演着调控各种生理与行为的重要角色，目前的研究表明TRP参与了温度感受、机械感受及化学感受（Fowler and Montell，2013），TRP在生物躲避危险中发挥重要作用。

    在果蝇中发现，具有驱避作用的香茅醛激活触角上的嗅觉信号通路的同时，激活了TRPA1通路，TRPA1和嗅觉通路一起参与了果蝇的驱避行为，果蝇中的TRPA1虽然没有直接参与识别气味分子，但是参与了嗅觉的传导。缺失TRPA1的果蝇对香茅醛不再表现出驱避行为。此外，在冈比亚按蚊Anopheles gambiae中发现，香茅醛可通过直接高效激活TRPA来实现其驱避行为（Kwon et al.，2010），随后的研究表明猫薄荷挥发物也能激活冈比亚按蚊中TRPA，从而介导驱避行为（Melo et al.，2021）。对蜜蜂的寄生螨狄斯瓦螨Varroa destructor的研究表明，植物源化合物香芹酚（Carvacrol）和α-松油醇（α-terpineol）通过激活狄斯瓦螨体内的TRPA1来实现对其驱避（Peng et al.，2015）。

    4.   展望

    昆虫行为调控技术是一种新的害虫防治策略，不同于常规的化学消杀防治方法，它利用行为调控剂，靶向害虫嗅觉介导的神经行为环路进行调控，有效实现对害虫的诱捕、驱避或者集成推拉策略形成生态隔离带，从而在保护作物的同时，减少了化学农药的使用，对于我国的粮食安全和农业可持续发展具有重要意义。在筛选昆虫行为调控剂时，除了常规的化学生态学方法外，利用反向化学生态学的方法研究昆虫的化学感受基因-嗅觉受体基因的功能来高通量筛选昆虫的潜在行为调控剂也是一种策略。同时，对重要嗅觉受体表达的干扰技术也具备对害虫、益虫实施靶向行为调控的应用潜力（郭丽娜等，2020；Chen et al.，2024）。

    对多种昆虫触角转录组数据研究发现，在昆虫的化学感受基因中，OR基因的数量一般高于OBP基因的数量，再加上部分行使嗅觉功能的IR基因，昆虫的嗅觉受体基因数量远高于OBP基因的数量。对昆虫而言，如果昆虫感受气味化合物需要OBP基因和嗅觉受体基因的共同参与，嗅觉受体基因的数量比OBP基因多，这就总体上决定了单个嗅觉受体识别气味分子的平均数量和OBP相比相对较少，即嗅觉受体的特异性更强。当然，昆虫性信息素结合蛋白（Pheromone binding protein，PBP）的特异性也很强，只结合少数的性信息素组分，但总体上其特异性仍不及相对应的昆虫信息素受体。虽然自然界的气味信息复杂多变，但是关键的气味化合物的嗅觉信息往往就能引发昆虫迅速产生先天行为反应。所以，我们在通过反向化学生态学来筛选昆虫的行为调控化合物时，一些特异表达或者高表达的气味受体基因就成为首选靶标，相对于OBP的低特异性和多个IR行使功能的复杂性，直接研究昆虫高特异性的气味受体功能可能会更快产生实际应用价值。此外，昆虫的TRP功能研究相对较少，鉴于TRP的功能和疼痛相关，TRP基因和功能在不同动物中又具有高度保守性，未来研究昆虫TRP的功能有可能快速筛选出昆虫的行为驱避剂。

    随着高通量技术的飞速发展，通过大规模获取基因组和转录组数据，研究人员可以系统性地比较不同昆虫气味受体基因家族、表达模式和功能，从而揭示其在不同昆虫物种中的功能多样性和演化路径，包括解析物理位置相关的受体基因其功能协同及所介导的跨物种表型演化（Li et al.，2023）。另外，对于昆虫嗅觉信号在大脑中枢传递模式的研究正不断深入，从果蝇触角叶中OR和IR的分离投射，到近期蝗虫触角叶中发现的环形编码模式（Jiang et al.，2024），为我们进一步理解昆虫嗅觉系统提供了宝贵参考，也为尝试归纳神经纤维球数量差异巨大的不同类群昆虫对气味感知的一般规律奠定了基础。

    随着结构预测精度的提高以及分子动力学模拟技术的进步，研究人员能够更迅速且准确地高通量筛选嗅觉受体的配体，进而揭示嗅觉受体的功能机制，这不仅可以深化对嗅觉受体功能的理解，而且对探究嗅觉受体的生物学作用及其潜在应用具有重要意义。目前，越来越多的昆虫嗅觉受体配体已得到确认，但这些配体的具体生态学功能仍需通过行为试验进一步验证，以便更有效地应用于昆虫行为调控和化学生态防治实践中。除了嗅觉外，触觉和视觉在昆虫感知外界信息的过程中也扮演着重要角色。这些感知系统相互作用，未来的研究应关注昆虫嗅觉系统与其他感知系统（如触觉、视觉）的交互作用，并探讨这些系统如何共同影响昆虫的行为和适应能力。

    附录：表 1 昆虫气味受体和离子型受体注释情况进展

    表 2 昆虫嗅觉受体功能研究进展

    详细数据见网络版增强出版附表（http://hjkcxb.alljournals.net/）

    附表  1  昆虫气味受体和离子型受体注释情况进展
    Appendix Table  1  Progress in the annotation of insect odorant receptors and ionotropic receptors
    目
    Order    		 科
    Family    		 物种
    Species    		 来源
    Source    		 气味受体
    OR    		 离子型受体
    IR    		 参考文献
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    鳞翅目Lepidoptera 尺蛾科Geometridae 槐尺蠖Semiothisa cinerearia 触角转录组Antennal transcriptome 52 23 Liu et al., 2020a
    鳞翅目Lepidoptera 麦蛾科Gelechiidae 番茄潜叶蛾Tuta absoluta 基因组Genome 58 44 Yin et al., 2021; Huang et al., 2024
    鳞翅目Lepidoptera 蛀果蛾科Carposinidae 桃蛀果蛾Carposina sasakii 触角转录组Antennal transcriptome 52 8 Tian et al., 2018
    鳞翅目Lepidoptera 灯蛾科Arctiidae 美国白蛾Hyphantria cunea 基因组Genome 47 44 Wu et al., 2019a; Yin et al., 2021
    鳞翅目Lepidoptera 裳蛾科Erebidae 舞毒蛾Lymantria dispar 基因组Genome; 触角转录组Antennal transcriptome 33 54 Clavijo McCormick et al., 2017; Yin et al., 2021
    鳞翅目Lepidoptera 粉蝶科Pieridae 菜粉蝶Pieris rapae 触角转录组Antennal transcriptome 60 34 Wang et al., 2023
    鳞翅目Lepidoptera 凤蝶科Papilionidae 柑橘凤蝶Papilio xuthus 基因组Genome 59 33 Yin et al., 2021; Yin et al., 2022
    鳞翅目Lepidoptera 凤蝶科Papilionidae 东方虎凤蝶Papilio glaucus 基因组Genome 61 34 Yin et al., 2021; Yin et al., 2022
    鳞翅目Lepidoptera 凤蝶科Papilionidae 金凤蝶Papilio machaon 基因组Genome 61 32 Yin et al., 2021; Yin et al., 2022
    鳞翅目Lepidoptera 凤蝶科Papilionidae 玉带凤蝶Papilio polytes 基因组Genome 67 37 Yin et al., 2021; Yin et al., 2022
    鳞翅目Lepidoptera 蛱蝶科Nymphalidae 黑脉金斑蝶Danaus plexippus 基因组Genome 64 32 Zhan et al., 2011; Engsontia et al., 2014; Yin et al., 2021
    鳞翅目Lepidoptera 蛱蝶科Nymphalidae 红带袖蝶Heliconius melpomene 基因组Genome 70 33 Yin et al., 2021
    鳞翅目Lepidoptera 枯叶蛾科Lasiocampidae 马尾松毛虫Dendrolimus punctatus 转录组Transcriptome 60 18 Zhang et al., 2017a
    鳞翅目Lepidoptera 木蠹蛾科Cossidae 沙棘木蠹蛾Eogystia hippophaecolus 触角转录组Antennal transcriptome 63 12 Hu et al.，2016
    鳞翅目Lepidoptera 夜蛾科Noctuidae 草地贪夜蛾Spodoptera frugiperda 基因组Genome 82 45 刘莹等，2019；Yin et al.，2021
    鳞翅目Lepidoptera 夜蛾科Noctuidae 甜菜夜蛾Spodoptera exigua 基因组Genome；触角转录组Antennal transcriptome 53 20 Du et al.，2018；Zhang et al.，2023a
    鳞翅目Lepidoptera 夜蛾科Noctuidae 海灰翅夜蛾Spodoptera littoralis 转录组Transcriptome 64 22 Koutroumpa et al.，2021
    鳞翅目Lepidoptera 夜蛾科Noctuidae 斜纹夜蛾Spodoptera litura 基因组Genome；触角转录组Antennal transcriptome 27 45 Zhu et al.，2018；Yang et al.，2024
    鳞翅目Lepidoptera 夜蛾科Noctuidae 东方粘虫Mythimna separata 触角转录组Antennal transcriptome 67 19 Tang et al., 2020
    鳞翅目Lepidoptera 夜蛾科Noctuidae 二点委夜蛾Athetis lepigone 触角转录组Antennal transcriptome 61 19 Zhang et al., 2016
    鳞翅目Lepidoptera 夜蛾科Noctuidae 疆夜蛾Peridroma saucia 触角转录组Antennal transcriptome 63 24 Sun et al., 2020
    鳞翅目Lepidoptera 夜蛾科Noctuidae 棉铃虫Helicoverpa armigera 基因组Genome; 触角转录组Antennal transcriptome 65 51 Zhang et al., 2015a; Liu et al., 2018a; Fan et al., 2022
    鳞翅目Lepidoptera 夜蛾科Noctuidae 烟青虫Helicoverpa assulta 触角转录组Antennal transcriptome 64 24 Xu et al., 2014; Zhang et al., 2015a
    鳞翅目Lepidoptera 夜蛾科Noctuidae 双委夜蛾Athetis dissimilis 触角转录组Antennal transcriptome 60 12 Dong et al., 2020
    鳞翅目Lepidoptera 夜蛾科Noctuidae 小地老虎Agrotis ipsilon 基因组Genome 86 39 Wang et al., 2021
    鳞翅目Lepidoptera 舟蛾科Notodontidae 仁扇舟蛾Clostera restitura 触角转录组Antennal transcriptome 78 15 Gu et al., 2019
    鞘翅目Coleoptera 吉丁科Buprestidae 白蜡窄吉丁Agrilus planipennis 基因组Genome 47 31 Andersson et al., 2019
    鞘翅目Coleoptera 吉丁科Buprestidae 花椒窄吉丁Agrilus zanthoxylum 触角转录组Antennal transcriptome 8 7 杨平等, 2019
    鞘翅目Coleoptera 象甲科Curculionidae 中欧山松大小蠹Dendroctonus ponderosae 基因组Genome 86 57 Andersson et al., 2019
    鞘翅目Coleoptera 象甲科Curculionidae 红脂大小蠹Dendroctonus valens 触角转录组Antennal transcriptome 22 3 Gu et al., 2015
    鞘翅目Coleoptera 象甲科Curculionidae 云南切梢小蠹Tomicus yunnanensis 转录组Transcriptome 9 3 Liu et al., 2018b
    鞘翅目Coleoptera 象甲科Curculionidae 红棕象甲Rhynchophorus ferrugineus 触角转录组Antennal transcriptome 76 10 Antony et al., 2016
    鞘翅目Coleoptera 象甲科Curculionidae 稻水象甲Lissorhoptrus oryzophilus 触角转录组Antennal transcriptome 41 10 Zhang et al., 2019a
    鞘翅目Coleoptera 象甲科Curculionidae 玉米象Sitophilus zeamais 触角转录组Antennal transcriptome 64 20 Tang et al., 2019; Chen et al., 2020
    鞘翅目Coleoptera 天牛科Cerambycidae 光肩星天牛Anoplophora glabripennis 转录组Transcriptome 132 59 Zhao et al., 2020b
    鞘翅目Coleoptera 天牛科Cerambycidae 管纹艳虎天牛Rhaphuma horsfieldi 转录组Transcriptome 84 20 Zhao et al., 2020b
    鞘翅目Coleoptera 天牛科Cerambycidae 灭字脊虎天牛Xylotrechus quadripes 触角转录组Antennal transcriptome 33 18 Pan et al., 2018
    鞘翅目Coleoptera 天牛科Cerambycidae 星天牛Anoplophora chinensis 触角转录组Antennal transcriptome 53 4 Sun et al., 2018
    鞘翅目Coleoptera 天牛科Cerambycidae 双条杉天牛Semanotus bifasciatus 触角转录组Antennal transcriptome 71 18 Li et al.，2022
    鞘翅目Coleoptera 拟步甲科Tenebrionidae 赤拟谷盗Tribolium castaneum 基因组Genome 341 23 Zhao et al.，2020b
    鞘翅目Coleoptera 金龟科Scarabaeidae 铜绿丽金龟Anomala corpulenta 触角转录组Antennal transcriptome 43 5 Li et al.，2015b
    鞘翅目Coleoptera 金龟科Scarabaeidae 华北大黑鳃金龟Holotrichia oblita 触角转录组Antennal transcriptome 44 9 Li et al.，2017b
    鞘翅目Coleoptera 金龟科Scarabaeidae 暗黑鳃金龟Holotrichia parallela 触角转录组Antennal transcriptome 47 27 Yi et al.，2018
    鞘翅目Coleoptera 金龟科Scarabaeidae 白星花金龟Protaetia brevitarsis 触角转录组Antennal transcriptome 72 8 Liu et al.，2019
    鞘翅目Coleoptera 叶甲科Chrysomelidae 大猿叶虫Colaphellus bowringi 触角转录组Antennal transcriptome 43 9 Li et al.，2015c
    鞘翅目Coleoptera 叶甲科Chrysomelidae 马铃薯甲虫Leptinotarsa decemlineata 触角转录组Antennal transcriptome 81 27 Cohen et al.，2024
    鞘翅目Coleoptera 叶甲科Chrysomelidae 紫榆叶甲Ambrostoma quadriimpressum 触角转录组Antennal transcriptome 34 20 Wang et al.，2016b
    鞘翅目Coleoptera 叶甲科Chrysomelidae 黄曲条跳甲Phyllotreta striolata 触角转录组Antennal transcriptome 73 49 Wu et al.，2016a
    鞘翅目Coleoptera 叶甲科Chrysomelidae 椰心叶甲Brontispa longissima 触角转录组Antennal transcriptome 48 19 Bin et al.，2017
    鞘翅目Coleoptera 叶甲科Chrysomelidae 茶角胸叶甲Basilepta melanopus 触角转录组Antennal transcriptome 63 18 Zhou et al.，2019a
    鞘翅目Coleoptera 叶甲科Chrysomelidae 绿豆象Callosobruchus chinensis 触角转录组Antennal transcriptome 116 1 郑海霞等，2018
    鞘翅目Coleoptera 三锥象科Brentidae 甘薯小象Cylas formicarius 基因组Genome 132 72 Hua et al.，2023
    鞘翅目Coleoptera 芫菁科Meloidae 眼斑沟芫菁Hycleus cichorii 基因组Genome 149 50 Wu et al.，2020
    鞘翅目Coleoptera 芫菁科Meloidae 大斑沟芫菁Hycleus phaleratus 基因组Genome 89 45 Wu et al.，2020
    半翅目Hemiptera 飞虱科Delphacidae 灰飞虱Laodelphax striatellus 基因组Genome 133 23 He et al.，2020
    半翅目Hemiptera 飞虱科Delphacidae 褐飞虱Nilaparvata lugens 基因组Genome 141 25 He et al.，2018
    半翅目Hemiptera 飞虱科Delphacidae 白背飞虱Sogatella furcifera 基因组Genome 135 16 He et al.，2018
    半翅目Hemiptera 蚜科Aphididae 大豆蚜Aphis glycines 基因组Genome 47 19 Robertson et al.，2019b
    半翅目Hemiptera 蚜科Aphididae 棉蚜Aphis gossypii 基因组Genome 34 23 Quan et al.，2019
    半翅目Hemiptera 蚜科Aphididae 豌豆蚜Acyrthosiphon pisum 基因组Genome 87 19 Robertson et al.，2019b
    半翅目Hemiptera 蚜科Aphididae 禾谷缢管蚜Rhopalosiphum padi 转录组Transcriptome 15 16 Kang et al.，2018
    半翅目Hemiptera 木虱科Psyllidae 柑橘木虱Diaphorina citri 触角转录组Antennal transcriptome 46 35 Wu et al.，2016b
    半翅目Hemiptera 木虱科Psyllidae 中国梨喀木虱Cacopsylla chinensis 转录组Transcriptome 7 4 Xu et al.，2019
    半翅目Hemiptera 盲蝽科Miridae 绿后丽盲蝽Apolygus lucorum 基因组Genome 135 33 Liu et al.，2021a
    半翅目Hemiptera 盲蝽科Miridae 苜蓿盲蝽Adelphocoris lineolatus 触角转录组Antennal transcriptome 88 12 Xiao et al.，2017
    半翅目Hemiptera 盲蝽科Miridae 黑肩绿盔盲蝽Cyrtorhinus lividipennis 触角转录组Antennal transcriptome 15 6 Wang et al.，2018
    半翅目Hemiptera 蝽科Pentatomidae 茶翅蝽Halyomorpha halys 基因组Genome 149 39 Sparks et al.，2020
    半翅目Hemiptera 荔蝽科Tessaratomidae 荔蝽Tessaratoma papillosa 触角转录组Antennal transcriptome 59 14 Wu et al.，2017
    半翅目Hemiptera 猎蝽科Reduviidae 普热猎蝽Rhodnius prolixus 基因组Genome 116 33 Lorenzo et al.，2024
    半翅目Hemiptera 蛛缘蝽科Alydidae 点蜂缘蝽Riptortus pedestris 基因组Genome 237 31 Liu et al.，2023
    蜚蠊目Blattodea 姬蜚蠊科Blattellidae 德国小蠊Blattella germanica 基因组Genome 134 897 Robertson et al.，2018a
    蜚蠊目Blattodea 蜚蠊科Blattidae 美洲大蠊Periplaneta americana 基因组Genome 58 233 Li et al.，2018；Zheng et al.，2022
    蜚蠊目Blattodea 原白蚁科Termopsidae 湿木白蚁Zootermopsis nevadensis 基因组Genome 61 141 Harrison et al.，2018
    蜚蠊目Blattodea 白蚁科Termitidae 纳塔尔大白蚁Macrotermes natalensis 基因组Genome 12 75 Harrison et al.，2018
    蜚蠊目Blattodea 木白蚁科Kalotermitidae Cryptotermes secundus 基因组Genome 54 135 Harrison et al.，2018
    膜翅目Hymenoptera 茧蜂科Braconidae 中红侧沟茧蜂Microplitis mediator 触角转录组Antennal transcriptome 169 17 Wang et al.，2017b；Wang et al.，2016c
    膜翅目Hymenoptera 茧蜂科Braconidae 菜蛾盘绒茧蜂Cotesia vestalis 触角转录组Antennal transcriptome 25 3 Liu et al.，2020b
    膜翅目Hymenoptera 茧蜂科Braconidae 腰带长体茧蜂Macrocentrus cingulum 触角转录组Antennal transcriptome 79 13 Ahmed et al.，2016
    膜翅目Hymenoptera 茧蜂科Braconidae 斑痣悬茧蜂Meteorus pulchricornis 触角转录组Antennal transcriptome 99 19 Sheng et al.，2017
    膜翅目Hymenoptera 茧蜂科Braconidae 阿维蚜茧蜂Aphidius ervi 基因组Genome 228 38 Dennis et al.，2020
    膜翅目Hymenoptera 茧蜂科Braconidae 烟蚜茧蜂Aphidius gifuensis 基因组Genome；转录组Transcriptome 80 25 Fan et al.，2018；Li et al.，2021a
    膜翅目Hymenoptera 茧蜂科Braconidae 豆柄瘤蚜茧蜂Lysiphlebus fabarum 基因组Genome 156 37 Dennis et al.，2020
    膜翅目Hymenoptera 赤眼蜂科Trichogrammatidae 螟黄赤眼蜂Trichogramma chilonis 转录组Transcriptome 45 14 Liu et al.，2018c
    膜翅目Hymenoptera 赤眼蜂科Trichogrammatidae 稻螟赤眼蜂Trichogramma japonicum 转录组Transcriptome 51 7 Li et al.，2021b
    膜翅目Hymenoptera 赤眼蜂科Trichogrammatidae 松毛虫赤眼蜂Trichogramma dendrolimi 基因组Genome 100 27 Zhang et al.，2023b
    膜翅目Hymenoptera 姬小蜂科Eulophidae 白蛾周氏啮小蜂Chouioia cunea 触角转录组Antennal transcriptome 80 10 Zhao et al.，2016
    膜翅目Hymenoptera 蜜蜂科Apidae 中华蜜蜂Apis cerana 基因组Genome 119 10 Park et al.，2015
    膜翅目Hymenoptera 蜜蜂科Apidae 意大利蜜蜂Apis mellifera 基因组Genome 163 21 Robertson and Wanner, 2006；Paoli and Galizia, 2021
    膜翅目Hymenoptera 金小蜂科Pteromalidae 丽蝇蛹集金小蜂Nasonia vitripennis 基因组Genome 301 111 Robertson et al.，2010；Harrison et al.，2018
    膜翅目Hymenoptera 茎蜂科Cephidae 麦茎蜂Cephus cinctus 基因组Genome 72 49 Robertson et al.，2018b
    膜翅目Hymenoptera 蚁科Formicidae 巨首芭切叶蚁Atta cephalotes 基因组Genome 376 18 Kock et al.，2013；Engsontia et al.，2015
    膜翅目Hymenoptera 蚁科Formicidae 佛罗里达弓背蚁Camponotus floridanus 转录组Transcriptome 407 31 Zhou et al.，2012
    膜翅目Hymenoptera 蚁科Formicidae 跳镰猛蚁Harpegnathos saltator 转录组Transcriptome 377 23 Zhou et al.，2012
    膜翅目Hymenoptera 蚁科Formicidae 阿根廷蚁Linepithema humile 基因组Genome 367 32 Smith et al.，2011
    膜翅目Hymenoptera 蚁科Formicidae 红胡须蚁Pogonomyrmex barbatus 基因组Genome 344 24 Smith et al.，2011
    膜翅目Hymenoptera 旋小蜂科Eupelmidae 荔蝽卵平腹小蜂Anastatus japonicus 触角转录组Antennal transcriptome 184 17 Zhan et al.，2023
    蜻蜓目Odonata 色蟌科Calopterygidae 华丽色蟌Calopteryx splendens 基因组Genome 5 21 Ioannidis et al.，2017
    直翅目Orthoptera 蝗科Acrididae 东亚飞蝗Locusta migratoria 基因组Genome；触角转录组Antennal
    transcriptome    		 142 32 Wang et al.，2015
    直翅目Orthoptera 蝗科Acrididae 青脊竹蝗Ceracris nigricornis 触角转录组Antennal transcriptome 71 8 Yuan et al.，2019
    直翅目Orthoptera 蝗科Acrididae 黄脊竹蝗Ceracris kiangsu 触角转录组Antennal transcriptome 91 13 Li et al.，2020a
    直翅目Orthoptera 蝗科Acrididae 亚洲小车蝗Oedaleus asiaticus 触角转录组Antennal transcriptome 60 6 Zhou et al.，2019b
    直翅目Orthoptera 蝗科Acrididae 中华稻蝗Oxya chinensis 触角转录组Antennal transcriptome 94 12 Cui et al.，2019
    附表  2  昆虫嗅觉受体功能研究进展
    Appendix Table  2  Progress in the functional deorphanization of insect olfactory receptors
    目Order 科Family 物种Species 受体(配体) Receptor (Ligand) 参考文献References
    双翅目Diptera 果蝇科Drosophilidae 黑腹果蝇Drosophila melanogaster OR67 (cis-vacceny acetate) Kurtovic et al.，2007
    双翅目Diptera 果蝇科Drosophilidae 黑腹果蝇Drosophila melanogaster IR75a+IR8a (acetic acid, propionic acid) Abuin et al.，2011
    双翅目Diptera 果蝇科Drosophilidae 黑腹果蝇Drosophila melanogaster IR84a+IR8a (phenylacetaldehyde) Abuin et al.，2011
    双翅目Diptera 果蝇科Drosophilidae 黑腹果蝇Drosophila melanogaster IR31a+IR8a (2-oxopentanoic acid) Silbering et al.，2011
    双翅目Diptera 果蝇科Drosophilidae 黑腹果蝇Drosophila melanogaster IR64a+IR8a (acetic acid) Ai et al.，2013
    双翅目Diptera 果蝇科Drosophilidae 黑腹果蝇Drosophila melanogaster IR92a (ammonia, amines) Min et al.，2013
    双翅目Diptera 果蝇科Drosophilidae 黑腹果蝇Drosophila melanogaster IR41a+IR76b (spermine, putrescine, 1,4-diamionbutane) Huassain et al.，2016
    双翅目Diptera 果蝇科Drosophilidae 黑腹果蝇Drosophila melanogaster IR25a+IR76b (acids) Chen and Amrein, 2017
    双翅目Diptera 果蝇科Drosophilidae 黑腹果蝇Drosophila melanogaster IR75b+IR75c+IR8a (butyric acid, propionic acid) Prieto-Godino et al.，2017
    双翅目Diptera 果蝇科Drosophilidae 大灰优食蚜蝇Eupeodes corollae OR25 (eugenol, p-cresol, methyl eugenol) Li et al.，2020b
    双翅目Diptera 果蝇科Drosophilidae 大灰优食蚜蝇Eupeodes corollae OR3 (E-β-farnesene) Wang et al.，2022b
    双翅目Diptera 蚊科Culicidae 冈比亚按蚊Anopheles gambiae IR41a,IR41c+IR76b+IR25a (amines) Pitts et al.，2017
    鳞翅目Lepidoptera 夜蛾科Noctuidae 棉铃虫Helicoverpa armigera OR6 (Z9-16:OH)；OR14b (Z9-14:Ald) Jiang et al.，2014
    鳞翅目Lepidoptera 夜蛾科Noctuidae 棉铃虫Helicoverpa armigera OR13 (Z11-16:Ald)；OR6 (Z9-16:Ald, Z9-14:Ald)；OR16 (Z11-16:OH) Liu et al.，2013a
    鳞翅目Lepidoptera 夜蛾科Noctuidae 棉铃虫Helicoverpa armigera OR42 (phenylacetaldehyde) Guo et al., 2021
    鳞翅目Lepidoptera 夜蛾科Noctuidae 棉铃虫Helicoverpa armigera OR13 (Z7-12:Ac) Sun et al., 2024
    鳞翅目Lepidoptera 夜蛾科Noctuidae 棉铃虫Helicoverpa armigera OR11 (3Z,6Z,9Z-21:H) Wang et al., 2024b
    鳞翅目Lepidoptera 夜蛾科Noctuidae 棉铃虫Helicoverpa armigera IR8a (acetic acid) Zhang et al., 2022a
    鳞翅目Lepidoptera 夜蛾科Noctuidae 烟青虫Helicoverpa assulta OR6 (Z9-16:OH); OR16 (Z9-14:Ald); Jiang et al., 2014
    鳞翅目Lepidoptera 夜蛾科Noctuidae 烟青虫Helicoverpa assulta OR23 (E-β-farnesene) Wu et al., 2019b
    鳞翅目Lepidoptera 夜蛾科Noctuidae 烟青虫Helicoverpa assulta OR14b (Z9-16:Ald) Yang et al., 2017
    鳞翅目Lepidoptera 夜蛾科Noctuidae 烟青虫Helicoverpa assulta OR13 (Z7-12:Ac) Sun et al., 2024
    鳞翅目Lepidoptera 夜蛾科Noctuidae 烟芽夜蛾Heliothis virescens OR13 (Z11-16:Ald) Grosse-Wilde et al., 2007
    鳞翅目Lepidoptera 夜蛾科Noctuidae 烟芽夜蛾Heliothis virescens OR6 (Z9-14:Ald); OR16 (Z11-16:OH) Wang et al., 2011
    鳞翅目Lepidoptera 夜蛾科Noctuidae 烟芽夜蛾Heliothis virescens OR13 (Z7-12:Ac) Sun et al., 2024
    鳞翅目Lepidoptera 夜蛾科Noctuidae 东方粘虫Mythimma separata OR3 (Z11-16:Ald); OR2 (Z9-14:Ald) Jiang et al., 2017
    鳞翅目Lepidoptera 夜蛾科Noctuidae 东方粘虫Mythimma separata IR8a (acetic acid) Tang et al., 2020
    鳞翅目Lepidoptera 夜蛾科Noctuidae 斜纹夜蛾Spodoptera litura OR12 (cis-3-hexenyl acetate); OR19 (4-ethylacetophenone) Zhang et al., 2013a
    鳞翅目Lepidoptera 夜蛾科Noctuidae 斜纹夜蛾Spodoptera litura OR51 (vanillin) Wei et al., 2023
    鳞翅目Lepidoptera 夜蛾科Noctuidae 斜纹夜蛾Spodoptera litura OR6 (Z9,E12-14:OAc); OR13 (Z9,E12-14:OAc, Z9-14:OAc); OR16 (Z9-14:OH) Zhang et al., 2015b
    鳞翅目Lepidoptera 夜蛾科Noctuidae 甜菜夜蛾Spodoptera exigua OR13 (Z9,E12-14:OAc, Z9-14:OAc); OR16 (Z9-14:OH) Liu et al., 2013b
    鳞翅目Lepidoptera 夜蛾科Noctuidae 甜菜夜蛾Spodoptera exigua OR3 (E-β-farnesene) Liu et al., 2014
    鳞翅目Lepidoptera 夜蛾科Noctuidae 美洲棉铃虫Helicoverpa zea OR14b (Z9-16:Ald, Z9-14:Ald); OR16 (Z11-16:OH) Cao et al., 2023
    鳞翅目Lepidoptera 夜蛾科Noctuidae 美洲棉铃虫Helicoverpa zea OR13 (Z11-16:Ald); OR6 (Z9-16:OH, Z9-14:Ald, Z9-16:Ald); Cao et al., 2023
    鳞翅目Lepidoptera 夜蛾科Noctuidae 阿根廷棉铃虫Helicoverpa gelotopoeon OR13 (Z11-16:Ald); OR14b (Z9-16:Ald, Z9-14:Ald); OR16 (Z9-14:Ald) Cao et al., 2023
    鳞翅目Lepidoptera 夜蛾科Noctuidae 阿根廷棉铃虫Helicoverpa gelotopoeon OR6 (Z9-16:OH, Z9-14:Ald, Z9-16:Ald); Cao et al., 2023
    鳞翅目Lepidoptera 夜蛾科Noctuidae 草地贪夜蛾Spodoptera frugiperda OR13 (Z9-14:Ac, Z9,E12-14:Ac); OR56 (Z7-12:Ac); OR62 (Z7-12:Ac) Guo et al., 2022a
    鳞翅目Lepidoptera 夜蛾科Noctuidae 草地贪夜蛾Spodoptera frugiperda OR6 (Z9,E12-14:Ac); OR16 (Z9-14:OH, Z9-14:Ald) Guo et al., 2022a
    鳞翅目Lepidoptera 夜蛾科Noctuidae 草地贪夜蛾Spodoptera frugiperda OR13 (Z7-12:Ac) Sun et al., 2024
    鳞翅目Lepidoptera 夜蛾科Noctuidae 草地贪夜蛾Spodoptera frugiperda IR75q.2 (nonanoic acid) Guo et al.，2023
    鳞翅目Lepidoptera 夜蛾科Noctuidae 海灰翅夜蛾Spodoptera littoralis OR6 (Z9,E12-14:OAc)； De Fouchier et al.，2017
    鳞翅目Lepidoptera 夜蛾科Noctuidae 黄地老虎Agrotis segetum IR75p.1 (hexanoic acid)；IR75q.1 (octanoic acid) Hou et al.，2022
    鳞翅目Lepidoptera 菜蛾科Plutellidae 小菜蛾Plutella xylostella OR6 (heptanal) Liu et al.，2024
    鳞翅目Lepidoptera 菜蛾科Plutellidae 小菜蛾Plutella xylostella OR11 (benzyl alcohol, salicylaldehyde, phenylacetaldehyde) Liu et al.，2022
    鳞翅目Lepidoptera 菜蛾科Plutellidae 小菜蛾Plutella xylostella OR41 (Z9-14:Ac, Z9-14:OH, Z9-14:Ald) Liu et al.，2018d
    鳞翅目Lepidoptera 蚕蛾科Bombycidae 家蚕Bombyx mori OR1 (bombykol) Sakurai et al.，2015
    鳞翅目Lepidoptera 蚕蛾科Bombycidae 家蚕Bombyx mori OR56 (cis-jasmone) Tanaka et al.，2009
    鳞翅目Lepidoptera 蚕蛾科Bombycidae 家蚕Bombyx mori OR19 (linalool) Anderson et al.，2009
    鳞翅目Lepidoptera 天蛾科Sphingidae 烟草天蛾Manduca sexta OR4 (Sex pheromone)；OR5 (linalool) Große-Wilde et al.，2010
    鳞翅目Lepidoptera 天蛾科Sphingidae 烟草天蛾Manduca sexta OR35 (α-copaene) Zhang et al.，2022b
    鳞翅目Lepidoptera 天蛾科Sphingidae 烟草天蛾Manduca sexta IR8a (hexanoic acid, 3-methylpentanoic) Zhang et al.，2019b
    鞘翅目Coleoptera 金龟科Scarabaeidae 暗黑鳃金龟Holotrichia parallela OR14 (L-isoleucine methyl) Wang et al.，2024c
    半翅目Hemiptera 盲蝽科Miridae 苜蓿盲蝽Adelphocoris lineolatus OR59 (menthyl salicylate) Xiao et al.，2020
    半翅目Hemiptera 蚜科Aphididae 绿盲蝽Apolygus lucorum OR47 (linalool) Zhang et al.，2022c
    半翅目Hemiptera 蚜科Aphididae 豌豆蚜Acyrthosiphon pisum OR23 (Green leaf volatiles) Huang et al.，2022
    半翅目Hemiptera 蚜科Aphididae 豌豆蚜Acyrthosiphon pisum OR5 (E-β-farnesene) Zhang et al.，2017b
    半翅目Hemiptera 叶蝉科Cicadellidae 茶小绿叶蝉Empoasca onukii IR25 (1-phenylethanol) Zhang et al.，2023c
    蜚蠊目Blattodea 蜚蠊科Blattidae 美洲大蠊Periplaneta americana OR53 (periplanone-A)；OR100 (periplanone-B) Li et al.，2024
    膜翅目Hymenoptera 姬蜂科Ichneumonidae 棉铃虫齿唇姬蜂Campoletis chlorideae OR18 (14:Ald)；OR47 (2-heptadecanone) Guo et al.，2022b

  • 图  1   昆虫的嗅觉受体

    注：A，气味受体（Odorant receptor，OR）；B，离子型受体（Ionotropic receptor，IR）（仿Fleischer et al.，2018；Wicher and Miazzi，2021）。

    Fig.  1   Olfactory receptor of insects

    下载: 全尺寸图片

    附表  1   昆虫气味受体和离子型受体注释情况进展

    Appendix Table  1   Progress in the annotation of insect odorant receptors and ionotropic receptors

    目
    Order    		 科
    Family    		 物种
    Species    		 来源
    Source    		 气味受体
    OR    		 离子型受体
    IR    		 参考文献
    References
    双翅目Diptera 果蝇科Drosophilidae 黑腹果蝇Drosophila melanogaster 基因组Genome 60 66 Benton et al., 2009; Rytz et al., 2013; Robertson, 2019a; 郭金梦等, 2020; 张夏瑄等, 2020
    双翅目Diptera 果蝇科Drosophilidae 斑翅果蝇Drosophila suzukii 基因组Genome 66 68 Crava et al., 2016; Ramasamy et al., 2016
    双翅目Diptera 实蝇科Tephritidae 桔小实蝇Bactrocera dorsalis 基因组Genome 104 86 Wang et al., 2022a
    双翅目Diptera 实蝇科Tephritidae 柑橘大实蝇Bactrocera minax 基因组Genome 59 59 Wang et al., 2022a
    双翅目Diptera 实蝇科Tephritidae 地中海实蝇Ceratitis capitata 基因组Genome 76 71 Papanicolaou et al., 2016
    双翅目Diptera 舌蝇科Glossinidae 奥斯汀舌蝇Glossina austeni 基因组Genome 40 28 Macharia et al., 2016
    双翅目Diptera 舌蝇科Glossinidae Glossina brevipalpis 基因组Genome 42 28 Macharia et al., 2016
    双翅目Diptera 舌蝇科Glossinidae Glossina fuscipes fuscipes 基因组Genome 42 31 Macharia et al., 2016
    双翅目Diptera 舌蝇科Glossinidae 刺舌蝇Glossina morsitans 基因组Genome 46 30 Macharia et al., 2016
    双翅目Diptera 舌蝇科Glossinidae 淡足舌蝇Glossina pallidipes 基因组Genome 42 30 Macharia et al., 2016
    双翅目Diptera 家蝇科Muscidae 家蝇Musca domestica 基因组Genome 86 110 Scott et al., 2014
    双翅目Diptera 丽蝇科Calliphoridae 幽暗丽蝇Calliphora stygia 触角转录组Antennal transcriptome 50 22 Leitch et al., 2015
    双翅目Diptera 食蚜蝇科Syrphidae 斜斑鼓额食蚜蝇Scaeva pyrastri 触角转录组Antennal transcriptome 38 16 Li et al., 2016
    双翅目Diptera 食蚜蝇科Syrphidae 大灰优食蚜蝇Eupeodes corollae 触角转录组Antennal transcriptome 42 23 Wang B et al., 2017a
    双翅目Diptera 食蚜蝇科Syrphidae 黑带食蚜蝇Episyrphus balteatus 触角转录组Antennal transcriptome 51 32 Wang B et al., 2017a
    双翅目Diptera 秆蝇科Chloropidae 稻秆潜蝇Chlorops oryzae 转录组Transcriptome 25 19 Qiu et al., 2018
    双翅目Diptera 蚊科Culicidae 中华按蚊Anopheles sinensis 基因组Genome; 转录组Transcriptome 59 35 Li et al., 2019; He et al., 2022
    双翅目Diptera 蚊科Culicidae 冈比亚按蚊Anopheles gambiae 基因组Genome 79 46 Pitts et al., 2017; He et al., 2022
    双翅目Diptera 蚊科Culicidae 埃及伊蚊Aedes aegypti 基因组Genome 117 135 Matthews et al., 2018
    双翅目Diptera 蚊科Culicidae 白纹伊蚊Aedes albopictus 基因组Genome 158 102 Chen et al., 2017; He et al., 2022
    双翅目Diptera 蚊科Culicidae 致倦库蚊Culex quinquefasciatus 基因组Genome 112 69 Croset et al., 2010; He et al., 2022
    双翅目Diptera 瘿蚊科Cecidomyiidae 黑森瘿蚊Mayetiola destructor 基因组Genome 122 39 Zhao et al., 2015
    双翅目Diptera 眼蕈蚊科Sciaridae 韭菜迟眼蕈蚊Bradysia odoriphaga 触角转录组Antennal transcriptome 71 18 Zhao et al., 2020a
    鳞翅目Lepidoptera 蝙蝠蛾科Hepialidae 剑川无钩蝠蛾Ahamus jianchuanensis 触角转录组Antennal transcriptome 10 7 Tang et al., 2024
    鳞翅目Lepidoptera 蝙蝠蛾科Hepialidae 虫草钩蝠蛾Thitarodes armoricanus 基因组Genome; 触角转录组Antennal transcriptome 16 32 Tang et al., 2024
    鳞翅目Lepidoptera 蝙蝠蛾科Hepialidae 小金蝠蛾Thitarodes xiaojinensis 基因组Genome; 触角转录组Antennal transcriptome 23 29 Tang et al., 2024
    鳞翅目Lepidoptera 菜蛾科Plutellidae 小菜蛾Plutella xylostella 基因组Genome; 触角转录组Antennal transcriptome 54 16 Yang et al., 2017
    鳞翅目Lepidoptera 蚕蛾科Bombycidae 家蚕Bombyx mori 基因组Genome 66 30 Yin et al., 2021; Morinaga et al., 2023
    鳞翅目Lepidoptera 天蛾科Sphingidae 烟草天蛾Manduca sexta 基因组Genome 73 34 Koenig et al., 2015; Yin et al., 2021
    鳞翅目Lepidoptera 卷蛾科Tortricidae 苹果蠹蛾Cydia pomonella 基因组Genome 85 43 Wan et al., 2019; Yin et al., 2021
    鳞翅目Lepidoptera 卷蛾科Tortricidae 梨小食心虫Grapholita molesta 触角转录组Antennal transcriptome 48 24 Li et al., 2015a
    鳞翅目Lepidoptera 卷蛾科Tortricidae 新西兰卷蛾Planotortrix excessana 触角转录组Antennal transcriptome 47 22 Steinwender et al., 2016; Grapputo et al., 2018
    鳞翅目Lepidoptera 螟蛾科Pyralidae 大蜡螟Galleria mellonella 基因组Genome; 触角转录组Antennal transcriptome 46 45 Zhao et al., 2019; Yin et al., 2021
    鳞翅目Lepidoptera 螟蛾科Pyralidae 稻纵卷叶螟Cnaphalocrocis medinalis 触角转录组Antennal transcriptome 29 15 Zeng et al., 2015
    鳞翅目Lepidoptera 螟蛾科Pyralidae 亚洲玉米螟Ostrinia furnacalis 触角转录组Antennal transcriptome 54 39 Yu et al., 2020; Yin et al., 2021
    鳞翅目Lepidoptera 螟蛾科Pyralidae 二化螟Chilo suppressalis 基因组Genome; 触角转录组Antennal transcriptome 47 36 Cao et al., 2014; Yin et al., 2021
    鳞翅目Lepidoptera 尺蛾科Geometridae 灰茶尺蠖Ectropis grisescens 触角转录组Antennal transcriptome 59 24 Li et al., 2017a
    鳞翅目Lepidoptera 尺蛾科Geometridae 槐尺蠖Semiothisa cinerearia 触角转录组Antennal transcriptome 52 23 Liu et al., 2020a
    鳞翅目Lepidoptera 麦蛾科Gelechiidae 番茄潜叶蛾Tuta absoluta 基因组Genome 58 44 Yin et al., 2021; Huang et al., 2024
    鳞翅目Lepidoptera 蛀果蛾科Carposinidae 桃蛀果蛾Carposina sasakii 触角转录组Antennal transcriptome 52 8 Tian et al., 2018
    鳞翅目Lepidoptera 灯蛾科Arctiidae 美国白蛾Hyphantria cunea 基因组Genome 47 44 Wu et al., 2019a; Yin et al., 2021
    鳞翅目Lepidoptera 裳蛾科Erebidae 舞毒蛾Lymantria dispar 基因组Genome; 触角转录组Antennal transcriptome 33 54 Clavijo McCormick et al., 2017; Yin et al., 2021
    鳞翅目Lepidoptera 粉蝶科Pieridae 菜粉蝶Pieris rapae 触角转录组Antennal transcriptome 60 34 Wang et al., 2023
    鳞翅目Lepidoptera 凤蝶科Papilionidae 柑橘凤蝶Papilio xuthus 基因组Genome 59 33 Yin et al., 2021; Yin et al., 2022
    鳞翅目Lepidoptera 凤蝶科Papilionidae 东方虎凤蝶Papilio glaucus 基因组Genome 61 34 Yin et al., 2021; Yin et al., 2022
    鳞翅目Lepidoptera 凤蝶科Papilionidae 金凤蝶Papilio machaon 基因组Genome 61 32 Yin et al., 2021; Yin et al., 2022
    鳞翅目Lepidoptera 凤蝶科Papilionidae 玉带凤蝶Papilio polytes 基因组Genome 67 37 Yin et al., 2021; Yin et al., 2022
    鳞翅目Lepidoptera 蛱蝶科Nymphalidae 黑脉金斑蝶Danaus plexippus 基因组Genome 64 32 Zhan et al., 2011; Engsontia et al., 2014; Yin et al., 2021
    鳞翅目Lepidoptera 蛱蝶科Nymphalidae 红带袖蝶Heliconius melpomene 基因组Genome 70 33 Yin et al., 2021
    鳞翅目Lepidoptera 枯叶蛾科Lasiocampidae 马尾松毛虫Dendrolimus punctatus 转录组Transcriptome 60 18 Zhang et al., 2017a
    鳞翅目Lepidoptera 木蠹蛾科Cossidae 沙棘木蠹蛾Eogystia hippophaecolus 触角转录组Antennal transcriptome 63 12 Hu et al.，2016
    鳞翅目Lepidoptera 夜蛾科Noctuidae 草地贪夜蛾Spodoptera frugiperda 基因组Genome 82 45 刘莹等，2019；Yin et al.，2021
    鳞翅目Lepidoptera 夜蛾科Noctuidae 甜菜夜蛾Spodoptera exigua 基因组Genome；触角转录组Antennal transcriptome 53 20 Du et al.，2018；Zhang et al.，2023a
    鳞翅目Lepidoptera 夜蛾科Noctuidae 海灰翅夜蛾Spodoptera littoralis 转录组Transcriptome 64 22 Koutroumpa et al.，2021
    鳞翅目Lepidoptera 夜蛾科Noctuidae 斜纹夜蛾Spodoptera litura 基因组Genome；触角转录组Antennal transcriptome 27 45 Zhu et al.，2018；Yang et al.，2024
    鳞翅目Lepidoptera 夜蛾科Noctuidae 东方粘虫Mythimna separata 触角转录组Antennal transcriptome 67 19 Tang et al., 2020
    鳞翅目Lepidoptera 夜蛾科Noctuidae 二点委夜蛾Athetis lepigone 触角转录组Antennal transcriptome 61 19 Zhang et al., 2016
    鳞翅目Lepidoptera 夜蛾科Noctuidae 疆夜蛾Peridroma saucia 触角转录组Antennal transcriptome 63 24 Sun et al., 2020
    鳞翅目Lepidoptera 夜蛾科Noctuidae 棉铃虫Helicoverpa armigera 基因组Genome; 触角转录组Antennal transcriptome 65 51 Zhang et al., 2015a; Liu et al., 2018a; Fan et al., 2022
    鳞翅目Lepidoptera 夜蛾科Noctuidae 烟青虫Helicoverpa assulta 触角转录组Antennal transcriptome 64 24 Xu et al., 2014; Zhang et al., 2015a
    鳞翅目Lepidoptera 夜蛾科Noctuidae 双委夜蛾Athetis dissimilis 触角转录组Antennal transcriptome 60 12 Dong et al., 2020
    鳞翅目Lepidoptera 夜蛾科Noctuidae 小地老虎Agrotis ipsilon 基因组Genome 86 39 Wang et al., 2021
    鳞翅目Lepidoptera 舟蛾科Notodontidae 仁扇舟蛾Clostera restitura 触角转录组Antennal transcriptome 78 15 Gu et al., 2019
    鞘翅目Coleoptera 吉丁科Buprestidae 白蜡窄吉丁Agrilus planipennis 基因组Genome 47 31 Andersson et al., 2019
    鞘翅目Coleoptera 吉丁科Buprestidae 花椒窄吉丁Agrilus zanthoxylum 触角转录组Antennal transcriptome 8 7 杨平等, 2019
    鞘翅目Coleoptera 象甲科Curculionidae 中欧山松大小蠹Dendroctonus ponderosae 基因组Genome 86 57 Andersson et al., 2019
    鞘翅目Coleoptera 象甲科Curculionidae 红脂大小蠹Dendroctonus valens 触角转录组Antennal transcriptome 22 3 Gu et al., 2015
    鞘翅目Coleoptera 象甲科Curculionidae 云南切梢小蠹Tomicus yunnanensis 转录组Transcriptome 9 3 Liu et al., 2018b
    鞘翅目Coleoptera 象甲科Curculionidae 红棕象甲Rhynchophorus ferrugineus 触角转录组Antennal transcriptome 76 10 Antony et al., 2016
    鞘翅目Coleoptera 象甲科Curculionidae 稻水象甲Lissorhoptrus oryzophilus 触角转录组Antennal transcriptome 41 10 Zhang et al., 2019a
    鞘翅目Coleoptera 象甲科Curculionidae 玉米象Sitophilus zeamais 触角转录组Antennal transcriptome 64 20 Tang et al., 2019; Chen et al., 2020
    鞘翅目Coleoptera 天牛科Cerambycidae 光肩星天牛Anoplophora glabripennis 转录组Transcriptome 132 59 Zhao et al., 2020b
    鞘翅目Coleoptera 天牛科Cerambycidae 管纹艳虎天牛Rhaphuma horsfieldi 转录组Transcriptome 84 20 Zhao et al., 2020b
    鞘翅目Coleoptera 天牛科Cerambycidae 灭字脊虎天牛Xylotrechus quadripes 触角转录组Antennal transcriptome 33 18 Pan et al., 2018
    鞘翅目Coleoptera 天牛科Cerambycidae 星天牛Anoplophora chinensis 触角转录组Antennal transcriptome 53 4 Sun et al., 2018
    鞘翅目Coleoptera 天牛科Cerambycidae 双条杉天牛Semanotus bifasciatus 触角转录组Antennal transcriptome 71 18 Li et al.，2022
    鞘翅目Coleoptera 拟步甲科Tenebrionidae 赤拟谷盗Tribolium castaneum 基因组Genome 341 23 Zhao et al.，2020b
    鞘翅目Coleoptera 金龟科Scarabaeidae 铜绿丽金龟Anomala corpulenta 触角转录组Antennal transcriptome 43 5 Li et al.，2015b
    鞘翅目Coleoptera 金龟科Scarabaeidae 华北大黑鳃金龟Holotrichia oblita 触角转录组Antennal transcriptome 44 9 Li et al.，2017b
    鞘翅目Coleoptera 金龟科Scarabaeidae 暗黑鳃金龟Holotrichia parallela 触角转录组Antennal transcriptome 47 27 Yi et al.，2018
    鞘翅目Coleoptera 金龟科Scarabaeidae 白星花金龟Protaetia brevitarsis 触角转录组Antennal transcriptome 72 8 Liu et al.，2019
    鞘翅目Coleoptera 叶甲科Chrysomelidae 大猿叶虫Colaphellus bowringi 触角转录组Antennal transcriptome 43 9 Li et al.，2015c
    鞘翅目Coleoptera 叶甲科Chrysomelidae 马铃薯甲虫Leptinotarsa decemlineata 触角转录组Antennal transcriptome 81 27 Cohen et al.，2024
    鞘翅目Coleoptera 叶甲科Chrysomelidae 紫榆叶甲Ambrostoma quadriimpressum 触角转录组Antennal transcriptome 34 20 Wang et al.，2016b
    鞘翅目Coleoptera 叶甲科Chrysomelidae 黄曲条跳甲Phyllotreta striolata 触角转录组Antennal transcriptome 73 49 Wu et al.，2016a
    鞘翅目Coleoptera 叶甲科Chrysomelidae 椰心叶甲Brontispa longissima 触角转录组Antennal transcriptome 48 19 Bin et al.，2017
    鞘翅目Coleoptera 叶甲科Chrysomelidae 茶角胸叶甲Basilepta melanopus 触角转录组Antennal transcriptome 63 18 Zhou et al.，2019a
    鞘翅目Coleoptera 叶甲科Chrysomelidae 绿豆象Callosobruchus chinensis 触角转录组Antennal transcriptome 116 1 郑海霞等，2018
    鞘翅目Coleoptera 三锥象科Brentidae 甘薯小象Cylas formicarius 基因组Genome 132 72 Hua et al.，2023
    鞘翅目Coleoptera 芫菁科Meloidae 眼斑沟芫菁Hycleus cichorii 基因组Genome 149 50 Wu et al.，2020
    鞘翅目Coleoptera 芫菁科Meloidae 大斑沟芫菁Hycleus phaleratus 基因组Genome 89 45 Wu et al.，2020
    半翅目Hemiptera 飞虱科Delphacidae 灰飞虱Laodelphax striatellus 基因组Genome 133 23 He et al.，2020
    半翅目Hemiptera 飞虱科Delphacidae 褐飞虱Nilaparvata lugens 基因组Genome 141 25 He et al.，2018
    半翅目Hemiptera 飞虱科Delphacidae 白背飞虱Sogatella furcifera 基因组Genome 135 16 He et al.，2018
    半翅目Hemiptera 蚜科Aphididae 大豆蚜Aphis glycines 基因组Genome 47 19 Robertson et al.，2019b
    半翅目Hemiptera 蚜科Aphididae 棉蚜Aphis gossypii 基因组Genome 34 23 Quan et al.，2019
    半翅目Hemiptera 蚜科Aphididae 豌豆蚜Acyrthosiphon pisum 基因组Genome 87 19 Robertson et al.，2019b
    半翅目Hemiptera 蚜科Aphididae 禾谷缢管蚜Rhopalosiphum padi 转录组Transcriptome 15 16 Kang et al.，2018
    半翅目Hemiptera 木虱科Psyllidae 柑橘木虱Diaphorina citri 触角转录组Antennal transcriptome 46 35 Wu et al.，2016b
    半翅目Hemiptera 木虱科Psyllidae 中国梨喀木虱Cacopsylla chinensis 转录组Transcriptome 7 4 Xu et al.，2019
    半翅目Hemiptera 盲蝽科Miridae 绿后丽盲蝽Apolygus lucorum 基因组Genome 135 33 Liu et al.，2021a
    半翅目Hemiptera 盲蝽科Miridae 苜蓿盲蝽Adelphocoris lineolatus 触角转录组Antennal transcriptome 88 12 Xiao et al.，2017
    半翅目Hemiptera 盲蝽科Miridae 黑肩绿盔盲蝽Cyrtorhinus lividipennis 触角转录组Antennal transcriptome 15 6 Wang et al.，2018
    半翅目Hemiptera 蝽科Pentatomidae 茶翅蝽Halyomorpha halys 基因组Genome 149 39 Sparks et al.，2020
    半翅目Hemiptera 荔蝽科Tessaratomidae 荔蝽Tessaratoma papillosa 触角转录组Antennal transcriptome 59 14 Wu et al.，2017
    半翅目Hemiptera 猎蝽科Reduviidae 普热猎蝽Rhodnius prolixus 基因组Genome 116 33 Lorenzo et al.，2024
    半翅目Hemiptera 蛛缘蝽科Alydidae 点蜂缘蝽Riptortus pedestris 基因组Genome 237 31 Liu et al.，2023
    蜚蠊目Blattodea 姬蜚蠊科Blattellidae 德国小蠊Blattella germanica 基因组Genome 134 897 Robertson et al.，2018a
    蜚蠊目Blattodea 蜚蠊科Blattidae 美洲大蠊Periplaneta americana 基因组Genome 58 233 Li et al.，2018；Zheng et al.，2022
    蜚蠊目Blattodea 原白蚁科Termopsidae 湿木白蚁Zootermopsis nevadensis 基因组Genome 61 141 Harrison et al.，2018
    蜚蠊目Blattodea 白蚁科Termitidae 纳塔尔大白蚁Macrotermes natalensis 基因组Genome 12 75 Harrison et al.，2018
    蜚蠊目Blattodea 木白蚁科Kalotermitidae Cryptotermes secundus 基因组Genome 54 135 Harrison et al.，2018
    膜翅目Hymenoptera 茧蜂科Braconidae 中红侧沟茧蜂Microplitis mediator 触角转录组Antennal transcriptome 169 17 Wang et al.，2017b；Wang et al.，2016c
    膜翅目Hymenoptera 茧蜂科Braconidae 菜蛾盘绒茧蜂Cotesia vestalis 触角转录组Antennal transcriptome 25 3 Liu et al.，2020b
    膜翅目Hymenoptera 茧蜂科Braconidae 腰带长体茧蜂Macrocentrus cingulum 触角转录组Antennal transcriptome 79 13 Ahmed et al.，2016
    膜翅目Hymenoptera 茧蜂科Braconidae 斑痣悬茧蜂Meteorus pulchricornis 触角转录组Antennal transcriptome 99 19 Sheng et al.，2017
    膜翅目Hymenoptera 茧蜂科Braconidae 阿维蚜茧蜂Aphidius ervi 基因组Genome 228 38 Dennis et al.，2020
    膜翅目Hymenoptera 茧蜂科Braconidae 烟蚜茧蜂Aphidius gifuensis 基因组Genome；转录组Transcriptome 80 25 Fan et al.，2018；Li et al.，2021a
    膜翅目Hymenoptera 茧蜂科Braconidae 豆柄瘤蚜茧蜂Lysiphlebus fabarum 基因组Genome 156 37 Dennis et al.，2020
    膜翅目Hymenoptera 赤眼蜂科Trichogrammatidae 螟黄赤眼蜂Trichogramma chilonis 转录组Transcriptome 45 14 Liu et al.，2018c
    膜翅目Hymenoptera 赤眼蜂科Trichogrammatidae 稻螟赤眼蜂Trichogramma japonicum 转录组Transcriptome 51 7 Li et al.，2021b
    膜翅目Hymenoptera 赤眼蜂科Trichogrammatidae 松毛虫赤眼蜂Trichogramma dendrolimi 基因组Genome 100 27 Zhang et al.，2023b
    膜翅目Hymenoptera 姬小蜂科Eulophidae 白蛾周氏啮小蜂Chouioia cunea 触角转录组Antennal transcriptome 80 10 Zhao et al.，2016
    膜翅目Hymenoptera 蜜蜂科Apidae 中华蜜蜂Apis cerana 基因组Genome 119 10 Park et al.，2015
    膜翅目Hymenoptera 蜜蜂科Apidae 意大利蜜蜂Apis mellifera 基因组Genome 163 21 Robertson and Wanner, 2006；Paoli and Galizia, 2021
    膜翅目Hymenoptera 金小蜂科Pteromalidae 丽蝇蛹集金小蜂Nasonia vitripennis 基因组Genome 301 111 Robertson et al.，2010；Harrison et al.，2018
    膜翅目Hymenoptera 茎蜂科Cephidae 麦茎蜂Cephus cinctus 基因组Genome 72 49 Robertson et al.，2018b
    膜翅目Hymenoptera 蚁科Formicidae 巨首芭切叶蚁Atta cephalotes 基因组Genome 376 18 Kock et al.，2013；Engsontia et al.，2015
    膜翅目Hymenoptera 蚁科Formicidae 佛罗里达弓背蚁Camponotus floridanus 转录组Transcriptome 407 31 Zhou et al.，2012
    膜翅目Hymenoptera 蚁科Formicidae 跳镰猛蚁Harpegnathos saltator 转录组Transcriptome 377 23 Zhou et al.，2012
    膜翅目Hymenoptera 蚁科Formicidae 阿根廷蚁Linepithema humile 基因组Genome 367 32 Smith et al.，2011
    膜翅目Hymenoptera 蚁科Formicidae 红胡须蚁Pogonomyrmex barbatus 基因组Genome 344 24 Smith et al.，2011
    膜翅目Hymenoptera 旋小蜂科Eupelmidae 荔蝽卵平腹小蜂Anastatus japonicus 触角转录组Antennal transcriptome 184 17 Zhan et al.，2023
    蜻蜓目Odonata 色蟌科Calopterygidae 华丽色蟌Calopteryx splendens 基因组Genome 5 21 Ioannidis et al.，2017
    直翅目Orthoptera 蝗科Acrididae 东亚飞蝗Locusta migratoria 基因组Genome；触角转录组Antennal
    transcriptome    		 142 32 Wang et al.，2015
    直翅目Orthoptera 蝗科Acrididae 青脊竹蝗Ceracris nigricornis 触角转录组Antennal transcriptome 71 8 Yuan et al.，2019
    直翅目Orthoptera 蝗科Acrididae 黄脊竹蝗Ceracris kiangsu 触角转录组Antennal transcriptome 91 13 Li et al.，2020a
    直翅目Orthoptera 蝗科Acrididae 亚洲小车蝗Oedaleus asiaticus 触角转录组Antennal transcriptome 60 6 Zhou et al.，2019b
    直翅目Orthoptera 蝗科Acrididae 中华稻蝗Oxya chinensis 触角转录组Antennal transcriptome 94 12 Cui et al.，2019

    附表  2   昆虫嗅觉受体功能研究进展

    Appendix Table  2   Progress in the functional deorphanization of insect olfactory receptors

    目Order 科Family 物种Species 受体(配体) Receptor (Ligand) 参考文献References
    双翅目Diptera 果蝇科Drosophilidae 黑腹果蝇Drosophila melanogaster OR67 (cis-vacceny acetate) Kurtovic et al.，2007
    双翅目Diptera 果蝇科Drosophilidae 黑腹果蝇Drosophila melanogaster IR75a+IR8a (acetic acid, propionic acid) Abuin et al.，2011
    双翅目Diptera 果蝇科Drosophilidae 黑腹果蝇Drosophila melanogaster IR84a+IR8a (phenylacetaldehyde) Abuin et al.，2011
    双翅目Diptera 果蝇科Drosophilidae 黑腹果蝇Drosophila melanogaster IR31a+IR8a (2-oxopentanoic acid) Silbering et al.，2011
    双翅目Diptera 果蝇科Drosophilidae 黑腹果蝇Drosophila melanogaster IR64a+IR8a (acetic acid) Ai et al.，2013
    双翅目Diptera 果蝇科Drosophilidae 黑腹果蝇Drosophila melanogaster IR92a (ammonia, amines) Min et al.，2013
    双翅目Diptera 果蝇科Drosophilidae 黑腹果蝇Drosophila melanogaster IR41a+IR76b (spermine, putrescine, 1,4-diamionbutane) Huassain et al.，2016
    双翅目Diptera 果蝇科Drosophilidae 黑腹果蝇Drosophila melanogaster IR25a+IR76b (acids) Chen and Amrein, 2017
    双翅目Diptera 果蝇科Drosophilidae 黑腹果蝇Drosophila melanogaster IR75b+IR75c+IR8a (butyric acid, propionic acid) Prieto-Godino et al.，2017
    双翅目Diptera 果蝇科Drosophilidae 大灰优食蚜蝇Eupeodes corollae OR25 (eugenol, p-cresol, methyl eugenol) Li et al.，2020b
    双翅目Diptera 果蝇科Drosophilidae 大灰优食蚜蝇Eupeodes corollae OR3 (E-β-farnesene) Wang et al.，2022b
    双翅目Diptera 蚊科Culicidae 冈比亚按蚊Anopheles gambiae IR41a,IR41c+IR76b+IR25a (amines) Pitts et al.，2017
    鳞翅目Lepidoptera 夜蛾科Noctuidae 棉铃虫Helicoverpa armigera OR6 (Z9-16:OH)；OR14b (Z9-14:Ald) Jiang et al.，2014
    鳞翅目Lepidoptera 夜蛾科Noctuidae 棉铃虫Helicoverpa armigera OR13 (Z11-16:Ald)；OR6 (Z9-16:Ald, Z9-14:Ald)；OR16 (Z11-16:OH) Liu et al.，2013a
    鳞翅目Lepidoptera 夜蛾科Noctuidae 棉铃虫Helicoverpa armigera OR42 (phenylacetaldehyde) Guo et al., 2021
    鳞翅目Lepidoptera 夜蛾科Noctuidae 棉铃虫Helicoverpa armigera OR13 (Z7-12:Ac) Sun et al., 2024
    鳞翅目Lepidoptera 夜蛾科Noctuidae 棉铃虫Helicoverpa armigera OR11 (3Z,6Z,9Z-21:H) Wang et al., 2024b
    鳞翅目Lepidoptera 夜蛾科Noctuidae 棉铃虫Helicoverpa armigera IR8a (acetic acid) Zhang et al., 2022a
    鳞翅目Lepidoptera 夜蛾科Noctuidae 烟青虫Helicoverpa assulta OR6 (Z9-16:OH); OR16 (Z9-14:Ald); Jiang et al., 2014
    鳞翅目Lepidoptera 夜蛾科Noctuidae 烟青虫Helicoverpa assulta OR23 (E-β-farnesene) Wu et al., 2019b
    鳞翅目Lepidoptera 夜蛾科Noctuidae 烟青虫Helicoverpa assulta OR14b (Z9-16:Ald) Yang et al., 2017
    鳞翅目Lepidoptera 夜蛾科Noctuidae 烟青虫Helicoverpa assulta OR13 (Z7-12:Ac) Sun et al., 2024
    鳞翅目Lepidoptera 夜蛾科Noctuidae 烟芽夜蛾Heliothis virescens OR13 (Z11-16:Ald) Grosse-Wilde et al., 2007
    鳞翅目Lepidoptera 夜蛾科Noctuidae 烟芽夜蛾Heliothis virescens OR6 (Z9-14:Ald); OR16 (Z11-16:OH) Wang et al., 2011
    鳞翅目Lepidoptera 夜蛾科Noctuidae 烟芽夜蛾Heliothis virescens OR13 (Z7-12:Ac) Sun et al., 2024
    鳞翅目Lepidoptera 夜蛾科Noctuidae 东方粘虫Mythimma separata OR3 (Z11-16:Ald); OR2 (Z9-14:Ald) Jiang et al., 2017
    鳞翅目Lepidoptera 夜蛾科Noctuidae 东方粘虫Mythimma separata IR8a (acetic acid) Tang et al., 2020
    鳞翅目Lepidoptera 夜蛾科Noctuidae 斜纹夜蛾Spodoptera litura OR12 (cis-3-hexenyl acetate); OR19 (4-ethylacetophenone) Zhang et al., 2013a
    鳞翅目Lepidoptera 夜蛾科Noctuidae 斜纹夜蛾Spodoptera litura OR51 (vanillin) Wei et al., 2023
    鳞翅目Lepidoptera 夜蛾科Noctuidae 斜纹夜蛾Spodoptera litura OR6 (Z9,E12-14:OAc); OR13 (Z9,E12-14:OAc, Z9-14:OAc); OR16 (Z9-14:OH) Zhang et al., 2015b
    鳞翅目Lepidoptera 夜蛾科Noctuidae 甜菜夜蛾Spodoptera exigua OR13 (Z9,E12-14:OAc, Z9-14:OAc); OR16 (Z9-14:OH) Liu et al., 2013b
    鳞翅目Lepidoptera 夜蛾科Noctuidae 甜菜夜蛾Spodoptera exigua OR3 (E-β-farnesene) Liu et al., 2014
    鳞翅目Lepidoptera 夜蛾科Noctuidae 美洲棉铃虫Helicoverpa zea OR14b (Z9-16:Ald, Z9-14:Ald); OR16 (Z11-16:OH) Cao et al., 2023
    鳞翅目Lepidoptera 夜蛾科Noctuidae 美洲棉铃虫Helicoverpa zea OR13 (Z11-16:Ald); OR6 (Z9-16:OH, Z9-14:Ald, Z9-16:Ald); Cao et al., 2023
    鳞翅目Lepidoptera 夜蛾科Noctuidae 阿根廷棉铃虫Helicoverpa gelotopoeon OR13 (Z11-16:Ald); OR14b (Z9-16:Ald, Z9-14:Ald); OR16 (Z9-14:Ald) Cao et al., 2023
    鳞翅目Lepidoptera 夜蛾科Noctuidae 阿根廷棉铃虫Helicoverpa gelotopoeon OR6 (Z9-16:OH, Z9-14:Ald, Z9-16:Ald); Cao et al., 2023
    鳞翅目Lepidoptera 夜蛾科Noctuidae 草地贪夜蛾Spodoptera frugiperda OR13 (Z9-14:Ac, Z9,E12-14:Ac); OR56 (Z7-12:Ac); OR62 (Z7-12:Ac) Guo et al., 2022a
    鳞翅目Lepidoptera 夜蛾科Noctuidae 草地贪夜蛾Spodoptera frugiperda OR6 (Z9,E12-14:Ac); OR16 (Z9-14:OH, Z9-14:Ald) Guo et al., 2022a
    鳞翅目Lepidoptera 夜蛾科Noctuidae 草地贪夜蛾Spodoptera frugiperda OR13 (Z7-12:Ac) Sun et al., 2024
    鳞翅目Lepidoptera 夜蛾科Noctuidae 草地贪夜蛾Spodoptera frugiperda IR75q.2 (nonanoic acid) Guo et al.，2023
    鳞翅目Lepidoptera 夜蛾科Noctuidae 海灰翅夜蛾Spodoptera littoralis OR6 (Z9,E12-14:OAc)； De Fouchier et al.，2017
    鳞翅目Lepidoptera 夜蛾科Noctuidae 黄地老虎Agrotis segetum IR75p.1 (hexanoic acid)；IR75q.1 (octanoic acid) Hou et al.，2022
    鳞翅目Lepidoptera 菜蛾科Plutellidae 小菜蛾Plutella xylostella OR6 (heptanal) Liu et al.，2024
    鳞翅目Lepidoptera 菜蛾科Plutellidae 小菜蛾Plutella xylostella OR11 (benzyl alcohol, salicylaldehyde, phenylacetaldehyde) Liu et al.，2022
    鳞翅目Lepidoptera 菜蛾科Plutellidae 小菜蛾Plutella xylostella OR41 (Z9-14:Ac, Z9-14:OH, Z9-14:Ald) Liu et al.，2018d
    鳞翅目Lepidoptera 蚕蛾科Bombycidae 家蚕Bombyx mori OR1 (bombykol) Sakurai et al.，2015
    鳞翅目Lepidoptera 蚕蛾科Bombycidae 家蚕Bombyx mori OR56 (cis-jasmone) Tanaka et al.，2009
    鳞翅目Lepidoptera 蚕蛾科Bombycidae 家蚕Bombyx mori OR19 (linalool) Anderson et al.，2009
    鳞翅目Lepidoptera 天蛾科Sphingidae 烟草天蛾Manduca sexta OR4 (Sex pheromone)；OR5 (linalool) Große-Wilde et al.，2010
    鳞翅目Lepidoptera 天蛾科Sphingidae 烟草天蛾Manduca sexta OR35 (α-copaene) Zhang et al.，2022b
    鳞翅目Lepidoptera 天蛾科Sphingidae 烟草天蛾Manduca sexta IR8a (hexanoic acid, 3-methylpentanoic) Zhang et al.，2019b
    鞘翅目Coleoptera 金龟科Scarabaeidae 暗黑鳃金龟Holotrichia parallela OR14 (L-isoleucine methyl) Wang et al.，2024c
    半翅目Hemiptera 盲蝽科Miridae 苜蓿盲蝽Adelphocoris lineolatus OR59 (menthyl salicylate) Xiao et al.，2020
    半翅目Hemiptera 蚜科Aphididae 绿盲蝽Apolygus lucorum OR47 (linalool) Zhang et al.，2022c
    半翅目Hemiptera 蚜科Aphididae 豌豆蚜Acyrthosiphon pisum OR23 (Green leaf volatiles) Huang et al.，2022
    半翅目Hemiptera 蚜科Aphididae 豌豆蚜Acyrthosiphon pisum OR5 (E-β-farnesene) Zhang et al.，2017b
    半翅目Hemiptera 叶蝉科Cicadellidae 茶小绿叶蝉Empoasca onukii IR25 (1-phenylethanol) Zhang et al.，2023c
    蜚蠊目Blattodea 蜚蠊科Blattidae 美洲大蠊Periplaneta americana OR53 (periplanone-A)；OR100 (periplanone-B) Li et al.，2024
    膜翅目Hymenoptera 姬蜂科Ichneumonidae 棉铃虫齿唇姬蜂Campoletis chlorideae OR18 (14:Ald)；OR47 (2-heptadecanone) Guo et al.，2022b
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  • 收稿日期:  2024-06-18
  • 修回日期:  2024-10-19
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