昆虫外周嗅觉系统研究进展
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摘要
嗅觉作为昆虫感知外界环境的主要手段之一,在觅食、防御、交配、繁殖、信息交流及栖息地选择等方面起着十分重要的作用。研究环境中或来自生物体的气味分子作用于昆虫感觉器官引起昆虫产生行为反应的机制,有助于人们了解昆虫的寄主识别、种间互作和种内交流等行为及制定害虫治理和益虫保护等策略。本文对昆虫嗅觉系统中气味分子的传导过程,及外周嗅觉系统相关蛋白的功能及研究现状进行综述。气味分子通过嗅觉感受器进入外周嗅觉系统,进而转化为电信号进入中枢神经系统进行加工处理。气味结合蛋白与气味分子进行结合,并将其运输到气味受体;化学感受蛋白也能够与气味分子结合,但其在昆虫体内具有更为广泛的功能;气味受体具有识别气味结合蛋白或化学感受蛋白传递而来的气味分子的功能,并将这种化学信号转化为电信号;离子受体在外周嗅觉系统具有和气味受体相似的作用,但其气味反应谱更窄;感觉神经元膜蛋白在气味分子的传递过程中可能起到的重要作用;气味降解酶也存在多种降解气味分子的假说。通过对昆虫外周嗅觉系统的研究进展进行梳理,为今后研究昆虫外周嗅觉系统提供参考。
As one of the most important methods to apperceive the environments,olfaction plays a crucial role in foraging,defense,mating,reproduction,information communication,and habitat selection of insects.To investigate the mechanism of how odorants from the environments or organisms act on sense organ of insects and elicit behavior response will help us understand the behavior of insects,such as host recognition,interspecific interaction and intraspecific communication,and develop strategies for pest management and beneficial insect protection.This paper reviews the advance in the research on the odorant transduction of insect peripheral olfac?tory system,and the function of related proteins.Through the olfactory receptor,odorants get into the peripheral olfactory system,and then convert into electrical signals for processing in the central nervous system.The odor?ant?binding proteins bind with the odorants and transport them to odorant receptors.The chemosensory proteins can also bind with the odorants,but the chemosensory proteins have a broader range of functions in insects.The odorant receptors can recognize the odorants transferred from odorant?binding proteins or chemosensory pro?teins,and convert the chemical signals into electrical signals.The ionotropic receptors have similar effects on odorant receptors in insect peripheral olfactory system,but the ionotropic receptors have a much narrower re?sponse spectrum of odorants.The sensory neuron membrane proteins may play an important role in the transmis?sion of odorants.And the odorant?degrading enzymes also have multiple hypotheses for degrading odorants.Combing the progress in the research on insect peripheral olfactory system would provide some references for further study of insect peripheral olfactory system.
As one of the most important methods to apperceive the environments,olfaction plays a crucial role in foraging,defense,mating,reproduction,information communication,and habitat selection of insects.To investigate the mechanism of how odorants from the environments or organisms act on sense organ of insects and elicit behavior response will help us understand the behavior of insects,such as host recognition,interspecific interaction and intraspecific communication,and develop strategies for pest management and beneficial insect protection.This paper reviews the advance in the research on the odorant transduction of insect peripheral olfac?tory system,and the function of related proteins.Through the olfactory receptor,odorants get into the peripheral olfactory system,and then convert into electrical signals for processing in the central nervous system.The odor?ant?binding proteins bind with the odorants and transport them to odorant receptors.The chemosensory proteins can also bind with the odorants,but the chemosensory proteins have a broader range of functions in insects.The odorant receptors can recognize the odorants transferred from odorant?binding proteins or chemosensory pro?teins,and convert the chemical signals into electrical signals.The ionotropic receptors have similar effects on odorant receptors in insect peripheral olfactory system,but the ionotropic receptors have a much narrower re?sponse spectrum of odorants.The sensory neuron membrane proteins may play an important role in the transmis?sion of odorants.And the odorant?degrading enzymes also have multiple hypotheses for degrading odorants.Combing the progress in the research on insect peripheral olfactory system would provide some references for further study of insect peripheral olfactory system.
引文
[1]Zwiebel L J,Takken W.Olfactory regulation of mosquito-host interaction[J].Insect Biochemistry and molecular Biology,2004,34(7):645-652.
[2]Brito N F,Moreira M F,Melo A C.A look inside odorant-binding proteins in insect chemoreception[J].Journal of InsectPhysiolog,2016,95:51-65.
[3]Leal W S.Odorant reception in insects:roles of receptors binding proteins,and degrading enzymes[J].Annual Review ofEntomology,2013,58(1):373-391.
[4]Schneider D.Insect antennae[J].Annual Review of Entomology,1964,9(1):103-122.
[5]Sato K,Touhara K. Insect olfactory:receptors,signal transduction,and behavior[J]. Results and Problems in CellDifferentiation,2008,47:121-138.
[6]Manoharan M,Chong M N F,Vaitinadapoule A,et al.Comparative genomics of odorant b inding proteins in anophelesgambiae,Aedes aegypti,and Culex quinquefasciatus[J].Genome Biology and Evolution,2013,5(1):163-180.
[7]Rützler M,Zwiebel L J. Molecular biology of insect olfaction:recent progress and conceptual models[J]. Journal ofComparative Physiology A,2005,191(9):777-790.
[8]Sandler B H,Nikonova L,Leal W S,et al.Sexual attraction in the silkworm moth:structure of the pheromone-binding-protein-bombykol complex[J].Cell Chemical Biology,2000,7(2):143-151.
[9]那杰,白旭,郭瑞,等.昆虫气味认知的嗅觉神经结构及分子机制[J].现代生物医学进展,2015,15(4):749-755.Na J,Bai X,Guo R,et al.Olfactory neuralstructure and molecular mechanism of ordor perception in insect[J].Progress inModern Biomedicine,2015,15(4):749-755.
[10]Peng G,Leal W S.Identification and cloning of a pheromone-binding protein from the Oriental beetle,Exomala orientalis[J].Journal of Chemical Ecology,2001,27(11):2183-2192.
[11]Zhou J J,He X L,Pickett J A,et al.Identification of odorant-binding proteins of the yellow fever mosquito Aedes aegypti:genome annotation and comparative analyses[J].Insect Molecular Biology,2008,17(2):147-163.
[12]Steinbrecht R A.Odorant-binding proteins:expression and function[J].Annals of the New York Academy of Sciences,1998,855(1):323-332.
[13]Vogt R G,Riddiford L M.Pheromone binding and inactivation by moth antennae[J].Nature,1981,293(5828):161-163.
[14]Xu Y L,He P,Zhang L,et al.Large-scale identification of odorant-binding proteins and chemosensory proteins fromexpressed sequence tags in insects[J].BMC Genomics,2009,10(1):632.
[15]Li Z Q,He P,Zhang Y N,et al.Molecular and functional characterization of three odorant-b inding protein from Periplanetaamericana[J].PLoS One,2017,12(1):e0170072.
[16]Yuko I,Jun I,Leal W S.Fatty acid solubilizer from the oral disk of the blowfly[J].PLoS One,2013,8(1):e51779.
[17]Ribeiro J M,Genta F A,Sorgine M H,et al.An insight into the transcriptome of the digestive tract of the bloodsucking bug,Rhodnius prolixus[J].PLoS Neglected Tropical Diseases,2014,8(1):2594.
[18]Sun Y L,Huang L Q,Pelosi P,et al.Expression in antennae and reproductive organs suggests a dual role of an odorant-binding protein in two sibling Helicoverpa Species[J].PLoS One,2012,7(1):e30040.
[19]张瑜,张胜男,张媛媛,等.昆虫嗅觉机制的研究进展[J].福建农业学报,2016,31(5):538-544.Zhang Y,Zhang S N,Zhang Y Y,et al.Advances in insect olfactory mechanism[J].Fujian Journal of Agricultural Sciences,2016,31(5):538-544.
[20]Li H L,Ni C X,Zhang L Y,et al.Chemosensory proteins of the eastern honeybee,Apis cerana:identification,tissuedistribution and olfactory related functional characterization[J]. Comparative Biochemistry and Physiology Part B:Biochemistry and Molecular Biology,2016,194-195(4):11-19.
[21]Xue W,Fan J,Zhang Y,et al.Identification and expression analysis of candidate odorantbinding protein and chemosensoryprotein genes by antennal transcriptome of Sitobion avenae[J].PLoS One,2016,11(8):e0161839.
[22]Yuan X,Jiang Y D,Wang G Y,et al. Odorant-binding proteins and chemosensory proteins from an invasive pestLissorhoptrus oryzophilus(Coleoptera:Curculionidae)[J].Environmental Entomology,2016,45(5):1276-1286.
[23]Yao C A,Ignell R,Carlson J R.Chemosensory coding by neurons in the coeloconic sensilla of the Drosophila antenna[J].The Journal of Neuroscience,2005,25(37):8359-8367.
[24]Mckenzie S K,Oxley P R,Kronauer D J.Comparative genomics and transcriptomics in ants provide new insights into theevolution and function of odorant binding and chemosensory proteins[J].BMC Genomics,2014,15(1):718.
[25]Forêt S,Wanner K W,Maleszka R. Chemosensory proteins in the honey bee:insights from the annotated genome,comparative analyses and expressional profiling[J].Insect Biochemistry and Molecular Biology,2007,37(1):19-28.
[26]Vieira F G,Rozas J. Comparative genomics of the odorant-binding and chemosensory protein gene families across theArthropoda:originandevolutionaryhistoryofthechemosensorysystem[J].GenomeBiologyandEvolution,2011,3(1):476-490.
[27]梅婷,何正波,王小婷,等.中华按蚊化学感受蛋白(CSP)家族基因的全基因组鉴定和特征分析[J].昆虫学报,2017,60(2):136-147.Mei T,He Z B,Wang X T,et al.Genome-wide identification and characterization of genes of the chemosensory protein(CSP)family in Anopheles sinensis(Diptera:Culicidae)[J].Acta Entomologica Sinica,2017,60(2):136-147.
[28]Kamikouchi A,Morioka M,Kubo T.Identification of honeybee antennal proteins/genes expressed in a sex-and/or casteselective manner[J].Zoological Science,2004,21(1):53-62.
[29]Steller M M,Kambhampati S,Caragea D.Comparative analysis of expressed sequence tags from three castes and two lifestages of the termite Reticulitermes flavipes[J].BMC Genomics,2010,11(1):463.
[30]Guo W,Wang X,Ma Z,et al.CSP and takeout genes modulate the switch between attraction and repulsion during behavioralphase change in the migratory locust[J].PLOS Genetics,2011,7(2):e1001291.
[31]Ozaki M,Wada-Katsumata A,Fujikawa K,et al.Ant nestmate and nonnestmate discrimination by a chemosensory sensillum[J].Science,2005,309(5732):311-314.
[32]Dani F R,Michelucci E,Francese S,et al.Odorant-binding proteins and chemosensory proteins in pheromone detection andrelease in the silkmoth Bombyx mori[J].Chemical Senses,2011,36(4):335-344.
[33]邓培渊,乔惠丽,李丹丹,等.家蚕化学感受蛋白BmCSP4表达谱及结合特性分析[J].昆虫学报,2011,53(8):869-876.Deng P Y,Qiao H L,Li D D,et al.Expression pattern and binding specificity of chemosensory protein BmCSP4 in the silk?worm,Bombyx mori[J].Acta Entomologica Sinica,2011,53(8):869-876.
[34]李庆荣,肖阳,邢东旭,等.化学感受蛋白家族基因在家蚕5龄幼虫表皮中的表达分析[J].蚕业科学,2017,43(1):39-44.Li Q R,Xiao Y,Xin D X,et al.Expression analysis of chemosensory protein family genes in epidermis of the 5th instar silk?worm larvae[J].Science of Sericulture,2017,43(1):39-44.
[35]Guo Z,Cheng Z Y,Huang F,et al.Microarray analysis of global gene regulation in the Cry1Ab-resistant and Cry1Ab-susceptible strains of Diatraea saccharalis[J].Pest Management Science,2012,68(5):718-730.
[36]Liu Y L,Guo H,Huang L Q,et al.Unique function of a chemosensory protein in the proboscis of two Helicoverpa species[J].Journal of Experimental Biology,2014,217(10):1821-1826.
[37]Leal W S,Chen A M,Ishida Y,et al.Kinetics and molecular properties of pheromone binding and release[J].PNAS,2005,102(15):5386-5391.
[38]Clyne P J,Warr C G,Freeman M R,et al.A novel family of divergent seven-transmembrane proteins:candidate odorantreceptors in Drosophila[J].Neuron,1999,22(2):327-328.
[39]Larsson M C,Domingos A I,Jones W D,et al.Or83b encodes a broadly expressed odorant receptor essential for Drosophilaolfaction[J].Neuron,2004,43(5):703-714.
[40]Leal W S,Ishida Y,Pelletier J,et al.Olfactory proteins mediating chemical communication in the navel orangeworm moth,Amyelois transitella[J].PLoS One,2009,4(9):e7235.
[41]Vosshall L B,Hansson B S.A unified nomenclature system for the insect olfactory coreceptor[J].Chemical Senses,2011,36(6):497-498.
[42]Nakagawa T,Pellegrino M,Sato K,et al.Amino acid residues contributing to function of the heteromeric insect olfactoryreceptor complex[J].PLoS One,2012,7(3):e32372.
[43]Hildebrand J G.Analysis of chemical signals by nervous systems[J].PNAS,1995,92(1):67-74.
[44]Neuhaus E M,Gisselmann G,Zhang W,et al.Odorant receptor heterodimerization in the olfactory system of Drosophilamelanogaster[J].Nature Neuroscience,2005,8(1):15-17.
[45]Benton R,Sachse S,Michnick S W,et al.Atypical membrane topology and heteromeric function of Drosophila odorantreceptors in vivo[J].PLoS Biology,2006,4(2):240-257.
[46]代玉华,赵忠懿,程鹏,等.蚊虫嗅觉受体的研究进展[J].寄生虫与医学昆虫学报,2014,21(1):59-64.Dai Y H,Zhao Z Y,Cheng P,et al.Advances in research of odorant receptor proteins of mosquitoes[J].Acta Parasitologicaet Medica Entomologica Sinica,2014,21(1):59-64.
[47]Dobritsa A A,Naters W,Warr C G,et al.Integrating the molecular and cellular basis of odor coding in the Drosophilaantenna[J].Neuron,2003,37(5):827-841.
[48]Sakurai T,Nakagawa T,Mitsuno H,et al.Identification and functional characterization of a sex pheromone receptor in thesilkmoth Bombyx mori[J].PNAS,2004,101(47):16653-16658.
[49]Nakagawa T,Sakurai T,Nishioka T,et al.Insect sex-pheromone signals mediated by specific combinations of olfactoryreceptors[J].Science,2005,307(5715):1638-1642.
[50]Liu Y,Liu C,Lin K,et al.unctional specificity of sex pheromone receptors in the cotton bollworm Helicoverpa armigera[J].PLoS One,2013,8(4):e62094.
[51]Wanner K W,Nichols A S,Allen J E,et al.Sex pheromone receptor specificity in the European corn borer moth,Ostrinianubilalis[J].PLoS One,2010,5(1):e8685.
[52]Miura N,Nakagawa T,Touhara K,et al.Broadly and narrowly tuned odorant receptors are involved in female sex pheromonereception in Ostrinia moths[J].Insect Biochemistry and Molecular Biology,2010,40(1):64-73.
[53]Chang H,Liu Y,Yang T,et al.Pheromone binding proteins enhance the sensitivity of olfactory receptors to sex pheromonesin Chilo suppressalis[J].Scientific Reports,2015,5(1):13093.
[54]Benton R,Vannice K S,Gomez-Diaz C,et al.Variant ionotropic glutamate receptors as chemosensory receptors in Drosophila[J].Cell,2009,136(1):149-162.
[55]Montell C.A taste of the Drosophila gustatory receptors[J].Current Opinion in Neurobiology,2009,19(4):345-353.
[56]Koh T W,He Z,Gorur-Shandilya S,et al.The Drosophila IR20a clade of ionotropic receptors are candidate taste andpheromone receptors[J].Neuron,2014,83(4):850-865.
[57]Chen C,Buhl E,Xu M,et al.Drosophila Ionotropic Receptor 25a mediates circadian clock resetting by temperature[J].Nature,2015,527(7579):516-520.
[58]Croset V,Rytz R,Cummins S F,et al.Ancient protostome origin of chemosensory ionotropic glutamate receptors and theevolution of insect taste and olfaction[J].PLoS Genetics,2010,6(8):e1001064.
[59]Olivier V,Monsempes C,Fran?ois M C,et al.Candidate chemosensory ionotropic receptors in a Lepidoptera[J].InsectMolecular Biology,2011,20(2):189-199.
[60]Bengtsson J M,Trona F,MontagnéN,et al.Putative chemosensory receptors of the codling moth,Cydia pomonella,identifiedby antennal transcriptome analysis[J].PLoS One,2012,7(2):e31620.
[61]Liu Y,Gu S,Zhang Y,et al.Candidate olfaction genes identified within the Helicoverpa armigera antennal transcriptome[J].PLoS One,2012,7(10):e48260.
[62]Getahun M N,Wicher D,Hansson B S,et al.Temporal response dynamics of Drosophila olfactory sensory neurons dependson receptor type and response polarity[J].Frontiers in Cellular Neuroscience,2012,6(47):54-65.
[63]孙畅仪.小菜蛾气味受体和离子型受体的克隆与表达[D].哈尔滨:东北林业大学,2014.Sun C Y.Gene cloning and expression analysis of odorant receptors and ionotropic receptors in the Diamondback Moth(Plutella xylostella)[D].Harbin:Northeast Forestry University,2014.
[64]Su C Y,Menuz K,Carlson J R.Olfactory perception:receptors,cells,and circuits[J].Cell,2009,139(1):45-59.
[65]Rasmussen J T,Berglund L,Rasmussen M S,et al.Assignment of disulfide bridges in bovine CD36[J].European Journal ofBiochemistry,1998,257(2):488-494.
[66]Rogers M E,Sun M,Lerner M R,et al.Snmp-1,a novel membrane protein of olfactory neurons of the silk moth Antheraeapolyphemus with homology to the CD36 family of membrane proteins[J].Journal of Biological Chemistry,1997,272(23):14792-14799.
[67]Liu S,Zhang Y R,Zhou W W,et al.Identification and characterization of two sensory neuron membrane proteins fromCnaphalocrocis medinalis(Lepidoptera:Pyralidae)[J].Archives of Insect Biochemistry and Physiology,2013,82(1):29-42.
[68]Forstner M,Gohl T,Gondesen I,et al.Differential expression of SNMP-1 and SNMP-2 proteins in pheromone-sensitive hairsof moths[J].Chemical Senses,2008,33(3):291-299.
[69]罗梅浩,蔡永萍,赵艳艳,等.烟夜蛾感觉神经元膜蛋白基因的克隆、序列分析与组织特异性表达[J].中国烟草学报,2012,18(5):91-95.Luo M H,Cai Y P,Zhao Y Y,et al.Cloning,sequence analysis and tissue-specific expression of sensory neuron membraneprotein gene from Helicoverpa assulta(Guenée)[J].Acta Tabacaria Sinica,2012,18(5):91-95.
[70]Gu S H,Yang R N,Guo M B,et al.Molecular identification and differential expression of sensory neuron membrane proteinsin the antennae of the black cutworm moth Agrotis ipsilon[J].Journal of Insect Physiology,2013,59(4):430-443.
[71]Rogers M E,Krieger J,Vogt R G.Antennal SNMPs(sensory neuron membrane proteins)of lepidoptera define a uniquefamily of invertebrate CD36-like proteins[J].Journal of Neurobiology,2001,49(1):47-61.
[72]Vogt R G,Miller N E,Litvack R,et al.The insect SNMP gene family[J].Insect Biochemistry and Molecular Biology,2009,39(7):448-456.
[73]Kaissling K E.Olfactory perireceptor and receptor events in moths:a kinetic model revised[J].Journal of ComparativePhysiology A,2009,195(10):895-922.
[74]Merlin C,Fran?ois M C,Bozzolan F,et al.A new aldehyde oxidase selectively expressed in chemosensory organs of insects[J].Biochemical and Biophysical Research Communications,2005,332(1):4-10.
[75]Durand N,Carot-Sans G,Bozzolan F,et al.Degradation of pheromone and plant volatile components by a same odorant-degrading enzyme in the cotton leafworm,Spodoptera littoralis[J].PLoS One,2011,6(12):e29147.
[76]Ishida Y,Leaal W S.Chiral discrimination of the Japanese beetle sex pheromone and a behavioral antagonist by a pheromone-degrading enzyme[J].PNAS,2008,105(26):9076-9080.
[2]Brito N F,Moreira M F,Melo A C.A look inside odorant-binding proteins in insect chemoreception[J].Journal of InsectPhysiolog,2016,95:51-65.
[3]Leal W S.Odorant reception in insects:roles of receptors binding proteins,and degrading enzymes[J].Annual Review ofEntomology,2013,58(1):373-391.
[4]Schneider D.Insect antennae[J].Annual Review of Entomology,1964,9(1):103-122.
[5]Sato K,Touhara K. Insect olfactory:receptors,signal transduction,and behavior[J]. Results and Problems in CellDifferentiation,2008,47:121-138.
[6]Manoharan M,Chong M N F,Vaitinadapoule A,et al.Comparative genomics of odorant b inding proteins in anophelesgambiae,Aedes aegypti,and Culex quinquefasciatus[J].Genome Biology and Evolution,2013,5(1):163-180.
[7]Rützler M,Zwiebel L J. Molecular biology of insect olfaction:recent progress and conceptual models[J]. Journal ofComparative Physiology A,2005,191(9):777-790.
[8]Sandler B H,Nikonova L,Leal W S,et al.Sexual attraction in the silkworm moth:structure of the pheromone-binding-protein-bombykol complex[J].Cell Chemical Biology,2000,7(2):143-151.
[9]那杰,白旭,郭瑞,等.昆虫气味认知的嗅觉神经结构及分子机制[J].现代生物医学进展,2015,15(4):749-755.Na J,Bai X,Guo R,et al.Olfactory neuralstructure and molecular mechanism of ordor perception in insect[J].Progress inModern Biomedicine,2015,15(4):749-755.
[10]Peng G,Leal W S.Identification and cloning of a pheromone-binding protein from the Oriental beetle,Exomala orientalis[J].Journal of Chemical Ecology,2001,27(11):2183-2192.
[11]Zhou J J,He X L,Pickett J A,et al.Identification of odorant-binding proteins of the yellow fever mosquito Aedes aegypti:genome annotation and comparative analyses[J].Insect Molecular Biology,2008,17(2):147-163.
[12]Steinbrecht R A.Odorant-binding proteins:expression and function[J].Annals of the New York Academy of Sciences,1998,855(1):323-332.
[13]Vogt R G,Riddiford L M.Pheromone binding and inactivation by moth antennae[J].Nature,1981,293(5828):161-163.
[14]Xu Y L,He P,Zhang L,et al.Large-scale identification of odorant-binding proteins and chemosensory proteins fromexpressed sequence tags in insects[J].BMC Genomics,2009,10(1):632.
[15]Li Z Q,He P,Zhang Y N,et al.Molecular and functional characterization of three odorant-b inding protein from Periplanetaamericana[J].PLoS One,2017,12(1):e0170072.
[16]Yuko I,Jun I,Leal W S.Fatty acid solubilizer from the oral disk of the blowfly[J].PLoS One,2013,8(1):e51779.
[17]Ribeiro J M,Genta F A,Sorgine M H,et al.An insight into the transcriptome of the digestive tract of the bloodsucking bug,Rhodnius prolixus[J].PLoS Neglected Tropical Diseases,2014,8(1):2594.
[18]Sun Y L,Huang L Q,Pelosi P,et al.Expression in antennae and reproductive organs suggests a dual role of an odorant-binding protein in two sibling Helicoverpa Species[J].PLoS One,2012,7(1):e30040.
[19]张瑜,张胜男,张媛媛,等.昆虫嗅觉机制的研究进展[J].福建农业学报,2016,31(5):538-544.Zhang Y,Zhang S N,Zhang Y Y,et al.Advances in insect olfactory mechanism[J].Fujian Journal of Agricultural Sciences,2016,31(5):538-544.
[20]Li H L,Ni C X,Zhang L Y,et al.Chemosensory proteins of the eastern honeybee,Apis cerana:identification,tissuedistribution and olfactory related functional characterization[J]. Comparative Biochemistry and Physiology Part B:Biochemistry and Molecular Biology,2016,194-195(4):11-19.
[21]Xue W,Fan J,Zhang Y,et al.Identification and expression analysis of candidate odorantbinding protein and chemosensoryprotein genes by antennal transcriptome of Sitobion avenae[J].PLoS One,2016,11(8):e0161839.
[22]Yuan X,Jiang Y D,Wang G Y,et al. Odorant-binding proteins and chemosensory proteins from an invasive pestLissorhoptrus oryzophilus(Coleoptera:Curculionidae)[J].Environmental Entomology,2016,45(5):1276-1286.
[23]Yao C A,Ignell R,Carlson J R.Chemosensory coding by neurons in the coeloconic sensilla of the Drosophila antenna[J].The Journal of Neuroscience,2005,25(37):8359-8367.
[24]Mckenzie S K,Oxley P R,Kronauer D J.Comparative genomics and transcriptomics in ants provide new insights into theevolution and function of odorant binding and chemosensory proteins[J].BMC Genomics,2014,15(1):718.
[25]Forêt S,Wanner K W,Maleszka R. Chemosensory proteins in the honey bee:insights from the annotated genome,comparative analyses and expressional profiling[J].Insect Biochemistry and Molecular Biology,2007,37(1):19-28.
[26]Vieira F G,Rozas J. Comparative genomics of the odorant-binding and chemosensory protein gene families across theArthropoda:originandevolutionaryhistoryofthechemosensorysystem[J].GenomeBiologyandEvolution,2011,3(1):476-490.
[27]梅婷,何正波,王小婷,等.中华按蚊化学感受蛋白(CSP)家族基因的全基因组鉴定和特征分析[J].昆虫学报,2017,60(2):136-147.Mei T,He Z B,Wang X T,et al.Genome-wide identification and characterization of genes of the chemosensory protein(CSP)family in Anopheles sinensis(Diptera:Culicidae)[J].Acta Entomologica Sinica,2017,60(2):136-147.
[28]Kamikouchi A,Morioka M,Kubo T.Identification of honeybee antennal proteins/genes expressed in a sex-and/or casteselective manner[J].Zoological Science,2004,21(1):53-62.
[29]Steller M M,Kambhampati S,Caragea D.Comparative analysis of expressed sequence tags from three castes and two lifestages of the termite Reticulitermes flavipes[J].BMC Genomics,2010,11(1):463.
[30]Guo W,Wang X,Ma Z,et al.CSP and takeout genes modulate the switch between attraction and repulsion during behavioralphase change in the migratory locust[J].PLOS Genetics,2011,7(2):e1001291.
[31]Ozaki M,Wada-Katsumata A,Fujikawa K,et al.Ant nestmate and nonnestmate discrimination by a chemosensory sensillum[J].Science,2005,309(5732):311-314.
[32]Dani F R,Michelucci E,Francese S,et al.Odorant-binding proteins and chemosensory proteins in pheromone detection andrelease in the silkmoth Bombyx mori[J].Chemical Senses,2011,36(4):335-344.
[33]邓培渊,乔惠丽,李丹丹,等.家蚕化学感受蛋白BmCSP4表达谱及结合特性分析[J].昆虫学报,2011,53(8):869-876.Deng P Y,Qiao H L,Li D D,et al.Expression pattern and binding specificity of chemosensory protein BmCSP4 in the silk?worm,Bombyx mori[J].Acta Entomologica Sinica,2011,53(8):869-876.
[34]李庆荣,肖阳,邢东旭,等.化学感受蛋白家族基因在家蚕5龄幼虫表皮中的表达分析[J].蚕业科学,2017,43(1):39-44.Li Q R,Xiao Y,Xin D X,et al.Expression analysis of chemosensory protein family genes in epidermis of the 5th instar silk?worm larvae[J].Science of Sericulture,2017,43(1):39-44.
[35]Guo Z,Cheng Z Y,Huang F,et al.Microarray analysis of global gene regulation in the Cry1Ab-resistant and Cry1Ab-susceptible strains of Diatraea saccharalis[J].Pest Management Science,2012,68(5):718-730.
[36]Liu Y L,Guo H,Huang L Q,et al.Unique function of a chemosensory protein in the proboscis of two Helicoverpa species[J].Journal of Experimental Biology,2014,217(10):1821-1826.
[37]Leal W S,Chen A M,Ishida Y,et al.Kinetics and molecular properties of pheromone binding and release[J].PNAS,2005,102(15):5386-5391.
[38]Clyne P J,Warr C G,Freeman M R,et al.A novel family of divergent seven-transmembrane proteins:candidate odorantreceptors in Drosophila[J].Neuron,1999,22(2):327-328.
[39]Larsson M C,Domingos A I,Jones W D,et al.Or83b encodes a broadly expressed odorant receptor essential for Drosophilaolfaction[J].Neuron,2004,43(5):703-714.
[40]Leal W S,Ishida Y,Pelletier J,et al.Olfactory proteins mediating chemical communication in the navel orangeworm moth,Amyelois transitella[J].PLoS One,2009,4(9):e7235.
[41]Vosshall L B,Hansson B S.A unified nomenclature system for the insect olfactory coreceptor[J].Chemical Senses,2011,36(6):497-498.
[42]Nakagawa T,Pellegrino M,Sato K,et al.Amino acid residues contributing to function of the heteromeric insect olfactoryreceptor complex[J].PLoS One,2012,7(3):e32372.
[43]Hildebrand J G.Analysis of chemical signals by nervous systems[J].PNAS,1995,92(1):67-74.
[44]Neuhaus E M,Gisselmann G,Zhang W,et al.Odorant receptor heterodimerization in the olfactory system of Drosophilamelanogaster[J].Nature Neuroscience,2005,8(1):15-17.
[45]Benton R,Sachse S,Michnick S W,et al.Atypical membrane topology and heteromeric function of Drosophila odorantreceptors in vivo[J].PLoS Biology,2006,4(2):240-257.
[46]代玉华,赵忠懿,程鹏,等.蚊虫嗅觉受体的研究进展[J].寄生虫与医学昆虫学报,2014,21(1):59-64.Dai Y H,Zhao Z Y,Cheng P,et al.Advances in research of odorant receptor proteins of mosquitoes[J].Acta Parasitologicaet Medica Entomologica Sinica,2014,21(1):59-64.
[47]Dobritsa A A,Naters W,Warr C G,et al.Integrating the molecular and cellular basis of odor coding in the Drosophilaantenna[J].Neuron,2003,37(5):827-841.
[48]Sakurai T,Nakagawa T,Mitsuno H,et al.Identification and functional characterization of a sex pheromone receptor in thesilkmoth Bombyx mori[J].PNAS,2004,101(47):16653-16658.
[49]Nakagawa T,Sakurai T,Nishioka T,et al.Insect sex-pheromone signals mediated by specific combinations of olfactoryreceptors[J].Science,2005,307(5715):1638-1642.
[50]Liu Y,Liu C,Lin K,et al.unctional specificity of sex pheromone receptors in the cotton bollworm Helicoverpa armigera[J].PLoS One,2013,8(4):e62094.
[51]Wanner K W,Nichols A S,Allen J E,et al.Sex pheromone receptor specificity in the European corn borer moth,Ostrinianubilalis[J].PLoS One,2010,5(1):e8685.
[52]Miura N,Nakagawa T,Touhara K,et al.Broadly and narrowly tuned odorant receptors are involved in female sex pheromonereception in Ostrinia moths[J].Insect Biochemistry and Molecular Biology,2010,40(1):64-73.
[53]Chang H,Liu Y,Yang T,et al.Pheromone binding proteins enhance the sensitivity of olfactory receptors to sex pheromonesin Chilo suppressalis[J].Scientific Reports,2015,5(1):13093.
[54]Benton R,Vannice K S,Gomez-Diaz C,et al.Variant ionotropic glutamate receptors as chemosensory receptors in Drosophila[J].Cell,2009,136(1):149-162.
[55]Montell C.A taste of the Drosophila gustatory receptors[J].Current Opinion in Neurobiology,2009,19(4):345-353.
[56]Koh T W,He Z,Gorur-Shandilya S,et al.The Drosophila IR20a clade of ionotropic receptors are candidate taste andpheromone receptors[J].Neuron,2014,83(4):850-865.
[57]Chen C,Buhl E,Xu M,et al.Drosophila Ionotropic Receptor 25a mediates circadian clock resetting by temperature[J].Nature,2015,527(7579):516-520.
[58]Croset V,Rytz R,Cummins S F,et al.Ancient protostome origin of chemosensory ionotropic glutamate receptors and theevolution of insect taste and olfaction[J].PLoS Genetics,2010,6(8):e1001064.
[59]Olivier V,Monsempes C,Fran?ois M C,et al.Candidate chemosensory ionotropic receptors in a Lepidoptera[J].InsectMolecular Biology,2011,20(2):189-199.
[60]Bengtsson J M,Trona F,MontagnéN,et al.Putative chemosensory receptors of the codling moth,Cydia pomonella,identifiedby antennal transcriptome analysis[J].PLoS One,2012,7(2):e31620.
[61]Liu Y,Gu S,Zhang Y,et al.Candidate olfaction genes identified within the Helicoverpa armigera antennal transcriptome[J].PLoS One,2012,7(10):e48260.
[62]Getahun M N,Wicher D,Hansson B S,et al.Temporal response dynamics of Drosophila olfactory sensory neurons dependson receptor type and response polarity[J].Frontiers in Cellular Neuroscience,2012,6(47):54-65.
[63]孙畅仪.小菜蛾气味受体和离子型受体的克隆与表达[D].哈尔滨:东北林业大学,2014.Sun C Y.Gene cloning and expression analysis of odorant receptors and ionotropic receptors in the Diamondback Moth(Plutella xylostella)[D].Harbin:Northeast Forestry University,2014.
[64]Su C Y,Menuz K,Carlson J R.Olfactory perception:receptors,cells,and circuits[J].Cell,2009,139(1):45-59.
[65]Rasmussen J T,Berglund L,Rasmussen M S,et al.Assignment of disulfide bridges in bovine CD36[J].European Journal ofBiochemistry,1998,257(2):488-494.
[66]Rogers M E,Sun M,Lerner M R,et al.Snmp-1,a novel membrane protein of olfactory neurons of the silk moth Antheraeapolyphemus with homology to the CD36 family of membrane proteins[J].Journal of Biological Chemistry,1997,272(23):14792-14799.
[67]Liu S,Zhang Y R,Zhou W W,et al.Identification and characterization of two sensory neuron membrane proteins fromCnaphalocrocis medinalis(Lepidoptera:Pyralidae)[J].Archives of Insect Biochemistry and Physiology,2013,82(1):29-42.
[68]Forstner M,Gohl T,Gondesen I,et al.Differential expression of SNMP-1 and SNMP-2 proteins in pheromone-sensitive hairsof moths[J].Chemical Senses,2008,33(3):291-299.
[69]罗梅浩,蔡永萍,赵艳艳,等.烟夜蛾感觉神经元膜蛋白基因的克隆、序列分析与组织特异性表达[J].中国烟草学报,2012,18(5):91-95.Luo M H,Cai Y P,Zhao Y Y,et al.Cloning,sequence analysis and tissue-specific expression of sensory neuron membraneprotein gene from Helicoverpa assulta(Guenée)[J].Acta Tabacaria Sinica,2012,18(5):91-95.
[70]Gu S H,Yang R N,Guo M B,et al.Molecular identification and differential expression of sensory neuron membrane proteinsin the antennae of the black cutworm moth Agrotis ipsilon[J].Journal of Insect Physiology,2013,59(4):430-443.
[71]Rogers M E,Krieger J,Vogt R G.Antennal SNMPs(sensory neuron membrane proteins)of lepidoptera define a uniquefamily of invertebrate CD36-like proteins[J].Journal of Neurobiology,2001,49(1):47-61.
[72]Vogt R G,Miller N E,Litvack R,et al.The insect SNMP gene family[J].Insect Biochemistry and Molecular Biology,2009,39(7):448-456.
[73]Kaissling K E.Olfactory perireceptor and receptor events in moths:a kinetic model revised[J].Journal of ComparativePhysiology A,2009,195(10):895-922.
[74]Merlin C,Fran?ois M C,Bozzolan F,et al.A new aldehyde oxidase selectively expressed in chemosensory organs of insects[J].Biochemical and Biophysical Research Communications,2005,332(1):4-10.
[75]Durand N,Carot-Sans G,Bozzolan F,et al.Degradation of pheromone and plant volatile components by a same odorant-degrading enzyme in the cotton leafworm,Spodoptera littoralis[J].PLoS One,2011,6(12):e29147.
[76]Ishida Y,Leaal W S.Chiral discrimination of the Japanese beetle sex pheromone and a behavioral antagonist by a pheromone-degrading enzyme[J].PNAS,2008,105(26):9076-9080.