异色瓢虫气味结合蛋白基因HaxyOBP1和HaxyOBP6的克隆及表达谱分析
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摘要
【目的】本研究旨在探索异色瓢虫Harmonia axyridis气味结合蛋白的结构和分布,更好地了解气味结合蛋白在异色瓢虫嗅觉系统中的作用。【方法】利用生物信息学方法克隆异色瓢虫2个气味结合蛋白基因序列并对其蛋白结构进行分析;通过实时荧光定量PCR分析克隆获得的这2个基因在异色瓢虫各个发育阶段和成虫不同组织中的表达水平。【结果】本研究成功克隆得到异色瓢虫两个气味结合蛋白基因HaxyOBP1和HaxyOBP6(GenBank登录号分别为MG757923和MG757927)。HaxyOBP1开放阅读框全长447 bp,编码148个氨基酸,具有6个保守的半胱氨酸,表明HaxyOBP1属于Classic OBPs。HaxyOBP1在异色瓢虫各发育阶段均有表达,在雄性成虫中表达量最高;组织表达谱分析表明HaxyOBP1在雄虫头部表达量最高。HaxyOBP6开放阅读框全长414 bp,编码137个氨基酸,具有4个保守的半胱氨酸,表明HaxyOBP6属于Minus-C OBPs。HaxyOBP6在成虫期的表达量明显高于幼虫阶段,并且在雄成虫中的表达量显著高于雌成虫;组织表达谱分析表明HaxyOBP6在雄成虫头部和雌成虫翅中表达量最高。【结论】本研究克隆得到异色瓢虫两个气味结合蛋白基因,表达谱分析表明HaxyOBP1和HaxyOBP6分别在异色瓢虫成虫头和翅中具有较高的表达水平,说明气味结合蛋白基因可能在异色瓢虫非嗅觉组织中同样具有重要作用。本研究结果为深入研究异色瓢虫气味结合蛋白结构和功能奠定了基础。
【Aim】 The study aims to explore the structure and distribution of odorant binding proteins of Harmonia axyridis, so as to provide a basis for further understanding the functions of odorant binding proteins in the olfactory system of H. axyridis. 【Methods】 Two odorant binding protein genes were cloned from H. axyridis, and their protein structures were analyzed by bioinformatics methods. The expression levels of the two genes in different developmental stages and adult tissues of H. axyridis were analyzed by qRT-PCR. 【Results】 Two odorant binding proteins genes of H.axyridis, HaxyOBP1 and HaxyOBP6(GenBank accession numbers: MG757923 and MG757927, respectively), were successfully cloned and sequenced. The open reading frame of HaxyOBP1 is 447 bp in length. It encodes 148 amino acids with six conserved cysteines, indicating that HaxyOBP1 belongs to Classical OBPs subfamily. HaxyOBP1 was expressed in various developmental stages of H. axyridis and had the highest expression level in male adults. Tissue expression profiles showed that HaxyOBP1 had the highest expression level in the head of male adult. The open reading frame of HaxyOBP6 is 414 bp in length. It encodes 137 amino acids with four conserved cysteines, indicating that HaxyOBP6 belongs to Minus-C OBPs subfamily. The expression level of HaxyOBP6 in adults was higher than that in larvae, and the expression level in male adults was significantly higher than that in female adults. Tissue expression profiles showed that HaxyOBP6 had the highest expression levels in the head of male adult and the wing of female adult. 【Conclusion】 In this study, two odorant binding protein genes were cloned from H. axyridis. The expression profiles showed that HaxyOBP1 and HaxyOBP6 were highly expressed in the head and the wing of adults, respectively, suggesting that the OBPs of H. axyridis may also play important roles in non-olfactory tissues. The results of this study lay a foundation for further study on the structure and function of OBPs of H. axyridis.
【Aim】 The study aims to explore the structure and distribution of odorant binding proteins of Harmonia axyridis, so as to provide a basis for further understanding the functions of odorant binding proteins in the olfactory system of H. axyridis. 【Methods】 Two odorant binding protein genes were cloned from H. axyridis, and their protein structures were analyzed by bioinformatics methods. The expression levels of the two genes in different developmental stages and adult tissues of H. axyridis were analyzed by qRT-PCR. 【Results】 Two odorant binding proteins genes of H.axyridis, HaxyOBP1 and HaxyOBP6(GenBank accession numbers: MG757923 and MG757927, respectively), were successfully cloned and sequenced. The open reading frame of HaxyOBP1 is 447 bp in length. It encodes 148 amino acids with six conserved cysteines, indicating that HaxyOBP1 belongs to Classical OBPs subfamily. HaxyOBP1 was expressed in various developmental stages of H. axyridis and had the highest expression level in male adults. Tissue expression profiles showed that HaxyOBP1 had the highest expression level in the head of male adult. The open reading frame of HaxyOBP6 is 414 bp in length. It encodes 137 amino acids with four conserved cysteines, indicating that HaxyOBP6 belongs to Minus-C OBPs subfamily. The expression level of HaxyOBP6 in adults was higher than that in larvae, and the expression level in male adults was significantly higher than that in female adults. Tissue expression profiles showed that HaxyOBP6 had the highest expression levels in the head of male adult and the wing of female adult. 【Conclusion】 In this study, two odorant binding protein genes were cloned from H. axyridis. The expression profiles showed that HaxyOBP1 and HaxyOBP6 were highly expressed in the head and the wing of adults, respectively, suggesting that the OBPs of H. axyridis may also play important roles in non-olfactory tissues. The results of this study lay a foundation for further study on the structure and function of OBPs of H. axyridis.
引文
Deng SS, 2011. Functional Analysis of Odorant-binding Proteins in Holotrichia oblita Faldermann (Coleoptera: Scarabaeida). MSc Thesis, Chinese Academy of Agricultural Sciences, Beijing. [邓思思, 2011. 华北大黑鳃金龟气味结合蛋白功能分析. 北京: 中国农业科学院硕士学位论文]
Dippel S, Oberhofer G, Kahnt J, Gerischer L, Opitz L, Schachtner J, Stanke M, Schütz S, Wimmer EA, Angeli S, 2014. Tissue-specific transcriptomics, chromosomal localization, and phylogeny of chemosensory and odorant binding proteins from the red flour beetle Tribolium castaneum reveal subgroup specificities for olfaction or more general functions. BMC Genomics, 15(1): 1141.
Du WM, Sun GZ, Zang LS, Wang XM, Ruan CC, Liu XJ, 2016. Effect of long-term cold storage on fecundity of pre-wintering Harmonia axyridis (Pallas) and the fitness of its progeny. J. Environ. Entomol., 38(2): 286-292. [杜文梅, 孙光芝, 臧连生, 王秀梅, 阮长春, 刘显娇, 2016. 长期冷藏对越冬代异色瓢虫繁殖及子代生长发育和捕食能力的影响. 环境昆虫学报, 38(2): 286-292]
Gu SH, Wu KM, Guo YY, Pickett JA, Field LM, Zhou JJ, Zhang YJ, 2013. Identification of genes expressed in the sex pheromone gland of the black cutworm Agrotis ipsilon with putative roles in sex pheromone biosynthesis and transport. BMC Genomics, 14(1): 636.
Gu SH, Zhang XY, Zhang YJ, Wu KM, Guo YY, 2010. Cloning and expression pattern analysis of an odorant binding protein gene Alin-OBP1 in the lucerne plant bug, Adelphocoris lineolatus (Goeze) (Hemiptera: Miridae). Acta Entomol. Sin., 53(5): 487-496. [谷少华, 张雪莹, 张永军, 吴孔明, 郭予元, 2010. 苜蓿盲蝽气味结合蛋白基因Alin-OBP1的克隆及表达谱分析. 昆虫学报, 53(5): 487-496]
Heidari M, Copland MJW, 1992. Host finding by Cryptolaemus montrouzieri (Col., Coccinellidae) a predator of mealybugs (Hom., Pseudococcidae). Entomophaga, 37(4): 621-625.
Koch RL, 2003. The multicolored Asian lady beetle, Harmonia axyridis: a review of its biology, uses in biological control, and non-target impacts. J. Insect Sci., 3: 32.
Li DZ, Yu GQ, Yi SC, Zhang Y, Kong DX, Wang MQ, 2015. Structure-based analysis of the ligand-binding mechanism for DhelOBP21, a C-minus odorant binding protein, from Dastarcus helophoroides (Fairmaire; Coleoptera: Bothrideridae). Int. J. Biol. Sci., 11(11): 1281-1295.
Li L, Zhou YT, Tan Y, Zhou XR, Pang BP, 2017. Identification of odorant-binding protein genes in Galeruca daurica (Coleoptera: Chrysomelidae) and analysis of their expression profiles. Bull. Entomol. Res., 107(4): 550-561.
Li W, Zhang YF, Xie YP, Niu XP, 2016. Selection response of Harmonia axyridis (Pallas) to body volatile of Ceroplastes japonicus Green. J. Environ. Entomol., 38(2): 329-336. [李伟, 张艳峰, 谢映平, 牛秀萍, 2016. 异色瓢虫对日本龟蜡蚧虫体挥发物的趋性反应. 环境昆虫学报, 38(2): 329-336]
Li XM, Zhu XY, Wang ZQ, Wang Y, He P, Chen G, Sun L, Deng DG, Zhang YN, 2015. Candidate chemosensory genes identified in Colaphellus bowringi by antennal transcriptome analysis. BMC Genomics, 16(1): 1028.
Li ZQ, Zhang S, Zhou SF, Luo JY, Cui JJ, 2017. Tissue expression profiling and ligand-binding properties of HarmOBP16 of the cotton bollworm, Helicoverpa armigera (Lepidoptera: Noctuidae). Acta Entomol. Sin., 60(8): 891-899. [李兆群, 张帅, 周淑芬, 雒珺瑜, 崔金杰, 2017. 棉铃虫气味结合蛋白HarmOBP16的组织表达谱及配基结合特性分析. 昆虫学报, 60(8): 891-899]
Liu S, Rao XJ, Li MY, Feng MF, He MZ, Li SG, 2015. Identification of candidate chemosensory genes in the antennal transcriptome of Tenebrio molitor (Coleoptera: Tenebrionidae). Comp. Biochem. Physiol. D, 13: 44-51.
Livak KJ, Schmittgen TD, 2001. Analysis of relative gene expression data using real-time quantitative PCR and the 2-ΔΔCT method. Methods, 25(4): 402-408.
Mombaerts P, 1999. Seven-transmembrane proteins as odorant and chemosensory receptors. Science, 286(5440): 707-711.
Niu DJ, Liu Y, Dong XT, Dong SL, 2016. Transcriptome based identification and tissue expression profiles of chemosensory genes in Blattella germanica (Blattaria: Blattidae). Comp. Biochem. Physiol. D, 18: 30-43.
Pan C, 2016. Cloning and Spatio-temporal Expression of the Odorant Binding Protein ComOBP1 and CmonOBP2 Genes from Cryptolaemus montrouzieri (Coleoptera: Coccinellidae). MSc Thesis, Sun Yat-sen University, Guangzhou. [潘畅, 2016. 孟氏隐唇瓢虫气味结合蛋白CmonOBP1和CmonOBP2基因的克隆和时空表达. 广州: 中山大学硕士学位论文]
Pan C, Zhang YH, Xie JQ, Li HS, Pang H, 2016. Cloning and spatio-temporal expression of the odorant binding protein ComOBP1 gene from Cryptolaemus montrouzieri Mulsant (Coleoptera: Coccinellidae). J. Environ. Entomol., 38(2): 249-260. [潘畅, 张宇宏, 谢佳沁, 李浩森, 庞虹, 2016. 孟氏隐唇瓢虫气味结合蛋白ComOBP1基因的克隆和时空表达. 环境昆虫学报, 38(2): 249-260]
Pelosi P, Maida R, 1995. Odorant-binding proteins in insects. Comp. Biochem. Physiol. B, 111(3): 503-514.
Teng SB, Xu ZQ, 2005. The cold storage conditions of Harmonia axyridis eggs and adults in mass-rearing. Chin. Bull. Entomol., 42(2): 180-183. [滕树兵, 徐志强, 2005. 人工扩繁代异色瓢虫卵和成虫最适冷藏条件的探讨. 昆虫知识, 42(2): 180-183]
Wang GR, Guo YY, Wu KM, 2002. Progress in the studies of antenna odorant binding proteins of insects. Acta Entomol. Sin., 45(1): 131-137. [王桂荣, 郭予元, 吴孔明, 2002. 昆虫触角气味结合蛋白的研究进展. 昆虫学报, 45(1): 131-137]
Wang HT, Zhang WD, Chen XZ, Zheng JF, Miao L, Qin QL, 2012. Mass rearing the multicolored Asian lady beetle on beet army worm larvae. Chin. J. Appl. Entomol., 49(6): 1726-1731. [王红托, 张伟东, 陈新中, 郑建峰, 苗麟, 秦启联, 2012. 异色瓢虫规模化生产技术及瓢虫工厂的建立. 应用昆虫学报, 49(6): 1726-1731]
Wang LY, 1986. Mass rearing and utilization in biological control of the lady beetle Leis axyridis (Palla). Acta Entomol. Sin., 29(1): 104. [王良衍, 1986. 异色瓢虫的人工饲养及野外释放和利用. 昆虫学报, 29(1): 104]
Wang YP, Lü F, Wang ZP, 2007. Progress of Harmonia axyridi (Pallas) utilization. Entomol. J. East China, 16(4): 310-314. [王延鹏, 吕飞, 王振鹏, 2007. 异色瓢虫开发利用研究进展. 华东昆虫学报, 16(4): 310-314]
Wei D, Ye ZF, Gao JQ, Dong SL, 2013. Molecular cloning and functional identification of a Minus-C odorant binding protein from the rice striped stem borer, Chilo suppressalis (Lepidoptera: Pyralidae). Acta Entomol. Sin., 56(7): 754-764. [魏丹, 叶占峰, 高建清, 董双林, 2013. 二化螟Minus-C气味结合蛋白的分子克隆及功能鉴定. 昆虫学报, 56(7): 754-764]
Xue JL, He J, Xie YP, 2008. Attractive effect of plant volatiles on Harmonia axyridis (Pallas). Chin. J. Appl. Environ. Biol., 14(4): 494-498. [薛皎亮, 贺珺, 谢映平, 2008. 植物挥发物对天敌昆虫异色瓢虫的引诱效应. 应用与环境生物学报, 14(4): 494-498]
Yang JC, Shen ZR, 2000. Effect of cold storage on survival of the ladybird beetles, Leis axyridis and Coccoinella septempunctata. Acta Entomol. Sin., 43(S1): 211-214. [杨俊成, 沈佐锐, 2000. 冷藏条件对瓢虫存活的影响. 昆虫学报, 43(S1): 211-214]
Zeng B, 2013. Studies on Mass Rearing and Field Release Technology of Harmonia axyridis. MSc Thesis, Huazhong Agricultural University, Wuhan. [曾斌, 2013. 异色瓢虫人工大量繁殖与田间释放技术研究. 武汉: 华中农业大学硕士学位论文]
Zhang J, Zhang L, 2006. Type and distribution of forewings sensilla of Locusta migratoria manilensis (Insecta, Orthoptera). Acta Agric. Bor. Sin., 21(S1): 122-126. [张杰, 张龙, 2006. 东亚飞蝗前翅感受器的类型与分布初步研究. 华北农学报, 21(S1): 122-126]
Zhou JJ, 2010. Odorant-binding proteins in insects. Vitam. Horm., 83: 241-272.
Zhuang XJ, Yin J, Li KB, Cao YZ, 2013. Bioinformatics analysis of the odorant-binding protein HoblOBP2 in olfactory sensilla of the scarab beetle Holotrichia oblita. Plant Prot., 39(1): 50-55. [庄绪静, 尹姣, 李克斌, 曹雅忠, 2013. 华北大黑鳃金龟气味结合蛋白HoblOBP2的生物信息学分析. 植物保护, 39(1): 50-55]
Dippel S, Oberhofer G, Kahnt J, Gerischer L, Opitz L, Schachtner J, Stanke M, Schütz S, Wimmer EA, Angeli S, 2014. Tissue-specific transcriptomics, chromosomal localization, and phylogeny of chemosensory and odorant binding proteins from the red flour beetle Tribolium castaneum reveal subgroup specificities for olfaction or more general functions. BMC Genomics, 15(1): 1141.
Du WM, Sun GZ, Zang LS, Wang XM, Ruan CC, Liu XJ, 2016. Effect of long-term cold storage on fecundity of pre-wintering Harmonia axyridis (Pallas) and the fitness of its progeny. J. Environ. Entomol., 38(2): 286-292. [杜文梅, 孙光芝, 臧连生, 王秀梅, 阮长春, 刘显娇, 2016. 长期冷藏对越冬代异色瓢虫繁殖及子代生长发育和捕食能力的影响. 环境昆虫学报, 38(2): 286-292]
Gu SH, Wu KM, Guo YY, Pickett JA, Field LM, Zhou JJ, Zhang YJ, 2013. Identification of genes expressed in the sex pheromone gland of the black cutworm Agrotis ipsilon with putative roles in sex pheromone biosynthesis and transport. BMC Genomics, 14(1): 636.
Gu SH, Zhang XY, Zhang YJ, Wu KM, Guo YY, 2010. Cloning and expression pattern analysis of an odorant binding protein gene Alin-OBP1 in the lucerne plant bug, Adelphocoris lineolatus (Goeze) (Hemiptera: Miridae). Acta Entomol. Sin., 53(5): 487-496. [谷少华, 张雪莹, 张永军, 吴孔明, 郭予元, 2010. 苜蓿盲蝽气味结合蛋白基因Alin-OBP1的克隆及表达谱分析. 昆虫学报, 53(5): 487-496]
Heidari M, Copland MJW, 1992. Host finding by Cryptolaemus montrouzieri (Col., Coccinellidae) a predator of mealybugs (Hom., Pseudococcidae). Entomophaga, 37(4): 621-625.
Koch RL, 2003. The multicolored Asian lady beetle, Harmonia axyridis: a review of its biology, uses in biological control, and non-target impacts. J. Insect Sci., 3: 32.
Li DZ, Yu GQ, Yi SC, Zhang Y, Kong DX, Wang MQ, 2015. Structure-based analysis of the ligand-binding mechanism for DhelOBP21, a C-minus odorant binding protein, from Dastarcus helophoroides (Fairmaire; Coleoptera: Bothrideridae). Int. J. Biol. Sci., 11(11): 1281-1295.
Li L, Zhou YT, Tan Y, Zhou XR, Pang BP, 2017. Identification of odorant-binding protein genes in Galeruca daurica (Coleoptera: Chrysomelidae) and analysis of their expression profiles. Bull. Entomol. Res., 107(4): 550-561.
Li W, Zhang YF, Xie YP, Niu XP, 2016. Selection response of Harmonia axyridis (Pallas) to body volatile of Ceroplastes japonicus Green. J. Environ. Entomol., 38(2): 329-336. [李伟, 张艳峰, 谢映平, 牛秀萍, 2016. 异色瓢虫对日本龟蜡蚧虫体挥发物的趋性反应. 环境昆虫学报, 38(2): 329-336]
Li XM, Zhu XY, Wang ZQ, Wang Y, He P, Chen G, Sun L, Deng DG, Zhang YN, 2015. Candidate chemosensory genes identified in Colaphellus bowringi by antennal transcriptome analysis. BMC Genomics, 16(1): 1028.
Li ZQ, Zhang S, Zhou SF, Luo JY, Cui JJ, 2017. Tissue expression profiling and ligand-binding properties of HarmOBP16 of the cotton bollworm, Helicoverpa armigera (Lepidoptera: Noctuidae). Acta Entomol. Sin., 60(8): 891-899. [李兆群, 张帅, 周淑芬, 雒珺瑜, 崔金杰, 2017. 棉铃虫气味结合蛋白HarmOBP16的组织表达谱及配基结合特性分析. 昆虫学报, 60(8): 891-899]
Liu S, Rao XJ, Li MY, Feng MF, He MZ, Li SG, 2015. Identification of candidate chemosensory genes in the antennal transcriptome of Tenebrio molitor (Coleoptera: Tenebrionidae). Comp. Biochem. Physiol. D, 13: 44-51.
Livak KJ, Schmittgen TD, 2001. Analysis of relative gene expression data using real-time quantitative PCR and the 2-ΔΔCT method. Methods, 25(4): 402-408.
Mombaerts P, 1999. Seven-transmembrane proteins as odorant and chemosensory receptors. Science, 286(5440): 707-711.
Niu DJ, Liu Y, Dong XT, Dong SL, 2016. Transcriptome based identification and tissue expression profiles of chemosensory genes in Blattella germanica (Blattaria: Blattidae). Comp. Biochem. Physiol. D, 18: 30-43.
Pan C, 2016. Cloning and Spatio-temporal Expression of the Odorant Binding Protein ComOBP1 and CmonOBP2 Genes from Cryptolaemus montrouzieri (Coleoptera: Coccinellidae). MSc Thesis, Sun Yat-sen University, Guangzhou. [潘畅, 2016. 孟氏隐唇瓢虫气味结合蛋白CmonOBP1和CmonOBP2基因的克隆和时空表达. 广州: 中山大学硕士学位论文]
Pan C, Zhang YH, Xie JQ, Li HS, Pang H, 2016. Cloning and spatio-temporal expression of the odorant binding protein ComOBP1 gene from Cryptolaemus montrouzieri Mulsant (Coleoptera: Coccinellidae). J. Environ. Entomol., 38(2): 249-260. [潘畅, 张宇宏, 谢佳沁, 李浩森, 庞虹, 2016. 孟氏隐唇瓢虫气味结合蛋白ComOBP1基因的克隆和时空表达. 环境昆虫学报, 38(2): 249-260]
Pelosi P, Maida R, 1995. Odorant-binding proteins in insects. Comp. Biochem. Physiol. B, 111(3): 503-514.
Teng SB, Xu ZQ, 2005. The cold storage conditions of Harmonia axyridis eggs and adults in mass-rearing. Chin. Bull. Entomol., 42(2): 180-183. [滕树兵, 徐志强, 2005. 人工扩繁代异色瓢虫卵和成虫最适冷藏条件的探讨. 昆虫知识, 42(2): 180-183]
Wang GR, Guo YY, Wu KM, 2002. Progress in the studies of antenna odorant binding proteins of insects. Acta Entomol. Sin., 45(1): 131-137. [王桂荣, 郭予元, 吴孔明, 2002. 昆虫触角气味结合蛋白的研究进展. 昆虫学报, 45(1): 131-137]
Wang HT, Zhang WD, Chen XZ, Zheng JF, Miao L, Qin QL, 2012. Mass rearing the multicolored Asian lady beetle on beet army worm larvae. Chin. J. Appl. Entomol., 49(6): 1726-1731. [王红托, 张伟东, 陈新中, 郑建峰, 苗麟, 秦启联, 2012. 异色瓢虫规模化生产技术及瓢虫工厂的建立. 应用昆虫学报, 49(6): 1726-1731]
Wang LY, 1986. Mass rearing and utilization in biological control of the lady beetle Leis axyridis (Palla). Acta Entomol. Sin., 29(1): 104. [王良衍, 1986. 异色瓢虫的人工饲养及野外释放和利用. 昆虫学报, 29(1): 104]
Wang YP, Lü F, Wang ZP, 2007. Progress of Harmonia axyridi (Pallas) utilization. Entomol. J. East China, 16(4): 310-314. [王延鹏, 吕飞, 王振鹏, 2007. 异色瓢虫开发利用研究进展. 华东昆虫学报, 16(4): 310-314]
Wei D, Ye ZF, Gao JQ, Dong SL, 2013. Molecular cloning and functional identification of a Minus-C odorant binding protein from the rice striped stem borer, Chilo suppressalis (Lepidoptera: Pyralidae). Acta Entomol. Sin., 56(7): 754-764. [魏丹, 叶占峰, 高建清, 董双林, 2013. 二化螟Minus-C气味结合蛋白的分子克隆及功能鉴定. 昆虫学报, 56(7): 754-764]
Xue JL, He J, Xie YP, 2008. Attractive effect of plant volatiles on Harmonia axyridis (Pallas). Chin. J. Appl. Environ. Biol., 14(4): 494-498. [薛皎亮, 贺珺, 谢映平, 2008. 植物挥发物对天敌昆虫异色瓢虫的引诱效应. 应用与环境生物学报, 14(4): 494-498]
Yang JC, Shen ZR, 2000. Effect of cold storage on survival of the ladybird beetles, Leis axyridis and Coccoinella septempunctata. Acta Entomol. Sin., 43(S1): 211-214. [杨俊成, 沈佐锐, 2000. 冷藏条件对瓢虫存活的影响. 昆虫学报, 43(S1): 211-214]
Zeng B, 2013. Studies on Mass Rearing and Field Release Technology of Harmonia axyridis. MSc Thesis, Huazhong Agricultural University, Wuhan. [曾斌, 2013. 异色瓢虫人工大量繁殖与田间释放技术研究. 武汉: 华中农业大学硕士学位论文]
Zhang J, Zhang L, 2006. Type and distribution of forewings sensilla of Locusta migratoria manilensis (Insecta, Orthoptera). Acta Agric. Bor. Sin., 21(S1): 122-126. [张杰, 张龙, 2006. 东亚飞蝗前翅感受器的类型与分布初步研究. 华北农学报, 21(S1): 122-126]
Zhou JJ, 2010. Odorant-binding proteins in insects. Vitam. Horm., 83: 241-272.
Zhuang XJ, Yin J, Li KB, Cao YZ, 2013. Bioinformatics analysis of the odorant-binding protein HoblOBP2 in olfactory sensilla of the scarab beetle Holotrichia oblita. Plant Prot., 39(1): 50-55. [庄绪静, 尹姣, 李克斌, 曹雅忠, 2013. 华北大黑鳃金龟气味结合蛋白HoblOBP2的生物信息学分析. 植物保护, 39(1): 50-55]