昆虫气味受体研究技术及其在林业昆虫中的应用研究进展

doi:10.11707/j.1001-7488.20200517

摘要/Abstract

摘要：

嗅觉在昆虫的生存和种群繁衍过程中起着重要的作用，嗅觉感受神经元的多种蛋白质共同参与嗅觉识别过程，包括：气味结合蛋白（OBP）、气味受体（OR）、气味降解酶（ODE）、化学感受蛋白（CSP）和感受神经元膜蛋白（SNMP）等。OR作为其中一种重要的蛋白质，其所介导的气味分子与嗅觉神经元的专一性结合是昆虫行使嗅觉功能的重要基础。由于昆虫OR结构与其他脊椎动物OR结构不同，一直不能通过同源克隆的方式获得，其发现经历了一段漫长的时间。昆虫OR可以分为2大类，第1类是识别各种气味分子的功能受体，第2类是没有气味分子识别功能，与其他功能受体共表达，协助其他功能受体完成对气味分子的识别。关于OR的基础研究，主要集中在鉴定和功能分析方面。OR的功能分析目前常用的有3种体外表达系统，分别是爪蟾卵母细胞表达系统、异源细胞表达系统和果蝇空神经元表达系统。RNAi和Crispr/Cas9作为活体研究基因功能的方法，也在某些物种中用于OR基因的功能研究。昆虫OR作用机制研究目前主要集中在一些模式物种中，林业昆虫作为一类重要的昆虫，其OR研究起步较晚，且主要集中在OR基因鉴定上，功能研究相对较少。气味受体的研究为害虫嗅觉调控和种群控制提供了一个很好的研究方向，值得进一步加强机制研究和防控技术开发。

关键词: 气味受体, 基因功能, 研究技术, 森林昆虫, 森林害虫防治

Abstract:

Olfaction plays an important role in survival and reproduction of insects. Many proteins from olfactory sensory neurons participate in the olfactory recognition process,including odorant binding protein (OBP),odorant receptor (OR),odorant degrading enzyme (ODE),chemosensory protein (CSP),and sensory neuron membrane protein (SNMP). As an important protein,OR mediates the specific combination of odour molecules and olfactory neurons,which is an important basis for insects to perform olfactory functions. Because the OR structure of insects is different from that of vertebrates,it cannot be obtained by homologous cloning for long time. Thus,ORs discovery of insects has taken a long time. ORs can be divided into two categories. The first category is the functional receptors to recognize various odour molecules,the second one is the receptor without odour molecules recognition function,which co-expresses with other functional receptors to perform olfactory functions together. Researches on OR mainly focus on its identification and functional exploration. At present,there are three expression systems in vitro commonly used in the functional analyses of OR,including the Xenopus oocyte expression system,the heterologous cell expression system,and the Drosophila empty neuron expression system. As for in vivo methods for researching the gene functions,RNAi (RNA interference,RNAi) and Crispr/Cas9 (clustered regularly interspaced short palindromic repeats/Crispr-associated sequence) systems have also been used to explore the OR's function in some insects. Researches on the mechanism of insects OR mainly focus on some model insects. Although forest insects are important insects,their OR research started late. At present,researches about OR in forest insect are mainly gene identification,and only few work is on gene function. The research on the olfactory receptor mechanism provides a good direction for pest control,and it is worth to further strengthen researches about the mechanisms and pest control methods based on olfactory regulation.

Key words: odorant receptors, gene function, research techniques, forest insects, forest pest control

中图分类号:

S718.7

申思凡,张真,孔祥波,刘福,张苏芳. 昆虫气味受体研究技术及其在林业昆虫中的应用研究进展[J]. 林业科学, 2020, 56(5): 150-159.

Sifan Shen,Zhen Zhang,Xiangbo Kong,Fu Liu,Sufang Zhang. Research Techniques of Insect Odour Receptors and Their Application in Forest Insects[J]. Scientia Silvae Sinicae, 2020, 56(5): 150-159.

图/表 1

表1

农林类害虫气味受体鉴定和功能研究情况"

分类目科 Classification		物种 Species	气味受体 OR name	基因表达系统 Gene expression system	参考文献 References
鳞翅目 Lepidoptera	菜蛾科 Plutellidae	小菜蛾Plutella xylostella	OR1, 3	Xo	Mitsuno et al., 2008
			Or83b		李珣等，2011
			OR4	Xo	Sun et al., 2013
			OR9	Xo	刘一鹏等，2015
			OR16, 17, 18		孔畅仪等，2014
			OR8, 41, 45	Xo	Liu et al., 2018
	灯蛾科 Arctiidae	美国白蛾Hyphantria cunea	52 candidate OR genes		Zhang et al., 2016
	卷蛾科Tortricidae	苹淡褐卷蛾Epiphyas postvittana	E.pos OR1-3	Sf9	Jordan et al., 2009
			Orco and OR3	HEK	Corcoran et al., 2014
			70 candidate OR genes		Corcoran et al., 2015
		苹果蠹蛾Cydia pomonella	43 putative OR genes		Bengtsson et al., 2012
			OR3	DENES	Bengtsson et al., 2014
			52 candidate OR genes		Walker et al., 2016
		梨小食心虫Grapholita molesta	48 candidate OR genes		Li et al., 2015
	枯叶蛾科 Lasiocampidae	云南松毛虫Dendrolimus houi	33 candidate OR genes		Zhang et al., 2014
		思茅松毛虫Dendrolimus kekuchii	33 candidate OR genes		Zhang et al., 2014
		马尾松毛虫Dendrolimus punctatus	60 candidate OR genes		Zhang et al., 2017
		马尾松毛虫Dendrolimus punctatus	68 candidate OR genes		Zhang et al., 2018
	螟蛾科 Pyralidae	瓜绢野螟Diaphania indica	OR1, 3	Xo	Mitsuno et al., 2008
	螟蛾科 Pyralidae	玉米螟Ostrinia nubilalis	O.nubOR1, OR3-6	Xo	Wanner et al., 2010
	天蛾科Sphingidae	烟草天蛾Manduca sexta	OR1, 2, 3		Patch et al., 2009
	天蛾科Sphingidae	烟草天蛾Manduca sexta	M.sexOR1-6		Grosse-Wilde et al., 2010
	夜蛾科 Noctuidae	小地老虎 Agrotis ipsilon	42 OR genes annotated, 35 ORs expressed		Gu et al., 2014
		棉铃虫Helicoverpa armigera	OR1, 2, 3		Zhang et al., 2010
			47 putative OR genes		Liu et al., 2012
			OR9, 29		刘宁灿等, 2014
			11 OR genes	Xo	Jiang et al.,2014
			60 OR genes		Zhang et al., 2015
			OR12	Xo	Cao et al., 2016
			OR16		Chang et al., 2017
			17 OR genes		Di et al.,2017
		烟青虫Helicoverpa assulta	OR1, 2, 3		Zhang et al.,2010
			OR18		张元臣等，2013
			10 OR genes	Xo	Jiang et al., 2014
			64 putative OR genes		Zhang et al., 2015
			OR12	Xo	Cao et al., 2016
			OR24, 40, 41, 55		Cui et al., 2018
		烟芽夜蛾Heliothis virescens	H.vir OR6, 11-16		Krieger et al., 2004
			OR13		Gohl et al., 2006
			OR13, 14, 16	HEK	Grosse-Wilde et al., 2007
			OR13, 15		Soques et al., 2010
			OR6, 14, 15, 16		Vásquez et al., 2011
			OR6, 13, 14, 16	Xo	Wang et al., 2011
			OR12	Xo	Cao et al., 2016
		Heliothis subflexa	OR13, 15		Soques et al., 2010
		Heliothis subflexa	OR6, 14, 15, 16		Vásquez et al., 2011
		黏虫Mythimna separate	OR1, 3	Xo	Mitsuno et al., 2008
			60 OR genes		Liu et al., 2017
			71 OR genes		Chang et al., 2017
		甜菜夜蛾Spodoptera exigua	OR2		张逸凡等，2011
			S.exiOR6, 11, 13, 16	Xo	Liu et al., 2013
			OR18		刘程程等，2013
			OR3	Xo	Liu et al., 2014
			10 OR genes		Liu et al., 2015
			64 OR genes		Zhang et al., 2018
		斜纹夜蛾Spodoptera litura	OR2		陈茜等，2011
			OR12, 19, 44, 51	Xo	Zhang et al., 2013
			OR18		Zheng et al., 2014
			S.litOR6, 11, 13, 16	Xo	Zhang et al., 2015
			OR1		Zhang et al., 2017
			26 OR genes		Li et al., 2015
		灰翅夜蛾Spodoptera littorali	S.litt OR6	DENES	Montagné et al., 2012
			36 OR genes		Jacquin et al., 2012
			47 candidate ORs	Xo	de Fouchier et al., 2017
鞘翅目 Coleoptera	吉丁甲科 Buprestidae	白蜡窄吉丁Agrilus planipennis	Orco and OR64		Mamidala et al., 2013
	叶甲科 Chrysomelidae	紫榆叶甲Ambrostoma quadriimpressum	18 OR genes		Wang et al., 2016
	天牛科 Cerambycidae	胡桃胭脂天牛Meyacyllene caryae	57 OR genes		Mitchell et al., 2012
		光肩星天牛Anoplophora glabripennis	37 OR genes		Hu et al., 2016
			132 OR genes		Mckenna et al., 2016
			132 OR genes		Mitchell et al., 2017
	小蠹科 Scolytidae	云杉八齿小蠹Ips typographus	43 OR genes		Andersson et al., 2013
		中欧山松大小蠹Dendroctomus ponderosae	49 OR genes		Andersson et al., 2013
		红脂大小蠹Dendroctonus valens	22 OR genes		Gu et al., 2015
		云南切梢小蠹Tomicus yunnanensis	9 OR genes		Liu et al., 2018

表1

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