云斑天牛成虫聚集、交配行为的初步研究
|
摘要
云斑天牛Batocera horsfieldi(Hope)是一种大型多食性蛀干害虫,由于其生活隐蔽,抗逆性强而防治困难,成为许多天然林、经济林、城市园林树木的毁灭性害虫。目前,对其主要采用效率较低的人工和化学防治,防治效果不甚理想,对云斑天牛成虫间信息联系机制的研究还未有涉及。因此,加强成虫间信息联系机制方面的研究,将为开发高活性引诱剂,诱捕成虫,集中治理,初步建立云斑天牛的持续控制技术体系奠定基础。本文从化学生态学角度研究云斑天牛成虫之间的信息联系机制。主要结论如下:
1.采用林间观察和室内实验相结合的方法,对云斑天牛的聚集、交配行为进行研究。发现:1)云斑天牛成虫远距离的聚集是靠补充营养寄主树木的引诱作用,雌雄虫到达寄主树木后对寄主特定区段的趋性使其完成近距离内的聚集;2)云斑天牛成虫的交配行为大都是在雄虫接触到雌虫后,便立即快速爬向雌体的表现表明,在交配活动中,雌虫信息素对雄虫的接触性刺激作用非常重要。雄虫用触角识别雌虫后,便进行抱对骑乘,无明显的求偶行为。在抱对骑乘阶段,雄虫会用下颚须、下唇须不断的舐触雌虫的前胸背板和鞘翅基部,进而曲腹、插入内囊输精完成交配;3)寄主植物存在与否对云斑天牛交配行为有重要影响,当有寄主植物时,交配持续时间和交配成功率都显著高于无寄主植物时的交配成功率和交配持续时间。
2.利用Y型嗅觉仪测定云斑天牛雄虫对雌虫浸提液的嗅觉趋性反应及通过雄虫对雌虫浸提液的接触感应生测实验。实验结果初步表明:1)雄虫对雌性信息素提取物的气味反应不明显,即通过非接触性的嗅觉感应雄虫很难感受到雌性提取物的存在;2)雌性信息素提取物中对雄虫产生关键性引诱作用的物质的挥发性很弱或不具有挥发性;3)在识别配偶过程中,接触性的嗅觉判断起主导作用,云斑天牛雌性成虫的性信息素为接触性信息素,雄虫依靠触角、下唇须和下颚须上化感器的接触感应来接收雌成虫产生的接触性信息素的刺激。
3.详细描述了触角、下颚须和下唇须的外部形态,运用扫描电子显微镜对触角、下颚须及下唇须上的化学感器进行了观察,发现触角上具8类感器,下颚须和下唇须上具有4类感器。云斑天牛成虫触角感受器在种类、数量和分布上存在性别差异,雌雄虫的下颚须和下唇须感受器数量有所不同。其中刺形感器为机械感受器,锥形感器和毛形感器具嗅觉功能,具沟锥形感器和拴锥形感器具味觉功能。
Batocera horsfieldi (Hope) is a kind of polyphagous woodworm pest with large-scale. It is difficult to control for the characteristics of hidden life and strong resistence. Therefore, they become the main destructive pest to many kinds of natural forests, economy trees and urban garden trees, cost great losses of the economy and ecology in China. Nowadays, the main method of preventing B. horsfieldi is always by mankind with low-efficient or chemical means. By contrast, less researches refered to the information communication mechanisms between B. horsfieldi male and female adults. In order to development of high-activity attractants, mass-trapping, centralize-processing of adults, and establishment of the sustainable control system of B. horsfieldi initially, it is important to study the chemical information communication mechanism between B. horsfieldi male and female adults based on chemical ecology. The results are summarized as follows:
1. Aggregation and mating behavior of the forest borer, B. horsfieldi was observed both in the laboratory and field. The results showed that: 1) Adults aggregating on the trees that are the host plants, feed on the phloem of branch; 2) Contact sex pheromone should be accumulated on the body of female B. horsfieldi. Males differentiated between sexes by touching. When male and female of B. horsfieldi encountered, the male was observed to approach or hold the female only when he touched her with his antennae. The male was observed to hold and mount the female, during holding and mounting, the male frequently touched the female elytra and pronotum with his labipalpi and maxipalpi, and then to bend his abdomen toward her abdominal tip, ejaculation sequentially; 3) The host plants had evident influence on mating behavior of B. horsfieldi. When adults mating on the host plants, duration of mating behavior is significantly longer than the one which mating in the clearing, and the efficiency of mating is significantly increase.
2. The extracts from females were tested, checks up the adults' induction tropism and behavior reaction to the odor or flavour trail of the male. The results are as follows: The distinguishing and estimation abilities of B. horsfieldi adults' olfaction without contacting are not the leading factors to look for the mates. The mating action of adults generally started by touching. The sex pheromone of female adults is a contacting pheromone. The male adults receive the signal stimulation by the contacting inductor of the chemical inductors on the antennae, labipalpi and maxipalpi.
3. The external morphology of antennal、labialpalpus and maxillarypalpus sensilla was observed by SEM. Eight types of antennal sensilla and four types of labipalpi and maxipalpi sensilla were found. There are diferences in number, type and location of antenna senilla between male and female. But only in number of labialpalpus and maxillarypalpus senilla. Chaetic sensilla was considered to be a mechanoreceptor. Basieonie sensilla and Trichodea sensilla were considered to be olfactory receptors. Colpate basieonie sensillum and Styloconic sensillum might be a sensor of contact chemicals.
1.采用林间观察和室内实验相结合的方法,对云斑天牛的聚集、交配行为进行研究。发现:1)云斑天牛成虫远距离的聚集是靠补充营养寄主树木的引诱作用,雌雄虫到达寄主树木后对寄主特定区段的趋性使其完成近距离内的聚集;2)云斑天牛成虫的交配行为大都是在雄虫接触到雌虫后,便立即快速爬向雌体的表现表明,在交配活动中,雌虫信息素对雄虫的接触性刺激作用非常重要。雄虫用触角识别雌虫后,便进行抱对骑乘,无明显的求偶行为。在抱对骑乘阶段,雄虫会用下颚须、下唇须不断的舐触雌虫的前胸背板和鞘翅基部,进而曲腹、插入内囊输精完成交配;3)寄主植物存在与否对云斑天牛交配行为有重要影响,当有寄主植物时,交配持续时间和交配成功率都显著高于无寄主植物时的交配成功率和交配持续时间。
2.利用Y型嗅觉仪测定云斑天牛雄虫对雌虫浸提液的嗅觉趋性反应及通过雄虫对雌虫浸提液的接触感应生测实验。实验结果初步表明:1)雄虫对雌性信息素提取物的气味反应不明显,即通过非接触性的嗅觉感应雄虫很难感受到雌性提取物的存在;2)雌性信息素提取物中对雄虫产生关键性引诱作用的物质的挥发性很弱或不具有挥发性;3)在识别配偶过程中,接触性的嗅觉判断起主导作用,云斑天牛雌性成虫的性信息素为接触性信息素,雄虫依靠触角、下唇须和下颚须上化感器的接触感应来接收雌成虫产生的接触性信息素的刺激。
3.详细描述了触角、下颚须和下唇须的外部形态,运用扫描电子显微镜对触角、下颚须及下唇须上的化学感器进行了观察,发现触角上具8类感器,下颚须和下唇须上具有4类感器。云斑天牛成虫触角感受器在种类、数量和分布上存在性别差异,雌雄虫的下颚须和下唇须感受器数量有所不同。其中刺形感器为机械感受器,锥形感器和毛形感器具嗅觉功能,具沟锥形感器和拴锥形感器具味觉功能。
Batocera horsfieldi (Hope) is a kind of polyphagous woodworm pest with large-scale. It is difficult to control for the characteristics of hidden life and strong resistence. Therefore, they become the main destructive pest to many kinds of natural forests, economy trees and urban garden trees, cost great losses of the economy and ecology in China. Nowadays, the main method of preventing B. horsfieldi is always by mankind with low-efficient or chemical means. By contrast, less researches refered to the information communication mechanisms between B. horsfieldi male and female adults. In order to development of high-activity attractants, mass-trapping, centralize-processing of adults, and establishment of the sustainable control system of B. horsfieldi initially, it is important to study the chemical information communication mechanism between B. horsfieldi male and female adults based on chemical ecology. The results are summarized as follows:
1. Aggregation and mating behavior of the forest borer, B. horsfieldi was observed both in the laboratory and field. The results showed that: 1) Adults aggregating on the trees that are the host plants, feed on the phloem of branch; 2) Contact sex pheromone should be accumulated on the body of female B. horsfieldi. Males differentiated between sexes by touching. When male and female of B. horsfieldi encountered, the male was observed to approach or hold the female only when he touched her with his antennae. The male was observed to hold and mount the female, during holding and mounting, the male frequently touched the female elytra and pronotum with his labipalpi and maxipalpi, and then to bend his abdomen toward her abdominal tip, ejaculation sequentially; 3) The host plants had evident influence on mating behavior of B. horsfieldi. When adults mating on the host plants, duration of mating behavior is significantly longer than the one which mating in the clearing, and the efficiency of mating is significantly increase.
2. The extracts from females were tested, checks up the adults' induction tropism and behavior reaction to the odor or flavour trail of the male. The results are as follows: The distinguishing and estimation abilities of B. horsfieldi adults' olfaction without contacting are not the leading factors to look for the mates. The mating action of adults generally started by touching. The sex pheromone of female adults is a contacting pheromone. The male adults receive the signal stimulation by the contacting inductor of the chemical inductors on the antennae, labipalpi and maxipalpi.
3. The external morphology of antennal、labialpalpus and maxillarypalpus sensilla was observed by SEM. Eight types of antennal sensilla and four types of labipalpi and maxipalpi sensilla were found. There are diferences in number, type and location of antenna senilla between male and female. But only in number of labialpalpus and maxillarypalpus senilla. Chaetic sensilla was considered to be a mechanoreceptor. Basieonie sensilla and Trichodea sensilla were considered to be olfactory receptors. Colpate basieonie sensillum and Styloconic sensillum might be a sensor of contact chemicals.
引文
[1]严敖金,嵇保中,钱范俊.云斑天牛Batocera horsfieldi(Hope)的研究.南京林业大学学报,1997,21(1):1-6.
[2]Nakamuta K,Leal WS,Nakashima T.Increase of trap catches by a combination of male sex pheromones and floral attractant in longhonr beetle,Anaglyptus subfasciatus.Chem Ecol,1997,23:1635-1640.
[3]嵇保中,魏勇,黄振裕.天牛成虫研究的现状与展望.南京林业大学学报(自然科学版),2002,26(2):79-83.
[4]Ginzel MD,Blomquist GJ,Millar JG,LM Hanks.Role of contact pheromones in mate recognition in Xylotrechus colonus.Chem Ecol,2003a,129(3):533-545.
[5]Harman DM,Harman AL.Distribution pattern of admit locust borers,(Coleoptera:Cerambycidae)on nearby goldenrod,Solidago spp.(Asteraceae) at a forest field edge.Proc Entomol Sac Am,1987,(89):706-710
[6]Adachi I.Population studies of Anoplophora malasiaca adults(Coleopera:Cerambycidae)in a citrus grove.Res Popul Ecol,1990,(32):15-32.
[7]Iwabuchi K,Iakahashi J,Nakagava Y,et.al.Behavioral responses of female grape borer Xylotrechus pyrrhoderus(Coleopera:Cerambycidae) to synthetic male sex pheromone components.Appl Entomol Zool,1986,(21):21-27.
[8]Bento JMS,Albino FE,Delta Lucia TMC.Field trapping of Migdolus fryanus(Coleopera:Cerambycidae) using natural sex pheromone.Chem Ecol,1992:(18)245-851.
[9]Leal WS,Bento JMS,Vilela EF.Female sex pheromone of the longhorn beetle Migdolus fryanus Westwood:N(-2s)-methylbutanoyl-2-methylbutyl-amine.Expeientia,1994,50:853-856.
[10]Fettkother R,Dettner K,Sehroder F.The male sex pheromone of the old house borer Hylotrupes bajulus(Coleopera:Cerambycidae):identification and female response.Experiemiat,1995,(51):270-277.
[11]Kuwahara Y.Mathuyama S.Suzuki T.Identification of 2,3- octanediol,2- hydroxy -3-octanone and 3-hydroxy -2-octanone from male Xylotrechus chmensis as possible sex pheromones(Coleopera:Cerambycidae).Appl Eatomol Zool,1987,(22):25-28.
[12]Itami JK.Craig TP.Life history of Styloxus bicolor(Coleopera:Cerambycidae) on Juntperus monospeem in Northern Arizona.Ann Entomol Soc Am,1989,(82):582-587.
[13]Hanks LM.Influence of the larval host plant on reproductive strategies of cerambycid beetles.Ann Rev Entomol,1999,44:483-505.
[14]Wang Q,Zeng W,Chen LY.Circadian reproductive rhythms,pair-bonding,and evidence for sex-specific pheromones in Nadezhdiella cantori(Coleoptera:Cerambycidae).Journal of Insect Behavior,2002,15(4):527-539.
[15]Wang Q.Evidence for a contact female sex pheromone on Anoplophora chiuensis (Forster)(Coleoptera:Cerambycidae:Lamiinae).ColeoptBull.,1998,52:363-368.
[16]Ginzel MD,Hanks LM.Contact pheromones as mate recognition cues of four species of longhorned beetles(Coleoptera:Cerambycidae).Insect Behav.,2003,16(2):181-187.
[17]胡江,陈树琼,李义龙.雌性信息素在麻点豹天牛成虫交尾行为中的作用.昆虫知识,2006,43(2):251-254.
[18]Akutsu K,Kuboki M.Analysis of mating behavior of the udo longicorn beetle.Acalolepta luxuriosa Bates(Coleoptera:Cerambycidae).Jpn.J.Appl.Entomol.Zool.,1983,27:247-251.
[19]温硕洋.粗鞘双条杉天牛交配行为生物学及雌性识别信息素研究.植物保护学报,1991.18(2):167-172.
[20]Kim GH,Takabayashi J,Takahashi S.Function of contact pheromone in the mating behaviour of the cryptomeria bark borer,Semanotus japonicus Lacordaire(Coleoptera:Cerambycidae).Appl.Entomol.Zool.,1993,28:525-535.
[21]Fukaya M.Review recent advances in sex pheromone studies on the white-spotted longicorn beetle,Anoplophora malasiaca.Available online at http://www.jircas affrc.go.jp,2003
[22]Zhang AJ,James E,Oliver K,Zhao B,Xia L,Xu Z.Evidence for contact sex re cognition pheromone of the Asian longhorned beetle,Anoplophora g labripennis(Coleoptera:Cerambycidae).Naturwissenschaften,2007,90:410-413.
[23]Fukaya M,Yasuda T,Wakamura S.Reproductive biology of the yellow-spotted longicorn beetle,Psacothea hilaris(Pascoe)(Coleoptera:Cerambycidae).Identification of contact sex pheromone on female body surface.Chem.Ecol.,1996,22:259-270.
[24]Hanks LM.Influence of the larval host plant on reproductive strategies of cerambycid beetles.Ann.Rev Entomol,1999,44:483-505.
[25]张玉风,陆群,张宏世.光肩星天牛成虫特性及其行为的研究.内蒙古林业科技,1997,(4):7-9.
[26]高瑞桐.利用成虫取食习性防治3种杨树天牛.北京林业大学学报,1998,20(1):43-48.
[27]贺萍,黄竞芳.光肩星天牛成虫的行为.昆虫学报,1993,36(1):51-55.
[28]尹新明.狭胸天牛生殖行为的研究.河南农业大学学报,1996,30(4):347-349.
[29]李德家,所雅彦,中岛忠一.光肩星天牛成虫交配行为机制研究.北京林业大学学报,1999,21(4):33-36.
[30]Barata EN,Mustaparta H,Picket JA.Encoding of host and non-host plant odours by receptor neurons in the eucalyptus woodborer,Phoracantha s emipunctata(Coleoptera:Cerambycidae).Comp.Physiol A,2002,188:121-133.
[31]Fettkother R,Reddy GVP,Noldt U,Detmer K.Efect of host and larval frass volatiles on behavioral response of the old house borer,Hylotrupes bajulus(Coleoptera:Cerambycidae),in a wind tunnel bioassay.Chemoecology,2000,10:1-10.
[32]Higgs MD,Evans DA.Chemical mediators in the oviposition behaviour of the house longhorn beetle Hylotrupes bajulus.Experientia.,1978,34:46-47.
[33]Kobayashi F,Yamane A,Ikeda T.The Japanese pine sawyer beetle as the vector of pine wilt disease.Ann.Rew.Entomol.,1984,29:115-135.
[34]Sakai M,Yamasaki T.(+)-Juniperol and(+)-pimaral:attractants for the cerambycid beetle,Monochamus alternatus Hope.Chem.Ecol.,1990,16:3387-3392.
[35]Sakai M,Yamasaki T.(+)-cis-3-Pinen-2-ol:attractant for male cerambycid beetle,Monochamus alternatus Hope.J.Chem.Ecol,1991,17:757-765.
[36]Phillips TW,Wilkening A J,Atkinson TH.Synergism of turpentine and ethanol as attractants for certain pine-infesting beetles(Coleoptera).Enviorn.Entomol.,1988,17(3):456-462.
[37]Barata EN,Picket JA,Wadhams LJ.Identification of host semiochemicals of eucalyptus woodborer Phoracantha semipunctata by Gas Chromatography Electroantennography.Chem.Ecol.,2000,26(8):1877-1895.
[38]Suckling DM,Gibb AR,Daly JM.Behavioral and E lectorphysiological Responses of Arhopalustr istis to Burnt Pine and other Stimuli.Chem.Ecol.,2001,27(6):1091-1104.
[39]张洪波,赵云翔,蒋青年.咖啡灭字脊虎天牛捕捉器试验及其生物防治意义.云南热作科技,1999,22(1):12-13.
[40]Kobayashi F,Yamane A,Ikeda T.The Japanese pine sawyer beetle as the vector of pine wilt disease.Ann.Rev.Entomol,1984,29:115-135.
[41]Allison JD,Borden JH,Mcintosh RL.Kaiormonal response by four monochamus species(Coleoptera:Cerambycidae)to bark beetle pheromones.Chem.Ecol.,2001,27(4):633-646.
[42]Ikeda T,Ohya E,Makihara H.Olfactory responses of Anaglyptus subfasciatus PIC and Demonax transilis BATES(Coleoptera:Cerambycidae)to flower scents.Jpn.For.Soc.,1993,75(2):108-112.
[43]黄金水,何学友,叶剑雄.苦楝引诱防治星天牛研究.林业科学,2001,37(4):58-65.
[44]Landolt PJ,Heath RR.Sexual role reversal in mate-finding strategies of the cabbage looper moth.Science,1990,249:1026-1028.
[45]Nakamuta K,Sato H,Nakashima T.Behavioral and morphological evidence for a m ale-produced sex pheromone in the Cryptomeria twig borer,Anaglyptus subfasciatus Pic(C oleoptera:Cerambycidae).Jpn.J.Entomol.,1994,62:371-376.
[46]Linsley EG.Ecology of cerambycidae.Ann.Rev.Entomol.,1959,4:99-138.
[47]Iwabuchi K,Takahashi J,Sakai T.Occurrence of 2,3-octanediol and 2-hydroxy-3-octanone,possible male sex pheromone in Xylotrechus chinensis Chevrolat(C oleoptera:Cerambycidae).Appl.Ent.Zool.,1987,22:110-111.
[48]程惊秋,汤卫.构筒天牛交配行为生物学和性信息素研究.西南农业大学学报,1990,12(2):110-113.
[49]Ginzel MD,Hanks LM.Contact pheromones as mate recognition cues of four species of longhorned beetles(Coleoptera:Cerambycidae).Insect.Behav.,2003c,16(2):181-187.
[50]Wang Q,Chen L,Zeng W.Sex recognition by males and evidence for a female sex pheromone in paraglenea fortunei(Coleoptera:cerambycidae).Ann.Entomol.Soc.Am.,1991,84:107-110.
[51]Sakai T,Nakagawa Y,Takahashi J,Iwabuchi K,Ishii K.Isolation and identification of the male sex pheromone of the grape borer Xylotrechus pyrrhoderus Bates (Coleoptera:Cerambycidae).Chemistry Letters,1984,261:263-264.
[52]Doppelreiter H.Evidence for a female sex pheromone in the house longhorn beetle Hylotrupes bajulus.Z.Angew.Entomol,1979,88(1):56-59.
[53]Heintz A.Lepturinenras blombesok och sekundara konskaraktilrer.Entomol.Tidskr.,1925, 46:21-34.
[54]Fauziah BA,Hidaka T,Tabata K.There productive behavior of Monochamus alternatus Hope(Coleoptera:Cerambycidae).Appl.Entomol.Zool.,1987,22:272-285.
[55]Kim GH,Takabayashi J,Takahashi S,Tabata K.Function of pheromones in mating behavior of the Japanese pine sawyer beetle,Monochamus alternatus Hope.Appl.Entomol.Zool.,1992,27:489-497.
[56]周琳,马志卿,冯岗.天牛性信息素、引诱植物和植物性引诱剂的研究与应用.昆虫知识,2006,43(4):433-438.
[57]陆群,张玉凤,章海波.对光肩星天牛性信息素的探讨.内蒙古林业科技,1994,22(4):35-37.
[58]陆群,张玉风,王文祥.光肩星天牛雌虫产卵器及腺体分泌物的活性研究.内蒙古林业科技,2000(2):41-42.
[59]张岩,刘敬泽.昆虫的性信息素及其应用.生物学通报,2003,38(12):7-10.
[60]Raina Ak,Jaffe H,Kempe TG.Identification of a neuropeptide hormone that regulates sex pheromone production in female moths.Science,1989,244:796-797.
[61]黄勇平,沈君浑,王淑芬.昆虫性信息素变异研究的进展.中南林学院学报,1998,18(4):88-95.
[62]鲁玉杰,张孝羲,翟保平.固相微量萃取法在棉铃虫性信息素研究中的应用.南京农业大学学报,2001,24(2):49-52.
[63]Allison JD,Morewood WD,Borden JH.Diferentil bio-Activity of Ips and Dendorctonus(Coleoptera:Scolytidae)pheromone components for Monochamus clamator and M.scutellatus(Coleoptera:Ce rambycidae).Chem.Ecol.,2003,32(1):23-30.
[64]Morewood WD,Simmonds KE,Giles R.Disruption by c onophthorin of the kairomonal response of sawyer beetles to bark beetle pheromones.Chem.Ecol.,2003,29(9):2115-2129.
[65]Huber DPW,Giles R,Borden JH.Two pheromones of conifeorphagous bark beetles found in the bark of nonhost angiosperms.Chem.Ecol.,1999,25:805-816.
[66]Yamasaki T,Sato M,Sakoguchi H.(-)-Germacrene D:Masking substance of attractants for the cerambycid beetle,Monochmnus alternatus(Hope).Appl.EntotnoI.Zool.,1997,32(3):423-429.
[67]Anbutsu H,Togashi K.Oviposition deterrent by female reproductive gland secretion in Japanese pine Sawyer,Monochamus alternatus.Chem.Ecol.,2001,27(6):1151-1161.
[68]Anbutsu H,Togashi K.Oviposition behavior and response to the oviposition scars occupied by eggs in Monochamus saltuarius(Coleoptera:Cerambycidae).Appl.Entomol.Zool.,1997,32(4):541-549.
[69]Lacey ES,Ginzel MD,Millar JG,Hanks LM.Male-produced aggregation pheromone of the cerambycid beetle Neoclytus acuminatus acuminatus.J.Chem.Ecol.,2004,30:1493-1506.
[70]Thrum U,Kuppers J.Epithelial physiology of insect sensilla.In:Locke M,Smith DS,eds.Insect biology in the future.New York:Acad.Press.1980,355-385.
[71]雷宏,邱宇彤,T.A.Christensen.昆虫嗅觉系统的结构与功能.见:刘同先,康乐,编.昆虫学研究进展与展望,北京,科学出版社,2005,133-169.
[72]Altner H.Insect sensillum specificity and structure:an approach to a new typology.Olfaction Taste,1977,6:295-303.
[73]Altner H,Prillinger L.Ultrastructure of invertebrate chemo-,thermo-,and hygroreceptors and its functional significance.Int.Rev.Cytol.,1980,67:69-139.
[74]Keil T.Fine structure of the pheromone -sensitive sensilla on the antenna of the hawkmoth,Manduca sexta.Tissue Cell,1989,21:139-151.
[75]Keil T.Morphology and development of the peripheral olfactory organs.In:Hansson BS,eds.Insect olfaction.Germany:Springer-Verlag Berlin Heidelberg.1999,5-47.
[76]Keil T,Steinbrecht RA.Diffusion barriers in silkmoth sensory epithelia:application of lanthanum tracer to olfactory sensilla of Antheraea polyphemus and Bombyx mori.Tissue Cell,1987,19:119-134.
[77]Keil TA,Steinbrecht RA.Mechanosensitive and olfactory sensilla of insects.In:King RC,Akai H,eds.Insect ultrastructure.New York:Plenum,1994,1:402-433.
[78]Esslen J,Kaissling KE.Number and distribution of the sensilla on the antennal flagellum of the honeybee.Zoomorphologie,1976,83:227-251.
[79]Mclver SB.Review article:Sensilla of mosquitoes(Diptera:Culicidea).Med.Entomol.,1982,19:489-535.
[80]Steibrecht RA.The thermo-/hygrosensitive sensilla of the silkmoth,Bombyx mori:morphological changes after dry- and moisty-adaptation.Cell Tissue Res,1991,266:411-456.
[81]Ochieng SA,Hallberg E,Hansson BS.Fine structure and distribution of antennal sensilla of the desert locust,Schistocerca gregaria(Orthoptera:Acrididae).Cell Tissue Res.,1998,291:525-536.
[82]张瑛,魏传秀.寄生蜂对外界信号的感受及其与行为反应的关系.昆虫知识,1998,35(3):176-178.
[83]程家安,唐振华.昆虫与植物相互关系的分子基础.北京,科学出版社,2000,146-149.
[84]Vogt RG,Riddiford LM.Pheromone binding and inactivation by moth antennae.Nature,1981,293:161-163.
[85]Ishida Y,Ching VP,Haverty MI.Odorant-binding proteins from a primitive termite.Chem.Ecol.,2002,28(9):1887-1893.
[86]Park SK,Shanbhag SR,Wang Q.Expression patterns of two putative odorant-binding proteins in the olfactory organs of Drosophila melanogaster have different implications for their funcitons.Cell Tissue Res.,2000,300:181-192.
[87]Richard GV.Odorant binding protein homologues of the malaria mosquito Anopheles gambiae;Possible orthologues of the Os-E and Os-F obps Dorsophila melanogaster.Chem.Ecol.,2002,28(11):2371-2376.
[88]丁红建.用于昆虫嗅觉行为研究的四臂嗅觉仪的设计、制作和应用.昆虫知识.1996.33(4):241-243.
[2]Nakamuta K,Leal WS,Nakashima T.Increase of trap catches by a combination of male sex pheromones and floral attractant in longhonr beetle,Anaglyptus subfasciatus.Chem Ecol,1997,23:1635-1640.
[3]嵇保中,魏勇,黄振裕.天牛成虫研究的现状与展望.南京林业大学学报(自然科学版),2002,26(2):79-83.
[4]Ginzel MD,Blomquist GJ,Millar JG,LM Hanks.Role of contact pheromones in mate recognition in Xylotrechus colonus.Chem Ecol,2003a,129(3):533-545.
[5]Harman DM,Harman AL.Distribution pattern of admit locust borers,(Coleoptera:Cerambycidae)on nearby goldenrod,Solidago spp.(Asteraceae) at a forest field edge.Proc Entomol Sac Am,1987,(89):706-710
[6]Adachi I.Population studies of Anoplophora malasiaca adults(Coleopera:Cerambycidae)in a citrus grove.Res Popul Ecol,1990,(32):15-32.
[7]Iwabuchi K,Iakahashi J,Nakagava Y,et.al.Behavioral responses of female grape borer Xylotrechus pyrrhoderus(Coleopera:Cerambycidae) to synthetic male sex pheromone components.Appl Entomol Zool,1986,(21):21-27.
[8]Bento JMS,Albino FE,Delta Lucia TMC.Field trapping of Migdolus fryanus(Coleopera:Cerambycidae) using natural sex pheromone.Chem Ecol,1992:(18)245-851.
[9]Leal WS,Bento JMS,Vilela EF.Female sex pheromone of the longhorn beetle Migdolus fryanus Westwood:N(-2s)-methylbutanoyl-2-methylbutyl-amine.Expeientia,1994,50:853-856.
[10]Fettkother R,Dettner K,Sehroder F.The male sex pheromone of the old house borer Hylotrupes bajulus(Coleopera:Cerambycidae):identification and female response.Experiemiat,1995,(51):270-277.
[11]Kuwahara Y.Mathuyama S.Suzuki T.Identification of 2,3- octanediol,2- hydroxy -3-octanone and 3-hydroxy -2-octanone from male Xylotrechus chmensis as possible sex pheromones(Coleopera:Cerambycidae).Appl Eatomol Zool,1987,(22):25-28.
[12]Itami JK.Craig TP.Life history of Styloxus bicolor(Coleopera:Cerambycidae) on Juntperus monospeem in Northern Arizona.Ann Entomol Soc Am,1989,(82):582-587.
[13]Hanks LM.Influence of the larval host plant on reproductive strategies of cerambycid beetles.Ann Rev Entomol,1999,44:483-505.
[14]Wang Q,Zeng W,Chen LY.Circadian reproductive rhythms,pair-bonding,and evidence for sex-specific pheromones in Nadezhdiella cantori(Coleoptera:Cerambycidae).Journal of Insect Behavior,2002,15(4):527-539.
[15]Wang Q.Evidence for a contact female sex pheromone on Anoplophora chiuensis (Forster)(Coleoptera:Cerambycidae:Lamiinae).ColeoptBull.,1998,52:363-368.
[16]Ginzel MD,Hanks LM.Contact pheromones as mate recognition cues of four species of longhorned beetles(Coleoptera:Cerambycidae).Insect Behav.,2003,16(2):181-187.
[17]胡江,陈树琼,李义龙.雌性信息素在麻点豹天牛成虫交尾行为中的作用.昆虫知识,2006,43(2):251-254.
[18]Akutsu K,Kuboki M.Analysis of mating behavior of the udo longicorn beetle.Acalolepta luxuriosa Bates(Coleoptera:Cerambycidae).Jpn.J.Appl.Entomol.Zool.,1983,27:247-251.
[19]温硕洋.粗鞘双条杉天牛交配行为生物学及雌性识别信息素研究.植物保护学报,1991.18(2):167-172.
[20]Kim GH,Takabayashi J,Takahashi S.Function of contact pheromone in the mating behaviour of the cryptomeria bark borer,Semanotus japonicus Lacordaire(Coleoptera:Cerambycidae).Appl.Entomol.Zool.,1993,28:525-535.
[21]Fukaya M.Review recent advances in sex pheromone studies on the white-spotted longicorn beetle,Anoplophora malasiaca.Available online at http://www.jircas affrc.go.jp,2003
[22]Zhang AJ,James E,Oliver K,Zhao B,Xia L,Xu Z.Evidence for contact sex re cognition pheromone of the Asian longhorned beetle,Anoplophora g labripennis(Coleoptera:Cerambycidae).Naturwissenschaften,2007,90:410-413.
[23]Fukaya M,Yasuda T,Wakamura S.Reproductive biology of the yellow-spotted longicorn beetle,Psacothea hilaris(Pascoe)(Coleoptera:Cerambycidae).Identification of contact sex pheromone on female body surface.Chem.Ecol.,1996,22:259-270.
[24]Hanks LM.Influence of the larval host plant on reproductive strategies of cerambycid beetles.Ann.Rev Entomol,1999,44:483-505.
[25]张玉风,陆群,张宏世.光肩星天牛成虫特性及其行为的研究.内蒙古林业科技,1997,(4):7-9.
[26]高瑞桐.利用成虫取食习性防治3种杨树天牛.北京林业大学学报,1998,20(1):43-48.
[27]贺萍,黄竞芳.光肩星天牛成虫的行为.昆虫学报,1993,36(1):51-55.
[28]尹新明.狭胸天牛生殖行为的研究.河南农业大学学报,1996,30(4):347-349.
[29]李德家,所雅彦,中岛忠一.光肩星天牛成虫交配行为机制研究.北京林业大学学报,1999,21(4):33-36.
[30]Barata EN,Mustaparta H,Picket JA.Encoding of host and non-host plant odours by receptor neurons in the eucalyptus woodborer,Phoracantha s emipunctata(Coleoptera:Cerambycidae).Comp.Physiol A,2002,188:121-133.
[31]Fettkother R,Reddy GVP,Noldt U,Detmer K.Efect of host and larval frass volatiles on behavioral response of the old house borer,Hylotrupes bajulus(Coleoptera:Cerambycidae),in a wind tunnel bioassay.Chemoecology,2000,10:1-10.
[32]Higgs MD,Evans DA.Chemical mediators in the oviposition behaviour of the house longhorn beetle Hylotrupes bajulus.Experientia.,1978,34:46-47.
[33]Kobayashi F,Yamane A,Ikeda T.The Japanese pine sawyer beetle as the vector of pine wilt disease.Ann.Rew.Entomol.,1984,29:115-135.
[34]Sakai M,Yamasaki T.(+)-Juniperol and(+)-pimaral:attractants for the cerambycid beetle,Monochamus alternatus Hope.Chem.Ecol.,1990,16:3387-3392.
[35]Sakai M,Yamasaki T.(+)-cis-3-Pinen-2-ol:attractant for male cerambycid beetle,Monochamus alternatus Hope.J.Chem.Ecol,1991,17:757-765.
[36]Phillips TW,Wilkening A J,Atkinson TH.Synergism of turpentine and ethanol as attractants for certain pine-infesting beetles(Coleoptera).Enviorn.Entomol.,1988,17(3):456-462.
[37]Barata EN,Picket JA,Wadhams LJ.Identification of host semiochemicals of eucalyptus woodborer Phoracantha semipunctata by Gas Chromatography Electroantennography.Chem.Ecol.,2000,26(8):1877-1895.
[38]Suckling DM,Gibb AR,Daly JM.Behavioral and E lectorphysiological Responses of Arhopalustr istis to Burnt Pine and other Stimuli.Chem.Ecol.,2001,27(6):1091-1104.
[39]张洪波,赵云翔,蒋青年.咖啡灭字脊虎天牛捕捉器试验及其生物防治意义.云南热作科技,1999,22(1):12-13.
[40]Kobayashi F,Yamane A,Ikeda T.The Japanese pine sawyer beetle as the vector of pine wilt disease.Ann.Rev.Entomol,1984,29:115-135.
[41]Allison JD,Borden JH,Mcintosh RL.Kaiormonal response by four monochamus species(Coleoptera:Cerambycidae)to bark beetle pheromones.Chem.Ecol.,2001,27(4):633-646.
[42]Ikeda T,Ohya E,Makihara H.Olfactory responses of Anaglyptus subfasciatus PIC and Demonax transilis BATES(Coleoptera:Cerambycidae)to flower scents.Jpn.For.Soc.,1993,75(2):108-112.
[43]黄金水,何学友,叶剑雄.苦楝引诱防治星天牛研究.林业科学,2001,37(4):58-65.
[44]Landolt PJ,Heath RR.Sexual role reversal in mate-finding strategies of the cabbage looper moth.Science,1990,249:1026-1028.
[45]Nakamuta K,Sato H,Nakashima T.Behavioral and morphological evidence for a m ale-produced sex pheromone in the Cryptomeria twig borer,Anaglyptus subfasciatus Pic(C oleoptera:Cerambycidae).Jpn.J.Entomol.,1994,62:371-376.
[46]Linsley EG.Ecology of cerambycidae.Ann.Rev.Entomol.,1959,4:99-138.
[47]Iwabuchi K,Takahashi J,Sakai T.Occurrence of 2,3-octanediol and 2-hydroxy-3-octanone,possible male sex pheromone in Xylotrechus chinensis Chevrolat(C oleoptera:Cerambycidae).Appl.Ent.Zool.,1987,22:110-111.
[48]程惊秋,汤卫.构筒天牛交配行为生物学和性信息素研究.西南农业大学学报,1990,12(2):110-113.
[49]Ginzel MD,Hanks LM.Contact pheromones as mate recognition cues of four species of longhorned beetles(Coleoptera:Cerambycidae).Insect.Behav.,2003c,16(2):181-187.
[50]Wang Q,Chen L,Zeng W.Sex recognition by males and evidence for a female sex pheromone in paraglenea fortunei(Coleoptera:cerambycidae).Ann.Entomol.Soc.Am.,1991,84:107-110.
[51]Sakai T,Nakagawa Y,Takahashi J,Iwabuchi K,Ishii K.Isolation and identification of the male sex pheromone of the grape borer Xylotrechus pyrrhoderus Bates (Coleoptera:Cerambycidae).Chemistry Letters,1984,261:263-264.
[52]Doppelreiter H.Evidence for a female sex pheromone in the house longhorn beetle Hylotrupes bajulus.Z.Angew.Entomol,1979,88(1):56-59.
[53]Heintz A.Lepturinenras blombesok och sekundara konskaraktilrer.Entomol.Tidskr.,1925, 46:21-34.
[54]Fauziah BA,Hidaka T,Tabata K.There productive behavior of Monochamus alternatus Hope(Coleoptera:Cerambycidae).Appl.Entomol.Zool.,1987,22:272-285.
[55]Kim GH,Takabayashi J,Takahashi S,Tabata K.Function of pheromones in mating behavior of the Japanese pine sawyer beetle,Monochamus alternatus Hope.Appl.Entomol.Zool.,1992,27:489-497.
[56]周琳,马志卿,冯岗.天牛性信息素、引诱植物和植物性引诱剂的研究与应用.昆虫知识,2006,43(4):433-438.
[57]陆群,张玉凤,章海波.对光肩星天牛性信息素的探讨.内蒙古林业科技,1994,22(4):35-37.
[58]陆群,张玉风,王文祥.光肩星天牛雌虫产卵器及腺体分泌物的活性研究.内蒙古林业科技,2000(2):41-42.
[59]张岩,刘敬泽.昆虫的性信息素及其应用.生物学通报,2003,38(12):7-10.
[60]Raina Ak,Jaffe H,Kempe TG.Identification of a neuropeptide hormone that regulates sex pheromone production in female moths.Science,1989,244:796-797.
[61]黄勇平,沈君浑,王淑芬.昆虫性信息素变异研究的进展.中南林学院学报,1998,18(4):88-95.
[62]鲁玉杰,张孝羲,翟保平.固相微量萃取法在棉铃虫性信息素研究中的应用.南京农业大学学报,2001,24(2):49-52.
[63]Allison JD,Morewood WD,Borden JH.Diferentil bio-Activity of Ips and Dendorctonus(Coleoptera:Scolytidae)pheromone components for Monochamus clamator and M.scutellatus(Coleoptera:Ce rambycidae).Chem.Ecol.,2003,32(1):23-30.
[64]Morewood WD,Simmonds KE,Giles R.Disruption by c onophthorin of the kairomonal response of sawyer beetles to bark beetle pheromones.Chem.Ecol.,2003,29(9):2115-2129.
[65]Huber DPW,Giles R,Borden JH.Two pheromones of conifeorphagous bark beetles found in the bark of nonhost angiosperms.Chem.Ecol.,1999,25:805-816.
[66]Yamasaki T,Sato M,Sakoguchi H.(-)-Germacrene D:Masking substance of attractants for the cerambycid beetle,Monochmnus alternatus(Hope).Appl.EntotnoI.Zool.,1997,32(3):423-429.
[67]Anbutsu H,Togashi K.Oviposition deterrent by female reproductive gland secretion in Japanese pine Sawyer,Monochamus alternatus.Chem.Ecol.,2001,27(6):1151-1161.
[68]Anbutsu H,Togashi K.Oviposition behavior and response to the oviposition scars occupied by eggs in Monochamus saltuarius(Coleoptera:Cerambycidae).Appl.Entomol.Zool.,1997,32(4):541-549.
[69]Lacey ES,Ginzel MD,Millar JG,Hanks LM.Male-produced aggregation pheromone of the cerambycid beetle Neoclytus acuminatus acuminatus.J.Chem.Ecol.,2004,30:1493-1506.
[70]Thrum U,Kuppers J.Epithelial physiology of insect sensilla.In:Locke M,Smith DS,eds.Insect biology in the future.New York:Acad.Press.1980,355-385.
[71]雷宏,邱宇彤,T.A.Christensen.昆虫嗅觉系统的结构与功能.见:刘同先,康乐,编.昆虫学研究进展与展望,北京,科学出版社,2005,133-169.
[72]Altner H.Insect sensillum specificity and structure:an approach to a new typology.Olfaction Taste,1977,6:295-303.
[73]Altner H,Prillinger L.Ultrastructure of invertebrate chemo-,thermo-,and hygroreceptors and its functional significance.Int.Rev.Cytol.,1980,67:69-139.
[74]Keil T.Fine structure of the pheromone -sensitive sensilla on the antenna of the hawkmoth,Manduca sexta.Tissue Cell,1989,21:139-151.
[75]Keil T.Morphology and development of the peripheral olfactory organs.In:Hansson BS,eds.Insect olfaction.Germany:Springer-Verlag Berlin Heidelberg.1999,5-47.
[76]Keil T,Steinbrecht RA.Diffusion barriers in silkmoth sensory epithelia:application of lanthanum tracer to olfactory sensilla of Antheraea polyphemus and Bombyx mori.Tissue Cell,1987,19:119-134.
[77]Keil TA,Steinbrecht RA.Mechanosensitive and olfactory sensilla of insects.In:King RC,Akai H,eds.Insect ultrastructure.New York:Plenum,1994,1:402-433.
[78]Esslen J,Kaissling KE.Number and distribution of the sensilla on the antennal flagellum of the honeybee.Zoomorphologie,1976,83:227-251.
[79]Mclver SB.Review article:Sensilla of mosquitoes(Diptera:Culicidea).Med.Entomol.,1982,19:489-535.
[80]Steibrecht RA.The thermo-/hygrosensitive sensilla of the silkmoth,Bombyx mori:morphological changes after dry- and moisty-adaptation.Cell Tissue Res,1991,266:411-456.
[81]Ochieng SA,Hallberg E,Hansson BS.Fine structure and distribution of antennal sensilla of the desert locust,Schistocerca gregaria(Orthoptera:Acrididae).Cell Tissue Res.,1998,291:525-536.
[82]张瑛,魏传秀.寄生蜂对外界信号的感受及其与行为反应的关系.昆虫知识,1998,35(3):176-178.
[83]程家安,唐振华.昆虫与植物相互关系的分子基础.北京,科学出版社,2000,146-149.
[84]Vogt RG,Riddiford LM.Pheromone binding and inactivation by moth antennae.Nature,1981,293:161-163.
[85]Ishida Y,Ching VP,Haverty MI.Odorant-binding proteins from a primitive termite.Chem.Ecol.,2002,28(9):1887-1893.
[86]Park SK,Shanbhag SR,Wang Q.Expression patterns of two putative odorant-binding proteins in the olfactory organs of Drosophila melanogaster have different implications for their funcitons.Cell Tissue Res.,2000,300:181-192.
[87]Richard GV.Odorant binding protein homologues of the malaria mosquito Anopheles gambiae;Possible orthologues of the Os-E and Os-F obps Dorsophila melanogaster.Chem.Ecol.,2002,28(11):2371-2376.
[88]丁红建.用于昆虫嗅觉行为研究的四臂嗅觉仪的设计、制作和应用.昆虫知识.1996.33(4):241-243.