调控褐飞虱及其天敌稻虱缨小蜂行为的活性化合物筛选及其田间效果的初步测定
|
摘要
植物在遭受植食性昆虫为害后能特异性地释放挥发性物质,这些挥发物不仅能对植食性昆虫的寄生性或捕食性天敌产生引诱作用,而且亦能对周围的植物、植食性昆虫以及微生物等产生影响。鉴定这些挥发物中的活性组分,可望开发治理害虫的新方法。为此,本文以我们前期报道的褐飞虱为害诱导的水稻挥发物为基础,筛选了能对褐飞虱及其天敌稻虱缨小蜂行为产生影响的活性组分,并对能引诱稻虱缨小蜂的活性化合物或混合物进行了初步的田间试验。主要结果如下
室内测定了褐飞虱对人工合成挥发物组分的行为反应。结果表明,共有6种挥发物组分对褐飞虱的行为反应具有明显的影响。其中farnesene、MeSA和2-heptanone对褐飞虱表现为趋避作用,linalool、E-3-ocimene和2-heptanol对褐飞虱表现为引诱作用。在具有趋避作用的组分中,farnesene和MeSA的作用较强；在具有引诱作用的活性组分中,linalool的作用最强。
利用Y型嗅觉仪测定稻虱缨小蜂对18种单一挥发物组分和32种混配物的行为反应。结果表明,有5种单一挥发物组分,包括3种绿叶性气味(E-3-hexenyl acetate,1-penten-3-ol, E-3-hexenal均为50ppm)、1种萜类化合物(linalool,50ppm)和水杨酸甲酯(MeSA,5ppm和50ppm),对稻虱缨小蜂具有明显的引诱作用。有趣的是,我们也发现了2种挥发物组分,(+)-limonene和2-heptanol,对该蜂具有一定的趋避作用。在32种混配物中,发现有3种对稻虱缨小蜂具有显著引诱作用。
在田间初步测定了对稻虱缨小蜂具有引诱作用的5种单一活性组分和1种混配物对田间褐飞虱卵被寄生率和被捕食率的影响。结果表明,3种单一活性组分,Linalool, E-3-hexenal和E-3-hexenyl acetate,和混配物在田间使用后能显著提高褐飞虱卵的被寄生率,分别比对照上的被寄生率上升31.67%、17.47%、7.76%和34.93%。另2种组分,MeSA和1-penten-3-ol虽然没有显著提高褐飞虱卵的被寄生率,但与对照相比仍有上升趋势。所有单一活性组分和混配物均没有显著提高褐飞虱卵的被捕食率,但除了E-3-hexenal外,其余化合物的使用都比对照有增加的趋势。
上述研究结果表明,虫害诱导的水稻挥发物中确实存在能对褐飞虱及其天敌稻虱缨小蜂行为反应产生影响的活性组分,对这些活性组分的进一步研究,可望开发褐飞虱治理的新方法与新途径。
In response to herbivory, plants will emit specific blends of volatiles, which may not only attract the natural enemies of the herbivores, but also influence the behavior and/or performance of herbivores, microbes and the nearby plants. It will be important for finding new methods for pest control to identify active chemicals that manipulate behavioral responses of the herbivores and their natural enemies in herbivore-induced plant volatiles (HIPVs). Based on our previous identified volatiles from rice plants infested by rice brown planthopper (BPH) Nilaparvata lugens (Stal), we screened the bioactive chemicals that modulate behavioral response of BPH and its egg parasitoid Anagrus nilaparvatae. Moreover, the efficiency of the active chemicals in enhancing effect of the parasitoid in the field was tested. The main results are as follows:
Responses of N. lugens to synthetic HIPVs were determined under laboratory conditions. The results showed that 6 chemicals have significant influence on the behavioral response of BPH. Among the 6 chemicals, farnesene, MeSA and 2-heptanone significantly repelled BPH, while linalool, E-3-ocimene and 2-heptanol significantly attracted BPH. Farnesene and MeSA were more active in repelling BPH than 2-heptanone, and linalool, on the other hand, was the most active among the 3 chemicals that were attractive to BPH.
Attractiveness of 18 single synthetic HIPVs and 32 blends to A. nilaparvatae were tested in a Y-tube olfactometer. Five single chemicals, including three green leafy volatiles (E-3-hexenyl acetate, 1-penten-3-ol, E-3-hexenal)(50ppm), one terpenoid (linalool,50ppm) and methyl salicylate (MeSA,5ppm and 50ppm), were found to be strongly attractive to A. nilaparvatae. On the other hand, the parasitoid was also attracted by 3 blends. Interestingly, we also found that the parasitoid was repelled by 2 single chemicals, (+)-limonene and 2-heptanol.
Primary field experiments were conducted to evaluate the effectiveness of these bioactive compounds or blends in attracting A. nilaparvatae and predators of BPH. The results showed that parasitism rates of BPH eggs by the parasitoid in the field were significantly increased when plants were hang up a seprum containing linalool, E-3-hexenal, E-3-hexenyl acetate or the mixture; the parasitism rates, compared with those in the control field, were increased by 31.67%,17.47%,7.76% and 34.93%, respectively. Application of other 2 single compounds, MeSA and 1-penten-3-ol, also increased the parasitism rates of BPH eggs, though this difference was not significant compared to the control. Application of all of the single chemicals or the blend did not cause significant increases in predation rates of BPH eggs.
The results indicate that some chemicals from HIPVs did have influences on behavioral responses of rice brown planthopper N. lugens and its egg parasitoid A. nilaparvatae. Further elucidation of these active chemicals will exploit new methods and strategies for pest management.
室内测定了褐飞虱对人工合成挥发物组分的行为反应。结果表明,共有6种挥发物组分对褐飞虱的行为反应具有明显的影响。其中farnesene、MeSA和2-heptanone对褐飞虱表现为趋避作用,linalool、E-3-ocimene和2-heptanol对褐飞虱表现为引诱作用。在具有趋避作用的组分中,farnesene和MeSA的作用较强；在具有引诱作用的活性组分中,linalool的作用最强。
利用Y型嗅觉仪测定稻虱缨小蜂对18种单一挥发物组分和32种混配物的行为反应。结果表明,有5种单一挥发物组分,包括3种绿叶性气味(E-3-hexenyl acetate,1-penten-3-ol, E-3-hexenal均为50ppm)、1种萜类化合物(linalool,50ppm)和水杨酸甲酯(MeSA,5ppm和50ppm),对稻虱缨小蜂具有明显的引诱作用。有趣的是,我们也发现了2种挥发物组分,(+)-limonene和2-heptanol,对该蜂具有一定的趋避作用。在32种混配物中,发现有3种对稻虱缨小蜂具有显著引诱作用。
在田间初步测定了对稻虱缨小蜂具有引诱作用的5种单一活性组分和1种混配物对田间褐飞虱卵被寄生率和被捕食率的影响。结果表明,3种单一活性组分,Linalool, E-3-hexenal和E-3-hexenyl acetate,和混配物在田间使用后能显著提高褐飞虱卵的被寄生率,分别比对照上的被寄生率上升31.67%、17.47%、7.76%和34.93%。另2种组分,MeSA和1-penten-3-ol虽然没有显著提高褐飞虱卵的被寄生率,但与对照相比仍有上升趋势。所有单一活性组分和混配物均没有显著提高褐飞虱卵的被捕食率,但除了E-3-hexenal外,其余化合物的使用都比对照有增加的趋势。
上述研究结果表明,虫害诱导的水稻挥发物中确实存在能对褐飞虱及其天敌稻虱缨小蜂行为反应产生影响的活性组分,对这些活性组分的进一步研究,可望开发褐飞虱治理的新方法与新途径。
In response to herbivory, plants will emit specific blends of volatiles, which may not only attract the natural enemies of the herbivores, but also influence the behavior and/or performance of herbivores, microbes and the nearby plants. It will be important for finding new methods for pest control to identify active chemicals that manipulate behavioral responses of the herbivores and their natural enemies in herbivore-induced plant volatiles (HIPVs). Based on our previous identified volatiles from rice plants infested by rice brown planthopper (BPH) Nilaparvata lugens (Stal), we screened the bioactive chemicals that modulate behavioral response of BPH and its egg parasitoid Anagrus nilaparvatae. Moreover, the efficiency of the active chemicals in enhancing effect of the parasitoid in the field was tested. The main results are as follows:
Responses of N. lugens to synthetic HIPVs were determined under laboratory conditions. The results showed that 6 chemicals have significant influence on the behavioral response of BPH. Among the 6 chemicals, farnesene, MeSA and 2-heptanone significantly repelled BPH, while linalool, E-3-ocimene and 2-heptanol significantly attracted BPH. Farnesene and MeSA were more active in repelling BPH than 2-heptanone, and linalool, on the other hand, was the most active among the 3 chemicals that were attractive to BPH.
Attractiveness of 18 single synthetic HIPVs and 32 blends to A. nilaparvatae were tested in a Y-tube olfactometer. Five single chemicals, including three green leafy volatiles (E-3-hexenyl acetate, 1-penten-3-ol, E-3-hexenal)(50ppm), one terpenoid (linalool,50ppm) and methyl salicylate (MeSA,5ppm and 50ppm), were found to be strongly attractive to A. nilaparvatae. On the other hand, the parasitoid was also attracted by 3 blends. Interestingly, we also found that the parasitoid was repelled by 2 single chemicals, (+)-limonene and 2-heptanol.
Primary field experiments were conducted to evaluate the effectiveness of these bioactive compounds or blends in attracting A. nilaparvatae and predators of BPH. The results showed that parasitism rates of BPH eggs by the parasitoid in the field were significantly increased when plants were hang up a seprum containing linalool, E-3-hexenal, E-3-hexenyl acetate or the mixture; the parasitism rates, compared with those in the control field, were increased by 31.67%,17.47%,7.76% and 34.93%, respectively. Application of other 2 single compounds, MeSA and 1-penten-3-ol, also increased the parasitism rates of BPH eggs, though this difference was not significant compared to the control. Application of all of the single chemicals or the blend did not cause significant increases in predation rates of BPH eggs.
The results indicate that some chemicals from HIPVs did have influences on behavioral responses of rice brown planthopper N. lugens and its egg parasitoid A. nilaparvatae. Further elucidation of these active chemicals will exploit new methods and strategies for pest management.
引文
1.邓晓军,陈晓亚,杜佳伟。植物挥发性物质及其代谢工程。植物生理与分子生物学报,2004,30(1)：11-18.
2.杜丽娜,张存莉,朱玮,张高宏。植物次生代谢合成途径及生物学意义。西北林学院学报,2005,20(3)：150-155.
3.李新岗,刘惠霞,黄建。虫害诱导植物防御的分子机理研究进展。应用生态学报,2008,19(4)：893-900.
4.娄永根,程家安。虫害诱导的植物挥发物：基本特性、生态学功能及释放机制。生态学报,2000,20(6)：1097-1106.
5.娄永根,程家安。植物的诱导抗虫性。昆虫学报,1997,40(3)：320-331
6.祝传书,赵惠燕。虫害诱导的植物挥发物。西北农林科技大学学报,2003,31：183-186.
7. Alberto Arab, Jose Roberto Trigo, Andre Luiz Lourencao, Aiarie Michele Peixoto, Fernanda Ramos, Jose Mauricio Simoes Bento. Differential attractiveness of potato tuber volatiles to Phthorimaea operculella (Gelechiidae) and the predator Orius insidiosus (Anthocoridae). J Chem Ecol,2007,33:1845-1855.
8. Antonin Ferry, Sebastien Dugravot, Thomas Delattre, Jean-Philippe Christides, Jacques Auger, Anne-Genevieve Bagneres, Denis Poinsot, Anne-Marie Cortesero. Identification of a widespread monomolecular odor differentially attractive to several Delia radicum ground-dwelling predators in the field. J Chem Ecol,2007, 33:2064-2077.
9. Bernasconi ML, Turlings TCJ, Ambrosetti L, Bassetti P, Dorn S. Herbivore-induced emissions of maize volatiles repel the corn leaf aphid, Rhopalosiphum maidis. Entomol Exp Appl,1998,87:133-142.
10. Carroll MJ, Schmelz EA, MeagherRL, Teal PEA. Attraction of Spodoptera frugiperda Larvae to Volatiles from Herbivore-Damaged Maize Seedlings. J Chem Ecol,2006,32:1911-1924.
11. Cesar R. Rodriguez-Saona, Luis E. Rodriguez-Saona, Christopher J. Frost. Herbivore-induced volatiles in the perennial shrub, Vaccinium corymbosum, and their role in inter-branch signaling. J Chem Ecol,2009,35:163-175.
12. Christopher J. Frost, Mark C. Mescher, John E. Carlson, Consuelo M. De Moraes. Plant defense priming against herbivores:Getting ready for a different battle. Plant physiology,2008a,146:818-824
13. Christopher J.Frost, Mark C. Mescher, John E. Carlson, Consuelo M. De Moraes. Why do distance limitations exist on plant-plant signaling via airborne volatiles? Plant Signaling and Behavior,2008b,3(7):466-468.
14. Colazza S, McElfresh JS, Millar JG. Identification of volatile synomones, induced by Nezara viridula feeding and oviposition on bean spp., that attract the egg parasitoid Trissolcus basalis. J Chem Ecol,2004,30:945-964.
15. David G. James. Further field evaluation of synthetic herbivore-induced plant volatiles as attractants for beneficial insects. Journal of Chemical Ecology,2005,3 (31):481-487.
16. David G. James. Field evaluation of herbivore-induced plant volatiles as attractants for beneficial insects:methyl salicylate and the green lacewing, Chrysopa nigricornis. Journal of Chemical Ecology,2003a,29,1601-1609.
17. David G. James. Synthetic herbivore-induced plant volatiles as attractants for beneficial insects. Environmental Entomology,2003b,32,977-982.
18. David G. James. Synthetic herbivore-induced plant volatiles as field attractants for beneficial insects. The British Crop Protection Council International Congress-Crop Science and Technology,2003c,1217-1222.
19. David G. James. Grape and hop plants sprayed with botanical oil pesticides containing herbivore-induced plant volatiles attract insect predators and parasitoids. Environmental Entomology (submitted for publication),2008.
20. David G James, Castle, S.C., Grasswitz, T., Reyna, V. Using synthetic herbivore-induced plant volatiles to enhance conservation biological control in hops and grapes. In:Proceedings of the 2nd International Symposium on Biological Control of Arthropods, pp.,2005,192-205.
21. David G. James, Grasswitz, T.R. Field attraction of parasitic wasps, Metaphycus sp. and Anagrus spp. using synthetic herbivore-induced plant volatiles. Biocontrol,2005, 50,871-880.
22. David G. James, Price, T.S. Field-testing of methyl salicylate for recruitment and retention of beneficial insects in grapes and hops. Journal of Chemical Ecology,2004, 30,1613-1628.
23. Devendra Kumar Choudhary, Bhavdish N. Johri, Anil Prakash. Volatiles as priming agents that initiate plant growth and defence responses. current science,2008, 94(5):595-604.
24. Dicke, M., van Poecke, R.P., and de Boer, J.G. Inducible indirect defence of plants: From mechanisms to ecological functions. Basic Appl. Ecol.,2003,4:27-42.
25. Douglas M. Light. A pear-derived kairomne with pheromonal potency that attracts male and female codling moth, cydia pomonella (L.). Naturwissenschaften,2001,88: 333-338.
26. Emmanuel Gaquerel, Alexander Weinhold, Ian T. Baldwin. Molecular interactions between the specialist herbivore Manduca sexta (Lepidoptera, Sphigidae) and its natural host Nicotiana attenuata. Ⅷ. An unbiased GCxGC-ToFMS analysis of the plant's. Plant Physiology,2009,149:1408-1423.
27. Eric Rohrig, John Sivinski, Peter Teal, Charles Stuhl, Martin Aluja. A floral-derived compound attractive to the tephritid fruit fly parasitoid Diachasmimorpha longicaudata (Hymenoptera:Braconidae). J Chem Ecol,2008,34:549-557.
28. Gen-ichiro Arimura, Kenji Matsui, Junji Takabayashi. Chemical and molecular ecology of herbivore-induced plant volatiles:Proximate factors and their ultimate functions. Plant Cell Physiology,2009,50(5):911-923.
29. Hamid R. S. Moayeri, Ahmad Ashouri, Henrik F. Brodsgaard, Annie Enkegaard. Males of the predatory mirid bug Macrolophus caliginosus exploit plant volatiles induced by conspecifics as a sexual synomone. Entomologia Experimentalis et Applicata,2007,123(1):49-53.
30. Hayato Ishiwari, Takahisa Suzuki, Taro Maeda. Essential compounds in herbivore-induced plant volatiles that attract the predatory mite Neoseiulus womersleyi. J Chem Ecol,2007,33:1670-1681.
31. Huilin Yu, Yongjun Zhang, Kongming Wu, Xi Wu Gao, Yu Yuan Guo. Field-testing of synthetic herbivore-induced plant volatiles as attractants for beneficial insects. Environmental Entomology,2008,37(6):1410-1415.
32. Ivo Feussner, Claus Wasternack. The lipoxygenase pathway. Ann. Rev. Plant Biol. 2002,53:275-97.
33. Jared G. Ali, Hans T. Alborn, Lukasz L. Stelinski. Subterranean herbivore-induced volatiles released by citrus roots upon feeding by Diaprepes abbreviates recruit entomopathogenic nematodes. J Chem Ecol,2010,36:361-368.
34. Jianing Wei, Lizhong Wang, Junwei Zhu, Sufang Zhang, Owi I. Nandi, Le Kang.. Plants attract parasitic wasps to defend themselves against insect pests by releasing hexenol. PLoS ONE,2007,9:1-7.
35. Joshua S. Yuan, Tobias G. Kollner, Greg Wiggins, Jerome Grant, Jorg Degenhardt, Feng Chen. Molecular and genomic basis of volatile-mediated indirect defense against insects in rice. The Plant Journal,2008,55:491-503.
36. Julien Serandour, Stephane Reynaud. Ubiquitous Water-Soluble Molecules in Aquatic Plant Exudates Determine Specific Insect Attraction. PLoS ONE,2008, 10(3)3:1-8.
37. JunWei Zhu, Kye-Chung Park. Methyl salicylate, a soybean aphid-indeced plant volatiles attractive to the predator. Journal of Chemical Ecology,2005,8(31): 1733-1740.
38. Karban R, Maron J, Felton GW, Ervin G, Eichenseer H. Herbivore damage to sagebrush induces resistance in wild tobacco:evidence for eavesdropping between plants. Oikos,2003,100:325-32.
39. Kessler A, Baldwin IT. Defensive function of herbivore-induced plant volatile emissions in nature. Science,2001,291:2141-2144.
40. Kessler A, Baldwin IT. Plant responses to insect herbivory:The emerging molecular analysis. Annual Review of Plant Biology,2002,53:299-328.
41. Kerryl. Metlen, Erik T. Aschehoug, Ragan M. Callaway. Plant behavioural ecology: dynamic plasticity in secondary metabolites. Plant, Cell and Environment,2009, 32:641-653.
42. Landolt PJ, Tumlinson JH, Alborn DH. Attraction of Colorado potato beetle (Coleoptera:Chrysomelidae) to damaged and chemically induced potato plants. Environ Entomol,1999,28:973-978.
43. Leslie Hammack. Single and blended maize volatiles as attractants for diabroticite corn rootworm beetles. Journal of Chemical Ecology,2001,7(27):1373-1379.
44. Livy Williams Ⅲ, Cesar Rodriguez-Saona, Sandra C. Castle, Su Zhu,2008. EAG-active herbivore-induced plant volatiles modify behavioral responses and host attack by an egg parasitoid. J Chem Ecol,34:1190-1201.
45. Lou Y, Baldwin IT. Manduca sexta recognition and resistance among allopplyploid Nicotoana host plants. Proceedings of the National Academy of Sciences of the United States of America,2003,100:14581-14586.
46. Marco D'Alessandro, Matthias Held, Yann Triponez, Ted C. J. Turlings. The role of indole and other shikimic acid derived maize volatiles in the attraction of two parasitic wasps. J Chem Ecol,2006,32:2733-2748.
47. Marco D' Alessandro, Ted C. J. Turlings. Advances and challenges in the identification of volatiles that mediate interactions among plants and arthropods. The Royal Society of Chemistry,2006,131:24-32.
48. Marco D'Alessandro, Virginie Brunner, Georg von Merey, Ted C. J. Turlings. Strong attraction of the parasitoid Cotesia marginiventris towards minor volatile compounds of maize. J Chem Ecol,2009,35:999-1008.
49. Martin Pareja, Maria C. B. Moraes, Suzanne J. Clark, Michael A. Birkett & Wilf Powell. Response of the aphid parasitoid Aphidius funebris to volatiles from undamaged and aphid-infested Centaurea nigra. J Chem Ecol,2007,33:695-710.
50. Miklos Toth, Ferenc Szentkiralyi, Jozsef Vuts, Agostino Letardi, Maria Rosaria Tabilio, Gunnhild Jaastad, Geir K. Knudsen. Optimization of a phenylacetaldehyde-based attractant for common green lacewings (Chrysoperla carnea s.l.). J Chem Ecol,2009,35:449-458.
51. M.E. Maffei. Sites of synthesis, biochemistry and functional role of plant volatiles. South African Journal of Botany,2010.
52. Mohamed A. Ibrahim, Anne Nissinen, Jarmo K. Holopainen. Response of Plutella xylostella and its parasitoid Cotesia plutellae to volatile compounds. Journal of Chemical Ecology,2005,31(9):1969-1984.
53. N. Erbilgin, N. E. Gillette, S. R. Mori, J. D. Stein, D. R. Owen, D. L. Wood. Acetophenone as an anti-attractant for the western pine beetle, Dendroctonus Brevicomis LeConte (Coleoptera:Scolytidae). J Chem Ecol,2007,33:817-823.
54. Nicole M. van Dam. Belowground herbivory and plant defenses. The Annual Review of Ecology, Evolution, and Systematics.2009,40:373-391.
55. Nina E. Fatouros, Marcel Dicke, Roland Mumm, Torsten Meiners, Monika Hilker. Foraging behavior of egg parasitoids exploiting chemical information. Behavioral Ecology,2008,11:677-689.
56. Quiroz A, Pettersson J, Pickett JA, Wadhams LJ, Niemeyer HM. Semiochemicals mediating spacing behavior of bird cherry-oat aphid, Rhopalosiphum padi feeding on cereals. J Chem Ecol,1997,23:2599-2607.
57. Ryoko T. Ichiki, Yooichi Kainoh, Soichi Kugimiya, Junji Takabayashi, Satoshi Nakamura. Attraction to herbivore-induced plant volatiles by the host-foraging parasitoid fly Exorista japonica. J Chem Ecol,2008,34:614-621.
58. Raul A. Laumann, Michely F. S. Aquino, Maria C. B. Moraes, Martin Pareja, Miguel Borges. Response of the egg parasitoids Trissolcus basalis and Telenomus podisi to compounds from defensive secretions of stink bugs. J Chem Ecol,2009,35:8-19.
59. Roda AL, Baldwin IT. Molecular technology reveals how the induced direct defenses of plant work. Basic and Applied Ecology,2003,4:15-26.
60. Sabelis, M.W., Van de Baan, H.E.,1983. Location of distant spider mite colonies by phytoseiid predators:demonstration of specific kairomones emitted by Tetranychus urticae and Panonychus ulmi (Acari:Phytoseiidae, Tetranychidae). Entomologia Experimentalis et Applicata,1983,33:303-314.
61. Soichi Kugimiya, Takeshi Shimoda, Jun Tabata, Junji Takabayashi. Present or past herbivory:A screening of volatiles released from Brassica rapa under caterpillar attacks as attractants for the solitary parasitoid, Cotesia vestalis. J Chem Ecol,2010, 36:620-628.
62. TAKESHI SHIMODA, RIKA OZAWA, KOTA SANO, EIZI YANO, JUNJI TAKABAYASHI. The involvement of volatile infochemicals from spider mites and from food-plants in prey location of the generalist predatory mite Neoseiulus californicus. Journal of Chemical Ecology,2005,31(9):2019-2032.
63. Takeshi Shimoda. A key volatile infochemical that elicits a strong olfactory response of the predatory mite Neoseiulus californicus, an important natural enemy of the two-spotted spider mite Tetranychus urticae. Exp Appl Acarol,2001,50:9-22.
64. Turlings TGJ, Wackers F. Recruitment of predators and parasitoids by herbivore-injured plants. Insect Chemical Ecology,2004,2:21-75.
65. Turlings TCJ, Tumlinson JH, Lewis WJ. Exploitation of herbivore-induced plant odors by host-seeking parasitic wasps. Science,1990,250:1251-1253.
66. Virginie Gosset, Nicolas Harmel, Cornelia Gobel, Frederic Francis. Attacks by a piercing-sucking insect (Myzus persicae Sultzer) or a chewing insect (Leptinotarsa decemlineata Say) on potato plants (Solanum tuberosum L.) induce differential changes in volatile compound release and oxylipin synthesis. Journal.of Experimental Botany,2009,1-10.
67. Van Dam NM, Horn M, Mares M, et al. Ontogeny constrains systemic protease inhibitor response in Nicotiana attenuate. Journal of Chemical Ecology,2001, 22(3):547-568.
68. Venkatesan Radhika, Christian Kost, Stefan Bartram, Martin Heil, Wilhelm Boland. Testing the optimal defence hypothesis for two indirect defences:extrafloral nectar and volatile organic compounds. Planta,2008,228:449-457.
69. Vet LEM, Dicke M. Ecology of infochemical use by natural enemies in a tritrophic context. Annu Rev Entomol,1992,37:141-172.
70. XU Tao, ZHOU Qiang, XIA Qiang, ZHANG Wenqing, ZHANG Guren, GU Dexiang. Effects of herbivore-induced rice volatiles on the host selection behavior of brown planthopper, Nilaparvata lugens. Chinese Science Bulletin,2002,47(16):1355-1360.
71. YONG-GEN LOU, BO MA, and JIA-AN CHENG. Attraction of the parasitoid Anagrus nilaparvatae to rice volatiles induced by the rice brown planthopper Nilaparvata lugens. Journal of Chemical Ecology,2005,31(10):2357-2372.
72. Zeyaur R. Khan, David G. James, Charles A.O. Midega, John A. Pickett. Chemical ecology and conservation biological control. Biological Control,2008,45:210-224.
2.杜丽娜,张存莉,朱玮,张高宏。植物次生代谢合成途径及生物学意义。西北林学院学报,2005,20(3)：150-155.
3.李新岗,刘惠霞,黄建。虫害诱导植物防御的分子机理研究进展。应用生态学报,2008,19(4)：893-900.
4.娄永根,程家安。虫害诱导的植物挥发物：基本特性、生态学功能及释放机制。生态学报,2000,20(6)：1097-1106.
5.娄永根,程家安。植物的诱导抗虫性。昆虫学报,1997,40(3)：320-331
6.祝传书,赵惠燕。虫害诱导的植物挥发物。西北农林科技大学学报,2003,31：183-186.
7. Alberto Arab, Jose Roberto Trigo, Andre Luiz Lourencao, Aiarie Michele Peixoto, Fernanda Ramos, Jose Mauricio Simoes Bento. Differential attractiveness of potato tuber volatiles to Phthorimaea operculella (Gelechiidae) and the predator Orius insidiosus (Anthocoridae). J Chem Ecol,2007,33:1845-1855.
8. Antonin Ferry, Sebastien Dugravot, Thomas Delattre, Jean-Philippe Christides, Jacques Auger, Anne-Genevieve Bagneres, Denis Poinsot, Anne-Marie Cortesero. Identification of a widespread monomolecular odor differentially attractive to several Delia radicum ground-dwelling predators in the field. J Chem Ecol,2007, 33:2064-2077.
9. Bernasconi ML, Turlings TCJ, Ambrosetti L, Bassetti P, Dorn S. Herbivore-induced emissions of maize volatiles repel the corn leaf aphid, Rhopalosiphum maidis. Entomol Exp Appl,1998,87:133-142.
10. Carroll MJ, Schmelz EA, MeagherRL, Teal PEA. Attraction of Spodoptera frugiperda Larvae to Volatiles from Herbivore-Damaged Maize Seedlings. J Chem Ecol,2006,32:1911-1924.
11. Cesar R. Rodriguez-Saona, Luis E. Rodriguez-Saona, Christopher J. Frost. Herbivore-induced volatiles in the perennial shrub, Vaccinium corymbosum, and their role in inter-branch signaling. J Chem Ecol,2009,35:163-175.
12. Christopher J. Frost, Mark C. Mescher, John E. Carlson, Consuelo M. De Moraes. Plant defense priming against herbivores:Getting ready for a different battle. Plant physiology,2008a,146:818-824
13. Christopher J.Frost, Mark C. Mescher, John E. Carlson, Consuelo M. De Moraes. Why do distance limitations exist on plant-plant signaling via airborne volatiles? Plant Signaling and Behavior,2008b,3(7):466-468.
14. Colazza S, McElfresh JS, Millar JG. Identification of volatile synomones, induced by Nezara viridula feeding and oviposition on bean spp., that attract the egg parasitoid Trissolcus basalis. J Chem Ecol,2004,30:945-964.
15. David G. James. Further field evaluation of synthetic herbivore-induced plant volatiles as attractants for beneficial insects. Journal of Chemical Ecology,2005,3 (31):481-487.
16. David G. James. Field evaluation of herbivore-induced plant volatiles as attractants for beneficial insects:methyl salicylate and the green lacewing, Chrysopa nigricornis. Journal of Chemical Ecology,2003a,29,1601-1609.
17. David G. James. Synthetic herbivore-induced plant volatiles as attractants for beneficial insects. Environmental Entomology,2003b,32,977-982.
18. David G. James. Synthetic herbivore-induced plant volatiles as field attractants for beneficial insects. The British Crop Protection Council International Congress-Crop Science and Technology,2003c,1217-1222.
19. David G. James. Grape and hop plants sprayed with botanical oil pesticides containing herbivore-induced plant volatiles attract insect predators and parasitoids. Environmental Entomology (submitted for publication),2008.
20. David G James, Castle, S.C., Grasswitz, T., Reyna, V. Using synthetic herbivore-induced plant volatiles to enhance conservation biological control in hops and grapes. In:Proceedings of the 2nd International Symposium on Biological Control of Arthropods, pp.,2005,192-205.
21. David G. James, Grasswitz, T.R. Field attraction of parasitic wasps, Metaphycus sp. and Anagrus spp. using synthetic herbivore-induced plant volatiles. Biocontrol,2005, 50,871-880.
22. David G. James, Price, T.S. Field-testing of methyl salicylate for recruitment and retention of beneficial insects in grapes and hops. Journal of Chemical Ecology,2004, 30,1613-1628.
23. Devendra Kumar Choudhary, Bhavdish N. Johri, Anil Prakash. Volatiles as priming agents that initiate plant growth and defence responses. current science,2008, 94(5):595-604.
24. Dicke, M., van Poecke, R.P., and de Boer, J.G. Inducible indirect defence of plants: From mechanisms to ecological functions. Basic Appl. Ecol.,2003,4:27-42.
25. Douglas M. Light. A pear-derived kairomne with pheromonal potency that attracts male and female codling moth, cydia pomonella (L.). Naturwissenschaften,2001,88: 333-338.
26. Emmanuel Gaquerel, Alexander Weinhold, Ian T. Baldwin. Molecular interactions between the specialist herbivore Manduca sexta (Lepidoptera, Sphigidae) and its natural host Nicotiana attenuata. Ⅷ. An unbiased GCxGC-ToFMS analysis of the plant's. Plant Physiology,2009,149:1408-1423.
27. Eric Rohrig, John Sivinski, Peter Teal, Charles Stuhl, Martin Aluja. A floral-derived compound attractive to the tephritid fruit fly parasitoid Diachasmimorpha longicaudata (Hymenoptera:Braconidae). J Chem Ecol,2008,34:549-557.
28. Gen-ichiro Arimura, Kenji Matsui, Junji Takabayashi. Chemical and molecular ecology of herbivore-induced plant volatiles:Proximate factors and their ultimate functions. Plant Cell Physiology,2009,50(5):911-923.
29. Hamid R. S. Moayeri, Ahmad Ashouri, Henrik F. Brodsgaard, Annie Enkegaard. Males of the predatory mirid bug Macrolophus caliginosus exploit plant volatiles induced by conspecifics as a sexual synomone. Entomologia Experimentalis et Applicata,2007,123(1):49-53.
30. Hayato Ishiwari, Takahisa Suzuki, Taro Maeda. Essential compounds in herbivore-induced plant volatiles that attract the predatory mite Neoseiulus womersleyi. J Chem Ecol,2007,33:1670-1681.
31. Huilin Yu, Yongjun Zhang, Kongming Wu, Xi Wu Gao, Yu Yuan Guo. Field-testing of synthetic herbivore-induced plant volatiles as attractants for beneficial insects. Environmental Entomology,2008,37(6):1410-1415.
32. Ivo Feussner, Claus Wasternack. The lipoxygenase pathway. Ann. Rev. Plant Biol. 2002,53:275-97.
33. Jared G. Ali, Hans T. Alborn, Lukasz L. Stelinski. Subterranean herbivore-induced volatiles released by citrus roots upon feeding by Diaprepes abbreviates recruit entomopathogenic nematodes. J Chem Ecol,2010,36:361-368.
34. Jianing Wei, Lizhong Wang, Junwei Zhu, Sufang Zhang, Owi I. Nandi, Le Kang.. Plants attract parasitic wasps to defend themselves against insect pests by releasing hexenol. PLoS ONE,2007,9:1-7.
35. Joshua S. Yuan, Tobias G. Kollner, Greg Wiggins, Jerome Grant, Jorg Degenhardt, Feng Chen. Molecular and genomic basis of volatile-mediated indirect defense against insects in rice. The Plant Journal,2008,55:491-503.
36. Julien Serandour, Stephane Reynaud. Ubiquitous Water-Soluble Molecules in Aquatic Plant Exudates Determine Specific Insect Attraction. PLoS ONE,2008, 10(3)3:1-8.
37. JunWei Zhu, Kye-Chung Park. Methyl salicylate, a soybean aphid-indeced plant volatiles attractive to the predator. Journal of Chemical Ecology,2005,8(31): 1733-1740.
38. Karban R, Maron J, Felton GW, Ervin G, Eichenseer H. Herbivore damage to sagebrush induces resistance in wild tobacco:evidence for eavesdropping between plants. Oikos,2003,100:325-32.
39. Kessler A, Baldwin IT. Defensive function of herbivore-induced plant volatile emissions in nature. Science,2001,291:2141-2144.
40. Kessler A, Baldwin IT. Plant responses to insect herbivory:The emerging molecular analysis. Annual Review of Plant Biology,2002,53:299-328.
41. Kerryl. Metlen, Erik T. Aschehoug, Ragan M. Callaway. Plant behavioural ecology: dynamic plasticity in secondary metabolites. Plant, Cell and Environment,2009, 32:641-653.
42. Landolt PJ, Tumlinson JH, Alborn DH. Attraction of Colorado potato beetle (Coleoptera:Chrysomelidae) to damaged and chemically induced potato plants. Environ Entomol,1999,28:973-978.
43. Leslie Hammack. Single and blended maize volatiles as attractants for diabroticite corn rootworm beetles. Journal of Chemical Ecology,2001,7(27):1373-1379.
44. Livy Williams Ⅲ, Cesar Rodriguez-Saona, Sandra C. Castle, Su Zhu,2008. EAG-active herbivore-induced plant volatiles modify behavioral responses and host attack by an egg parasitoid. J Chem Ecol,34:1190-1201.
45. Lou Y, Baldwin IT. Manduca sexta recognition and resistance among allopplyploid Nicotoana host plants. Proceedings of the National Academy of Sciences of the United States of America,2003,100:14581-14586.
46. Marco D'Alessandro, Matthias Held, Yann Triponez, Ted C. J. Turlings. The role of indole and other shikimic acid derived maize volatiles in the attraction of two parasitic wasps. J Chem Ecol,2006,32:2733-2748.
47. Marco D' Alessandro, Ted C. J. Turlings. Advances and challenges in the identification of volatiles that mediate interactions among plants and arthropods. The Royal Society of Chemistry,2006,131:24-32.
48. Marco D'Alessandro, Virginie Brunner, Georg von Merey, Ted C. J. Turlings. Strong attraction of the parasitoid Cotesia marginiventris towards minor volatile compounds of maize. J Chem Ecol,2009,35:999-1008.
49. Martin Pareja, Maria C. B. Moraes, Suzanne J. Clark, Michael A. Birkett & Wilf Powell. Response of the aphid parasitoid Aphidius funebris to volatiles from undamaged and aphid-infested Centaurea nigra. J Chem Ecol,2007,33:695-710.
50. Miklos Toth, Ferenc Szentkiralyi, Jozsef Vuts, Agostino Letardi, Maria Rosaria Tabilio, Gunnhild Jaastad, Geir K. Knudsen. Optimization of a phenylacetaldehyde-based attractant for common green lacewings (Chrysoperla carnea s.l.). J Chem Ecol,2009,35:449-458.
51. M.E. Maffei. Sites of synthesis, biochemistry and functional role of plant volatiles. South African Journal of Botany,2010.
52. Mohamed A. Ibrahim, Anne Nissinen, Jarmo K. Holopainen. Response of Plutella xylostella and its parasitoid Cotesia plutellae to volatile compounds. Journal of Chemical Ecology,2005,31(9):1969-1984.
53. N. Erbilgin, N. E. Gillette, S. R. Mori, J. D. Stein, D. R. Owen, D. L. Wood. Acetophenone as an anti-attractant for the western pine beetle, Dendroctonus Brevicomis LeConte (Coleoptera:Scolytidae). J Chem Ecol,2007,33:817-823.
54. Nicole M. van Dam. Belowground herbivory and plant defenses. The Annual Review of Ecology, Evolution, and Systematics.2009,40:373-391.
55. Nina E. Fatouros, Marcel Dicke, Roland Mumm, Torsten Meiners, Monika Hilker. Foraging behavior of egg parasitoids exploiting chemical information. Behavioral Ecology,2008,11:677-689.
56. Quiroz A, Pettersson J, Pickett JA, Wadhams LJ, Niemeyer HM. Semiochemicals mediating spacing behavior of bird cherry-oat aphid, Rhopalosiphum padi feeding on cereals. J Chem Ecol,1997,23:2599-2607.
57. Ryoko T. Ichiki, Yooichi Kainoh, Soichi Kugimiya, Junji Takabayashi, Satoshi Nakamura. Attraction to herbivore-induced plant volatiles by the host-foraging parasitoid fly Exorista japonica. J Chem Ecol,2008,34:614-621.
58. Raul A. Laumann, Michely F. S. Aquino, Maria C. B. Moraes, Martin Pareja, Miguel Borges. Response of the egg parasitoids Trissolcus basalis and Telenomus podisi to compounds from defensive secretions of stink bugs. J Chem Ecol,2009,35:8-19.
59. Roda AL, Baldwin IT. Molecular technology reveals how the induced direct defenses of plant work. Basic and Applied Ecology,2003,4:15-26.
60. Sabelis, M.W., Van de Baan, H.E.,1983. Location of distant spider mite colonies by phytoseiid predators:demonstration of specific kairomones emitted by Tetranychus urticae and Panonychus ulmi (Acari:Phytoseiidae, Tetranychidae). Entomologia Experimentalis et Applicata,1983,33:303-314.
61. Soichi Kugimiya, Takeshi Shimoda, Jun Tabata, Junji Takabayashi. Present or past herbivory:A screening of volatiles released from Brassica rapa under caterpillar attacks as attractants for the solitary parasitoid, Cotesia vestalis. J Chem Ecol,2010, 36:620-628.
62. TAKESHI SHIMODA, RIKA OZAWA, KOTA SANO, EIZI YANO, JUNJI TAKABAYASHI. The involvement of volatile infochemicals from spider mites and from food-plants in prey location of the generalist predatory mite Neoseiulus californicus. Journal of Chemical Ecology,2005,31(9):2019-2032.
63. Takeshi Shimoda. A key volatile infochemical that elicits a strong olfactory response of the predatory mite Neoseiulus californicus, an important natural enemy of the two-spotted spider mite Tetranychus urticae. Exp Appl Acarol,2001,50:9-22.
64. Turlings TGJ, Wackers F. Recruitment of predators and parasitoids by herbivore-injured plants. Insect Chemical Ecology,2004,2:21-75.
65. Turlings TCJ, Tumlinson JH, Lewis WJ. Exploitation of herbivore-induced plant odors by host-seeking parasitic wasps. Science,1990,250:1251-1253.
66. Virginie Gosset, Nicolas Harmel, Cornelia Gobel, Frederic Francis. Attacks by a piercing-sucking insect (Myzus persicae Sultzer) or a chewing insect (Leptinotarsa decemlineata Say) on potato plants (Solanum tuberosum L.) induce differential changes in volatile compound release and oxylipin synthesis. Journal.of Experimental Botany,2009,1-10.
67. Van Dam NM, Horn M, Mares M, et al. Ontogeny constrains systemic protease inhibitor response in Nicotiana attenuate. Journal of Chemical Ecology,2001, 22(3):547-568.
68. Venkatesan Radhika, Christian Kost, Stefan Bartram, Martin Heil, Wilhelm Boland. Testing the optimal defence hypothesis for two indirect defences:extrafloral nectar and volatile organic compounds. Planta,2008,228:449-457.
69. Vet LEM, Dicke M. Ecology of infochemical use by natural enemies in a tritrophic context. Annu Rev Entomol,1992,37:141-172.
70. XU Tao, ZHOU Qiang, XIA Qiang, ZHANG Wenqing, ZHANG Guren, GU Dexiang. Effects of herbivore-induced rice volatiles on the host selection behavior of brown planthopper, Nilaparvata lugens. Chinese Science Bulletin,2002,47(16):1355-1360.
71. YONG-GEN LOU, BO MA, and JIA-AN CHENG. Attraction of the parasitoid Anagrus nilaparvatae to rice volatiles induced by the rice brown planthopper Nilaparvata lugens. Journal of Chemical Ecology,2005,31(10):2357-2372.
72. Zeyaur R. Khan, David G. James, Charles A.O. Midega, John A. Pickett. Chemical ecology and conservation biological control. Biological Control,2008,45:210-224.