亚洲小车蝗聚集爆发的化学信息学研究
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摘要
亚洲小车蝗(Oedaleus asiaticus B.-Bienko)是我国北方草原和农牧交错地带最重要的优势种蝗虫之一,尤其近些年来亚洲小车蝗在各地起飞严重,作为前锋聚集迁移,甚至成群结队出现在城区中心,实属历史罕见。因此,加强亚洲小车蝗聚集行为的研究,对于了解其爆发成灾机制具有重要的意义。我们在室内外首次从化学信息学的角度对亚洲小车蝗成虫的聚集行为进行系统地研究,探明引起亚洲小车蝗成虫聚集的化学信息物的主要来源及化学组成,为进一步的活性成分的确定及探索利用化学信息素调控其行为的无公害防治途径提供依据。主要结果如下:
1.亚洲小车蝗成虫的聚集行为表现不明显,只是在幼成虫时期表现出一定的聚集性,老熟雌成虫也有部分聚集行为反应。
2.除雄性幼成虫对相同发育阶段的异性(雌性幼成虫粪便)有聚集反应外,各发育阶段的雌、雄成虫均喜欢选择来自同性的体表或粪便挥发物。
3.亚洲小车蝗雌、雄成虫对各味源的选择趋势和释放头数(5或10头)没有明显关系,因此反应趋势受试虫密度影响不大。
4.亚洲小车蝗喜欢选择在已有卵囊和泡沫的有卵杯中产卵,而不喜欢在没有卵囊和泡沫的无卵杯中产卵,表明该蝗可利用挥发性气味区分有卵杯和无卵杯,并且可被其卵囊和泡沫挥发物吸引产卵。这些结果初步显示了亚洲小车蝗存在聚集产卵的现象,卵囊及泡沫物质有可能是调控其产卵行为的信息化合物的来源。
5.亚洲小车蝗雌、雄成虫体表挥发物的成分都比较少。雌成虫体表挥发物含有苯类、环烯类、萜烯类、醛类,雄成虫体表挥发物只检测到环烯类、萜烯类。在这两个不同性别来源的样品中,α-蒎烯、β-蒎烯是共同含有的成分。
6.亚洲小车蝗雌、雄成虫粪便挥发物的成分都比较多。雌成虫粪便挥发物中含有酮类、醛类和萜烯类,雄成虫粪便挥发物含有酯类、醛类、醇类、环烯类、酮类、萜烯类。雌、雄成虫粪便挥发物有很大的相似处,都含有醛类、酮类、萜烯类化合物,2-甲基丙醛、3-甲基丁醛、2-甲基丁醛、α-蒎烯是两类不同性别来源样品中共有的成分。
Oedaleus asiaticus B.-Bienko(Orthoptera: Acrididae),one of the most dominant grasshoppers on steppes and agriculture-animal husbandry ecotones in northern China,occurred seriously in many areas in recent years. This species of grasshopper migrated as a vanguard and even swarmed into the centers of cities which rarely happened in the history. Therefore, studying the aggregation behavior of the grasshopper is crucial to explore the causes eliciting its outbreak. Here, we investigated the aggregation behavior of adult Oedaleus asiaticus from the point of semiochemicals both in the laboratory and in the field,and try to explore the main sources and chemical compositions of aggregation pheromones in order to characterize the active components of pheromones and exploit the environmentally-friendly managements through mediating aggregation behaviors by using pheromones in future. The main results are as follows:
1. The adult of O. asiaticus showed less obvious aggregation behavior, which behaved in aggregation only in the period of young adults. The old female adults were also found to aggregate partially.
2. The young or old female adults of O. asiaticus preferred to be within the airborne volatiles of live adults and feces of same sex but were indifferent to volatiles of male adults. Conversely,the young or old male adults of O. asiaticus preferred to be within the airborne volatiles of live adults and feces of same sex but were indifferent to volatiles of female adults,except the young male adults preferred to be within the airborne volatiles of females (young female feces) with same developmental stage as males.
3. The behavior responses of O. asiaticus to different odours were not correlated to releasing ways of the grasshoppers(in groups of 5 or 10).
4. The gravid females were found to oviposit preferentially in cups with eggs relative to in cups without eggs in a two-choice experiment. This suggests that the grasshoppers are able to use volatile cues to discriminate between cups with and without eggs, and can be attracted to to aggregate and oviposit by odours from egg pods and foams. These results preliminary showed that this species of grasshopper behaved in aggregative oviposition, and the egg pod and foam are possible sources of the primer pheromone mediating the ovipositing behavior of the grasshoppers.
5. GC-MS analysis identified less compounds in the airborne volatiles trapped from live female and male adults of O. asiaticus, respectively. Benzenes, cycloalkenes, terpenes, and aldehydes were found in the volatiles of live female adults. Cycloalkenes and terpenes were contained in the volatiles of live male adults. The common compounds found in these two volatile samples wereα-pinene andβ-pinene.
6. GC-MS analysis identified many compounds in the fecal volatiles from female and male adults of O. asiaticus, respectively. Ketones, aldehydes, and terpenes were found in the fecal volatiles of female adults. Esters, aldehydes, alcohols, cycloalkenes, ketones, and terpenes were present in the fecal volatiles of male adults. Aldehydes, ketones and terpene were common kinds of compounds in these two volatile samples. The composition of fecal volatile substances of female adult was much similar as one of male adult, sharing 2-methyl-propionaldehyde, 3-methyl-butyraldehyde, 2-methyl-butyraldehyde, andα- pinene.
1.亚洲小车蝗成虫的聚集行为表现不明显,只是在幼成虫时期表现出一定的聚集性,老熟雌成虫也有部分聚集行为反应。
2.除雄性幼成虫对相同发育阶段的异性(雌性幼成虫粪便)有聚集反应外,各发育阶段的雌、雄成虫均喜欢选择来自同性的体表或粪便挥发物。
3.亚洲小车蝗雌、雄成虫对各味源的选择趋势和释放头数(5或10头)没有明显关系,因此反应趋势受试虫密度影响不大。
4.亚洲小车蝗喜欢选择在已有卵囊和泡沫的有卵杯中产卵,而不喜欢在没有卵囊和泡沫的无卵杯中产卵,表明该蝗可利用挥发性气味区分有卵杯和无卵杯,并且可被其卵囊和泡沫挥发物吸引产卵。这些结果初步显示了亚洲小车蝗存在聚集产卵的现象,卵囊及泡沫物质有可能是调控其产卵行为的信息化合物的来源。
5.亚洲小车蝗雌、雄成虫体表挥发物的成分都比较少。雌成虫体表挥发物含有苯类、环烯类、萜烯类、醛类,雄成虫体表挥发物只检测到环烯类、萜烯类。在这两个不同性别来源的样品中,α-蒎烯、β-蒎烯是共同含有的成分。
6.亚洲小车蝗雌、雄成虫粪便挥发物的成分都比较多。雌成虫粪便挥发物中含有酮类、醛类和萜烯类,雄成虫粪便挥发物含有酯类、醛类、醇类、环烯类、酮类、萜烯类。雌、雄成虫粪便挥发物有很大的相似处,都含有醛类、酮类、萜烯类化合物,2-甲基丙醛、3-甲基丁醛、2-甲基丁醛、α-蒎烯是两类不同性别来源样品中共有的成分。
Oedaleus asiaticus B.-Bienko(Orthoptera: Acrididae),one of the most dominant grasshoppers on steppes and agriculture-animal husbandry ecotones in northern China,occurred seriously in many areas in recent years. This species of grasshopper migrated as a vanguard and even swarmed into the centers of cities which rarely happened in the history. Therefore, studying the aggregation behavior of the grasshopper is crucial to explore the causes eliciting its outbreak. Here, we investigated the aggregation behavior of adult Oedaleus asiaticus from the point of semiochemicals both in the laboratory and in the field,and try to explore the main sources and chemical compositions of aggregation pheromones in order to characterize the active components of pheromones and exploit the environmentally-friendly managements through mediating aggregation behaviors by using pheromones in future. The main results are as follows:
1. The adult of O. asiaticus showed less obvious aggregation behavior, which behaved in aggregation only in the period of young adults. The old female adults were also found to aggregate partially.
2. The young or old female adults of O. asiaticus preferred to be within the airborne volatiles of live adults and feces of same sex but were indifferent to volatiles of male adults. Conversely,the young or old male adults of O. asiaticus preferred to be within the airborne volatiles of live adults and feces of same sex but were indifferent to volatiles of female adults,except the young male adults preferred to be within the airborne volatiles of females (young female feces) with same developmental stage as males.
3. The behavior responses of O. asiaticus to different odours were not correlated to releasing ways of the grasshoppers(in groups of 5 or 10).
4. The gravid females were found to oviposit preferentially in cups with eggs relative to in cups without eggs in a two-choice experiment. This suggests that the grasshoppers are able to use volatile cues to discriminate between cups with and without eggs, and can be attracted to to aggregate and oviposit by odours from egg pods and foams. These results preliminary showed that this species of grasshopper behaved in aggregative oviposition, and the egg pod and foam are possible sources of the primer pheromone mediating the ovipositing behavior of the grasshoppers.
5. GC-MS analysis identified less compounds in the airborne volatiles trapped from live female and male adults of O. asiaticus, respectively. Benzenes, cycloalkenes, terpenes, and aldehydes were found in the volatiles of live female adults. Cycloalkenes and terpenes were contained in the volatiles of live male adults. The common compounds found in these two volatile samples wereα-pinene andβ-pinene.
6. GC-MS analysis identified many compounds in the fecal volatiles from female and male adults of O. asiaticus, respectively. Ketones, aldehydes, and terpenes were found in the fecal volatiles of female adults. Esters, aldehydes, alcohols, cycloalkenes, ketones, and terpenes were present in the fecal volatiles of male adults. Aldehydes, ketones and terpene were common kinds of compounds in these two volatile samples. The composition of fecal volatile substances of female adult was much similar as one of male adult, sharing 2-methyl-propionaldehyde, 3-methyl-butyraldehyde, 2-methyl-butyraldehyde, andα- pinene.
引文
1 Nolte DJ, Eggers SH, May IR. A locust pheromone:locustol[J]. Journal of Insect Physiology, 1973, Vol.19:1547-1554
2杜家纬.昆虫信息素及其应用[M].北京:中国林业出版社, 1988
3杜家纬.我们在昆虫信息素方面的研究进展[J].中国科学院院刊, 1991, Vol.6(4):326-328
4苏茂文,张钟宁.昆虫信息化学物质的应用进展[J].昆虫知识, 2007, Vol.44(4):477-485
5刘孟英.昆虫信息素研究及应用的新进展[J].昆虫知识, 1994, Vol.31(1):56-59
6 Roessingh P, Bouaichi A, Simpson ST. Effects of sensory stimuli on the behavioural phase state of desert locust, Schistocerca gregaria[J]. Journal of Insect Physiology, 1998,Vol.44:883-893
7姜勇,雷朝亮,张钟宁.昆虫聚集信息素[J].昆虫学报, 2002, Vol.45(6):822-832
8 Ross DW, Daterman GE. Using pheromone-baited traps to control the amount and distribution of tree mortality during outbreaks of the Douglas fir beetle[J]. Forest Science, 1997, Vol.43(1):65-70
9 Byer JA. Pheromones and chemical ecology of locusts[J]. Biological Reviews, 1991, Vol.66:347-78
10 Hassanali A, Njagi PGN, Bashir MO. Chemical ecology of locusts and related acridids[J]. Annual Review of Entomology, 2005, Vol.50:223–245
11严毓骅.试论拓宽生物防治范围发展虫害可持续治理[J].昆虫学报,1998, Vol.41(增刊): 1-7
12张龙,严毓骅.持续治理飞蝗灾害的新对策[J].昆虫学报, 2000 ,Vol.43(增刊):180-185
13 Niassy A, Torto B, Njagi PGN, Hassanali A, Obeng-Ofori D, Ayertey JN. Intra-and interspecific aggregation responses of Locusta migratoria migratorioides and Schistocerca gregaria and a comparison of their pheromone emissions[J]. Journal of chemical ecology, 1999, Vol.25:1029-1042
14石旺鹏.蝗虫化学信息物质研究进展[J].昆虫知识, 2005, Vol.42(3):244-249
15 Obeng-Ofori D, Torto B, Njagi PGN, Hassanali A, Amiani H. Fecal volatiles as part of the aggregation pheromone complex of the desert locust, Schistocerca gregaria (Forsk?l) (Orthoptera: Acrididae)[J]. Journal of chemical ecology, 1994, Vol.20:2077-2087
16 Hassanali A, Torto B. Grasshoppers and locusts. In: Hardie J, Minks AK. ed. Pheromones of Non-Lepidopteran Insects Associated with Agricultural Plants[M]. Oxon: CABI Publ, 1999, pp.305-328
17 Torto B, Obeng-Ofori D, Njagi PGN, Hassanali A, Amiani H. Aggregation pheromone systemof adult gregarious desert locust, Schistocerca gregaria (Forsk?l)(Orthoptera: Acrididae)[J]. Journal of chemical ecology, 1994, Vol.20:1749-1762
18 Torto B, Njagi PGN, Hassanali A, Amiani H. Aggregation pheromone system of nymphal gregarious desert locust, Schistocerca gregaria (Forsk?l) (Orthoptera: Acrididae)[J]. Journal of chemical ecology, 1996, Vol.22:2273-2281
19 Rai MM, Hassanali A , Saini RK, Odongo H , Kahoro H. Identification of components of the oviposition aggregation pheromone of the gregarious desert locust,Schistocerca gregaria (Forsk?l)[J]. Journal of Insect Physiology, 1997,Vol.43:83-87
20 McCaffery AR, Simpson SJ, Islam M, Roessingh P. A gregarising factor present in the egg pod foam of the desert locust Schistocerca gregaria[J]. Journal of Experimental Biology, 1998, Vol.201:347-363
21 Gillett SD, Packham JM, Papworth SJ. Possible pheromonal effects on aggregation and dispersion in the desert locust, Schistocerca gregaria (Forsk?l)[J]. Acrida, 1976, Vol.5:287-297
22 Ferenz H-J, Seidelmann K. Pheromones in relation to aggregation and reproduction in desert locusts[J]. Physiological entomology, 2003, Vol.28:11-18
23 Assad YOH, Torto B, Hassanali A, Njagi PGN, Bashir NHH, Mahamat H. Seasonal variation in the essential oil composition of Commiphora quadricincta and its effect on the maturation of immature adults of the desert locust, Schistocerca gregaria. Phytochemistry, 1997, Vol.44:833-842
24 Obeng-Ofori D, Njagi PGN, Torto B, Hassanali A, Amiani H. Sex differentiation studies relating to releaser pheromones of the desert locust, Schistocerca gregaria (Forsk?l)[J]. Entomologia Experimentalis et Applicata, 1994, Vol.73:85-91
25 Obeng-Ofori D, Torto B, Hassanali A. Evidence for mediation of two releaser pheromones in aggregation behaviour of the gregarious desert locust Schistocerca gregaria (Orthoptera: Acrididae)[J]. Journal of chemical ecology, 1993, Vol.19:1665-1676
26 Pener MP, Yerushalmi Y. The physiology of locust phase polymorphism: an update[J]. Journal of Insect Physiology, 1998, Vol.44: 365-377
27 Deng AL, Torto B, Hassanali A, Ali EE. Effects of shifting to crowded or solitary conditions on pheromone release and morphometrics of the desert locust, Schistocerca gregaria(Forsk?l)(Orthoptera:Acrididae)[J]. Journal of Insect Physiology, 1996, Vol.42:771-776
28 Siedelmann K, Luber K, Ferenz H-J. Analysis of release and role of benzyl cyanide in male desert locusts, Schistocerca gregaria[J]. Journal of chemical ecology, 2000,Vol.26:1897-1910
29 Uvarov BP. Grasshoppers and Locusts[M]. London: Cent. Overseas Pest Research, 1977, Vol.2,613 pp
30 Deng AL. Studies of some factors that influence phase dynamics of the desert locust, Schistocerca gregaria (Forsk?l) (Orthoptera: Acridoidae)[D]. Sudan:University of Khartoum, 1995,176 pp
31 Saini RK, Rai MM, Hassanali A, Wawiye J, Odongo H. Semiochemicals from froth of egg pods attract ovipositing female Schistocerca gregaria[J]. Journal of Insect Physiology, 1995, Vol.41:711-716
32 Fuzeau-Braesch S, Genin E, Jullien R, Knowles E, Papin C. Composition and role of volatile substances in atmosphere surrounding two gregarious locusts, Locusta migratoria and Schistocerca gregaria[J]. Journal of chemical ecology, 1988, Vol.14:1023-1032
33 El-Bashir S, Abdel-Rahman HE. Factors inducing phase change in the desert locust, Schistocerca gregaria (Forsk?l), at recession and breeding habitats[C]. In: 19th Annual Report of ICIPE, Nairobi: ICIPE Sci, 1991, pp.85-86
34 Johnston HB, Buxton DR. Field observations on locusts in East Africa[J]. Anti- Locust Bull, 1949, 5 pp
35石旺鹏,严毓骅,张龙,王旭东.东亚飞蝗粪便挥发物对其蝗蝻的聚集作用初报[J].中国农业大学学报, 2000, Vol.5(5):54-58.
36石旺鹏.化学信息素对东亚飞蝗聚集行为的调控作用[D].北京:中国农业大学博士学位论文, 2000
37石旺鹏,严毓骅,闫跃英.东亚飞蝗不同来源聚集信息素的电生理活性[J].昆虫知识, 2003, Vol.40(2):145-150
38邵奇妙.东亚飞蝗对其粪便中信息素的行为反应与信息接收机理的初步研究[D].北京:中国农业大学硕士学位论文, 2001
39郭志永.东亚飞蝗粪便挥发物中活性成份鉴定[D].北京:中国农业大学硕士学位论文, 2004
40吴效东.乌兰察布市蝗虫发生规律及危害特点[J].内蒙古农业科技, 2007, Vol.(6):69-70
41王殿臣.苏尼特左旗草原蝗虫发生及危害现状的研究[J].内蒙古草业, 2004, Vol.2 (16):14-15
42潘建梅.内蒙古草原蝗虫发生原因及防治对策[J].中国草地, 2002, Vol.24(6):66-69
43全国畜牧兽医总站草业饲料处.内蒙古草原蝗虫形成大暴发之势[J].草业科学, 2002, Vol.19(4):78-78
44许富祯,孟正平,郭永华,申集平,韩常在.乌兰察布市农牧交错区亚洲小车蝗发生与防治[J].内蒙古农业科技, 2005, Vol.7:384-387
45冯光翰,樊树喜,刘秋芳,杨延彪,马隆喜,曹国顺.室外罩笼条件下几种草原蝗虫的食量测定[J].草地学报, 1995, Vol.3(3):230-235
46 Kang L, Chen Y L. Dynamics of grasshopper communities under different grazing intensities in Inner Mongolia steppes[J]. Entomologia Sinica, 1995,Vol.2:265-281
47刘举鹏.中国蝗虫鉴定手册[M].北京:天则出版社, 1990
48都瓦拉.草原蝗灾遥感监测与评估方法研究[D].呼和浩特:内蒙古师范大学硕士学位论文, 2006
49关敬群,魏增柱.亚洲小车蝗(Oedaleus asiaticus B.-bienko)食量测定[J].昆虫知识, 1989, Vol.26(1):8-10
50刘玲,郭安红. 2004年内蒙古草原蝗虫大发生的气象生态条件分析[J].气象, 2004, Vol.30(11):55-57
51蒋湘,买买提明,张龙.夜间迁飞的亚洲小车蝗[J].草地学报, 2003, Vol.11(1):75-77
52乔峰.蝗灾北移的主力军-亚洲小车蝗[J].人与生物圈, 2005, Vol.(3):23
53颜忠诚.草原蝗虫的栖境选择[D].北京:中国科学院研究生院(动物研究所)博士学位论文, 1995
54仵均祥.农业昆虫学[M].北京:中国农业出版社, 2002
55牛虎力,周强.草原蝗虫的红外灯光诱集实验初报[C].中国植物保护学会第九届会员代表大会暨2005年学术年会农业生物灾害预防与控制研究, 2005
56 Cheke RA. A migrant pest in the Sahel: the Senegalese Grasshopper Oedaleus Senegalensis. Philosophical Transactions of the Royal Society of London[J]. Series B, Biological Sciences, 1990, 328:1251,539-553
57 Riley JR, Reynolds DR. Nocturnal grasshopper migration in west Africa: transport and concentration by the wind, and the implications for air to air control[M]. Philosophical Transactions of the Royal Society of London. Series B,biological sciences, 1990, 328(1251),655-672
58 Malual AG, Hassanali A, Torto B, Assad YOH, Njagi PGN. The nature of the gregarizing signal responsible for maternal transfer of phase to the offspring in the desert locust, Schistocerca gregaria[J]. Journal of chemical ecology, 2001, Vol.27:1423-1435
59 Norris MJ. Laboratory experiments on oviposition responses of the desert locust,Schistocerca gregaria(Forsk?l)[J]. Anti-Locust Bull, 1968, Vol.43:1-47
60 Bashir MO, Hassanali A, Rai MM.Saini RK. Changing oviposition preferences of the desert locust, Schistocerca gregaria, suggest a strong species predisposition for gregarization[J]. Journal of chemical ecology, 2000, Vol.26:1721-1733
61 Torto B, Assad YOH, Njagi PGN, Hassanali A. Evidence for additional pheromonalcomponents mediating oviposition aggregation in Schistocerca gregaria[J]. Journal of chemical ecology, 1999, Vol.25:835-845
62 Seidelmann K. Courtship inhibition pheromone in desert locusts, Schistocerca gregaria[J]. Journal of Insect Physiology, 2002, Vol.48:991-996
63 Seidelmann K, Weinert HJ. Wings and legs are production sites for the desert locust courtship-inhibition pheromone, phenylacetonitrile[J]. Journal of Insect Physiology, 2003, Vol.49:1125-1133
64阎凤鸣.化学生态学[M].北京:科学出版社,2003
65任琴.马尾松快速诱导抗性及化学信号物质的研究[D].北京:北京林业大学博士论文, 2006
66张凤娟.光肩星天牛对槭树挥发物的响应及虫害诱导信号物质研究[D].北京:北京林业大学博士论文,2006
67 Rono E, Njagi PGN, Bashir MO, Hassanali A.Concentration-dependent parsimonious releaser roles of gregarious male pheromone of the desert locust, Schistocerca gregaria[J]. Journal of Insect Physiology, 2008, Vol.54:162-168
68 Assad YOH, Hassanali A, Torto B, Mahamat H, Bashir NHH, Bashir SE. Effects of fifth-instar volatiles on sexual maturation of adult desert locust Schistocerca gregaria[J]. Journal of Chemical Ecology, 1997, Vol.23 :1373-1388
2杜家纬.昆虫信息素及其应用[M].北京:中国林业出版社, 1988
3杜家纬.我们在昆虫信息素方面的研究进展[J].中国科学院院刊, 1991, Vol.6(4):326-328
4苏茂文,张钟宁.昆虫信息化学物质的应用进展[J].昆虫知识, 2007, Vol.44(4):477-485
5刘孟英.昆虫信息素研究及应用的新进展[J].昆虫知识, 1994, Vol.31(1):56-59
6 Roessingh P, Bouaichi A, Simpson ST. Effects of sensory stimuli on the behavioural phase state of desert locust, Schistocerca gregaria[J]. Journal of Insect Physiology, 1998,Vol.44:883-893
7姜勇,雷朝亮,张钟宁.昆虫聚集信息素[J].昆虫学报, 2002, Vol.45(6):822-832
8 Ross DW, Daterman GE. Using pheromone-baited traps to control the amount and distribution of tree mortality during outbreaks of the Douglas fir beetle[J]. Forest Science, 1997, Vol.43(1):65-70
9 Byer JA. Pheromones and chemical ecology of locusts[J]. Biological Reviews, 1991, Vol.66:347-78
10 Hassanali A, Njagi PGN, Bashir MO. Chemical ecology of locusts and related acridids[J]. Annual Review of Entomology, 2005, Vol.50:223–245
11严毓骅.试论拓宽生物防治范围发展虫害可持续治理[J].昆虫学报,1998, Vol.41(增刊): 1-7
12张龙,严毓骅.持续治理飞蝗灾害的新对策[J].昆虫学报, 2000 ,Vol.43(增刊):180-185
13 Niassy A, Torto B, Njagi PGN, Hassanali A, Obeng-Ofori D, Ayertey JN. Intra-and interspecific aggregation responses of Locusta migratoria migratorioides and Schistocerca gregaria and a comparison of their pheromone emissions[J]. Journal of chemical ecology, 1999, Vol.25:1029-1042
14石旺鹏.蝗虫化学信息物质研究进展[J].昆虫知识, 2005, Vol.42(3):244-249
15 Obeng-Ofori D, Torto B, Njagi PGN, Hassanali A, Amiani H. Fecal volatiles as part of the aggregation pheromone complex of the desert locust, Schistocerca gregaria (Forsk?l) (Orthoptera: Acrididae)[J]. Journal of chemical ecology, 1994, Vol.20:2077-2087
16 Hassanali A, Torto B. Grasshoppers and locusts. In: Hardie J, Minks AK. ed. Pheromones of Non-Lepidopteran Insects Associated with Agricultural Plants[M]. Oxon: CABI Publ, 1999, pp.305-328
17 Torto B, Obeng-Ofori D, Njagi PGN, Hassanali A, Amiani H. Aggregation pheromone systemof adult gregarious desert locust, Schistocerca gregaria (Forsk?l)(Orthoptera: Acrididae)[J]. Journal of chemical ecology, 1994, Vol.20:1749-1762
18 Torto B, Njagi PGN, Hassanali A, Amiani H. Aggregation pheromone system of nymphal gregarious desert locust, Schistocerca gregaria (Forsk?l) (Orthoptera: Acrididae)[J]. Journal of chemical ecology, 1996, Vol.22:2273-2281
19 Rai MM, Hassanali A , Saini RK, Odongo H , Kahoro H. Identification of components of the oviposition aggregation pheromone of the gregarious desert locust,Schistocerca gregaria (Forsk?l)[J]. Journal of Insect Physiology, 1997,Vol.43:83-87
20 McCaffery AR, Simpson SJ, Islam M, Roessingh P. A gregarising factor present in the egg pod foam of the desert locust Schistocerca gregaria[J]. Journal of Experimental Biology, 1998, Vol.201:347-363
21 Gillett SD, Packham JM, Papworth SJ. Possible pheromonal effects on aggregation and dispersion in the desert locust, Schistocerca gregaria (Forsk?l)[J]. Acrida, 1976, Vol.5:287-297
22 Ferenz H-J, Seidelmann K. Pheromones in relation to aggregation and reproduction in desert locusts[J]. Physiological entomology, 2003, Vol.28:11-18
23 Assad YOH, Torto B, Hassanali A, Njagi PGN, Bashir NHH, Mahamat H. Seasonal variation in the essential oil composition of Commiphora quadricincta and its effect on the maturation of immature adults of the desert locust, Schistocerca gregaria. Phytochemistry, 1997, Vol.44:833-842
24 Obeng-Ofori D, Njagi PGN, Torto B, Hassanali A, Amiani H. Sex differentiation studies relating to releaser pheromones of the desert locust, Schistocerca gregaria (Forsk?l)[J]. Entomologia Experimentalis et Applicata, 1994, Vol.73:85-91
25 Obeng-Ofori D, Torto B, Hassanali A. Evidence for mediation of two releaser pheromones in aggregation behaviour of the gregarious desert locust Schistocerca gregaria (Orthoptera: Acrididae)[J]. Journal of chemical ecology, 1993, Vol.19:1665-1676
26 Pener MP, Yerushalmi Y. The physiology of locust phase polymorphism: an update[J]. Journal of Insect Physiology, 1998, Vol.44: 365-377
27 Deng AL, Torto B, Hassanali A, Ali EE. Effects of shifting to crowded or solitary conditions on pheromone release and morphometrics of the desert locust, Schistocerca gregaria(Forsk?l)(Orthoptera:Acrididae)[J]. Journal of Insect Physiology, 1996, Vol.42:771-776
28 Siedelmann K, Luber K, Ferenz H-J. Analysis of release and role of benzyl cyanide in male desert locusts, Schistocerca gregaria[J]. Journal of chemical ecology, 2000,Vol.26:1897-1910
29 Uvarov BP. Grasshoppers and Locusts[M]. London: Cent. Overseas Pest Research, 1977, Vol.2,613 pp
30 Deng AL. Studies of some factors that influence phase dynamics of the desert locust, Schistocerca gregaria (Forsk?l) (Orthoptera: Acridoidae)[D]. Sudan:University of Khartoum, 1995,176 pp
31 Saini RK, Rai MM, Hassanali A, Wawiye J, Odongo H. Semiochemicals from froth of egg pods attract ovipositing female Schistocerca gregaria[J]. Journal of Insect Physiology, 1995, Vol.41:711-716
32 Fuzeau-Braesch S, Genin E, Jullien R, Knowles E, Papin C. Composition and role of volatile substances in atmosphere surrounding two gregarious locusts, Locusta migratoria and Schistocerca gregaria[J]. Journal of chemical ecology, 1988, Vol.14:1023-1032
33 El-Bashir S, Abdel-Rahman HE. Factors inducing phase change in the desert locust, Schistocerca gregaria (Forsk?l), at recession and breeding habitats[C]. In: 19th Annual Report of ICIPE, Nairobi: ICIPE Sci, 1991, pp.85-86
34 Johnston HB, Buxton DR. Field observations on locusts in East Africa[J]. Anti- Locust Bull, 1949, 5 pp
35石旺鹏,严毓骅,张龙,王旭东.东亚飞蝗粪便挥发物对其蝗蝻的聚集作用初报[J].中国农业大学学报, 2000, Vol.5(5):54-58.
36石旺鹏.化学信息素对东亚飞蝗聚集行为的调控作用[D].北京:中国农业大学博士学位论文, 2000
37石旺鹏,严毓骅,闫跃英.东亚飞蝗不同来源聚集信息素的电生理活性[J].昆虫知识, 2003, Vol.40(2):145-150
38邵奇妙.东亚飞蝗对其粪便中信息素的行为反应与信息接收机理的初步研究[D].北京:中国农业大学硕士学位论文, 2001
39郭志永.东亚飞蝗粪便挥发物中活性成份鉴定[D].北京:中国农业大学硕士学位论文, 2004
40吴效东.乌兰察布市蝗虫发生规律及危害特点[J].内蒙古农业科技, 2007, Vol.(6):69-70
41王殿臣.苏尼特左旗草原蝗虫发生及危害现状的研究[J].内蒙古草业, 2004, Vol.2 (16):14-15
42潘建梅.内蒙古草原蝗虫发生原因及防治对策[J].中国草地, 2002, Vol.24(6):66-69
43全国畜牧兽医总站草业饲料处.内蒙古草原蝗虫形成大暴发之势[J].草业科学, 2002, Vol.19(4):78-78
44许富祯,孟正平,郭永华,申集平,韩常在.乌兰察布市农牧交错区亚洲小车蝗发生与防治[J].内蒙古农业科技, 2005, Vol.7:384-387
45冯光翰,樊树喜,刘秋芳,杨延彪,马隆喜,曹国顺.室外罩笼条件下几种草原蝗虫的食量测定[J].草地学报, 1995, Vol.3(3):230-235
46 Kang L, Chen Y L. Dynamics of grasshopper communities under different grazing intensities in Inner Mongolia steppes[J]. Entomologia Sinica, 1995,Vol.2:265-281
47刘举鹏.中国蝗虫鉴定手册[M].北京:天则出版社, 1990
48都瓦拉.草原蝗灾遥感监测与评估方法研究[D].呼和浩特:内蒙古师范大学硕士学位论文, 2006
49关敬群,魏增柱.亚洲小车蝗(Oedaleus asiaticus B.-bienko)食量测定[J].昆虫知识, 1989, Vol.26(1):8-10
50刘玲,郭安红. 2004年内蒙古草原蝗虫大发生的气象生态条件分析[J].气象, 2004, Vol.30(11):55-57
51蒋湘,买买提明,张龙.夜间迁飞的亚洲小车蝗[J].草地学报, 2003, Vol.11(1):75-77
52乔峰.蝗灾北移的主力军-亚洲小车蝗[J].人与生物圈, 2005, Vol.(3):23
53颜忠诚.草原蝗虫的栖境选择[D].北京:中国科学院研究生院(动物研究所)博士学位论文, 1995
54仵均祥.农业昆虫学[M].北京:中国农业出版社, 2002
55牛虎力,周强.草原蝗虫的红外灯光诱集实验初报[C].中国植物保护学会第九届会员代表大会暨2005年学术年会农业生物灾害预防与控制研究, 2005
56 Cheke RA. A migrant pest in the Sahel: the Senegalese Grasshopper Oedaleus Senegalensis. Philosophical Transactions of the Royal Society of London[J]. Series B, Biological Sciences, 1990, 328:1251,539-553
57 Riley JR, Reynolds DR. Nocturnal grasshopper migration in west Africa: transport and concentration by the wind, and the implications for air to air control[M]. Philosophical Transactions of the Royal Society of London. Series B,biological sciences, 1990, 328(1251),655-672
58 Malual AG, Hassanali A, Torto B, Assad YOH, Njagi PGN. The nature of the gregarizing signal responsible for maternal transfer of phase to the offspring in the desert locust, Schistocerca gregaria[J]. Journal of chemical ecology, 2001, Vol.27:1423-1435
59 Norris MJ. Laboratory experiments on oviposition responses of the desert locust,Schistocerca gregaria(Forsk?l)[J]. Anti-Locust Bull, 1968, Vol.43:1-47
60 Bashir MO, Hassanali A, Rai MM.Saini RK. Changing oviposition preferences of the desert locust, Schistocerca gregaria, suggest a strong species predisposition for gregarization[J]. Journal of chemical ecology, 2000, Vol.26:1721-1733
61 Torto B, Assad YOH, Njagi PGN, Hassanali A. Evidence for additional pheromonalcomponents mediating oviposition aggregation in Schistocerca gregaria[J]. Journal of chemical ecology, 1999, Vol.25:835-845
62 Seidelmann K. Courtship inhibition pheromone in desert locusts, Schistocerca gregaria[J]. Journal of Insect Physiology, 2002, Vol.48:991-996
63 Seidelmann K, Weinert HJ. Wings and legs are production sites for the desert locust courtship-inhibition pheromone, phenylacetonitrile[J]. Journal of Insect Physiology, 2003, Vol.49:1125-1133
64阎凤鸣.化学生态学[M].北京:科学出版社,2003
65任琴.马尾松快速诱导抗性及化学信号物质的研究[D].北京:北京林业大学博士论文, 2006
66张凤娟.光肩星天牛对槭树挥发物的响应及虫害诱导信号物质研究[D].北京:北京林业大学博士论文,2006
67 Rono E, Njagi PGN, Bashir MO, Hassanali A.Concentration-dependent parsimonious releaser roles of gregarious male pheromone of the desert locust, Schistocerca gregaria[J]. Journal of Insect Physiology, 2008, Vol.54:162-168
68 Assad YOH, Hassanali A, Torto B, Mahamat H, Bashir NHH, Bashir SE. Effects of fifth-instar volatiles on sexual maturation of adult desert locust Schistocerca gregaria[J]. Journal of Chemical Ecology, 1997, Vol.23 :1373-1388