B型和Q型烟粉虱竞争取代的取食行为学机制
|
摘要
烟粉虱是一种由多种生物型组成的世界性重要农业害虫，其中的B型和Q型是入侵性最强、危害最大的两种生物型。B型烟粉虱于20世纪90年代中后期入侵我国，Q型烟粉虱则于2003年最早于云南昆明发现。近些年来，烟粉虱在我国广大地区持续暴发危害，由其传播的番茄黄化曲叶病毒（TYLCV）病更是席卷我国主要番茄产区，给我国农业生产和国民经济造成了巨大损失。本论文利用EPG技术研究了寄主植物、病毒、内共生菌、化学农药等因子对B型和Q型烟粉虱取食行为的影响，相关研究结果对于阐明B型和Q型烟粉虱竞争取代机制以及TYLCV的流行机制具有重要意义。具体研究结果如下：
（1）B型和Q型烟粉虱在不同寄主植物上具有不同的取食优势。B型烟粉虱在甘蓝和黄瓜上的取食能力显著强于Q型，而Q型烟粉虱在番茄、棉花和一品红上的取食能力显著强于B型。B型和Q型烟粉虱取食行为的差异表明二者具有寄主适应性差异。
（2）TYLCV能以间接（植物感染）和直接（烟粉虱感染）2种方式修饰B型和Q型烟粉虱的取食行为。无论是B型还是Q型，TYLCV感染均能促进烟粉虱唾液分泌来促进病毒自身的快速传播。无论是植物感染还是烟粉虱感染TYLCV，Q型烟粉虱较B型具有适应性和传毒优势。
（3）在感染TYLCV的番茄苗上研究噻虫嗪对B型和Q型烟粉虱取食行为的影响发现，噻虫嗪处理后，B型烟粉虱刺吸起始时间提前，而Q型烟粉虱则表现了更强的连续性刺探、分泌了更多的唾液。可见噻虫嗪处理对Q型烟粉虱表现了更强的行为修饰作用，Q型烟粉虱对药剂处理表现了更强的适应性。烟粉虱的唾液分泌行为与TYLCV传播密切相关，显然噻虫嗪处理促进了Q型烟粉虱对TYLCV的传播。
（4）经抗生素处理和基因融合获得具有相同遗传背景，分别携带内共生菌Hamiltonella和不含Hamiltonella的两个Q型烟粉虱种群。两种群在黄瓜上的取食行为研究发现，与去除Hamiltonella的Q型烟粉虱相比，携带Hamiltonella的Q型烟粉虱能够较快、较容易的刺探到寄主植物韧皮部，并可显著提高烟粉虱唾液分泌频率。携带Hamiltonella提高Q型烟粉虱对寄主的适应性，同样Hamiltonella有可能通过修饰取食行为来促进TYLCV病毒的传播。
（5）将B型烟粉虱分别用甘蓝、番茄和黄瓜隔离饲养6年形成烟粉虱甘蓝种群、番茄种群和黄瓜种群。比较3个寄主种群B型烟粉虱分别在甘蓝、番茄和黄瓜上的取食行为发现，与番茄和黄瓜种群相比，甘蓝种群无论是在其原始寄主甘蓝上、还是在转移寄主番茄和黄瓜上均能较容易地刺探到韧皮部、分泌较多的唾液并具有较强的韧皮部取食能力。显然，长期在单一的寄主植物上取食可以对B型烟粉虱取食行为产生驯化作用，并且这种驯化作用强弱依寄主植物而异。
（6）对以上长期寄主隔离的甘蓝种群、番茄种群和黄瓜种群分子标记研究发现，寄主植物的长期隔离可以诱导B型烟粉虱种群发生明显的遗传分化，但是与Q型烟粉虱种群相比，B型烟粉虱不同寄主种群间的遗传距离均小于B和Q型烟粉虱生物型间的遗传距离。
综上所述，与B型烟粉虱相比，Q型烟粉虱具有较好的寄主适应性、较高的传毒效率、对化学农药适应性更强等行为优势，这可能是Q型烟粉虱取代B型烟粉虱的重要原因之一。
The whitefly Bemisia tabaci (Gennadius)(Hemiptera: Aleyrodidae) is a complex speciescomposed of at least24morphologically indistinguishable biotypes. Among them, biotypes B and Q areregarded as the two most invasive and destructive whiteflies. In China, B biotype was first detected inthe mid-1990s. Since then, B has gradually replaced the indigenous species, and has been the majorwhitefly pest till2007. The situation started to change in2003when Q arrived in Yunnan Province,China. In the following years, biotype Q has rapidly displaced B in most part of China and has becomethe most dominant whitefly. In recent years, B. tabaci causes excessive crop losses through directfeeding and transmission of plant viruses. For example, Tomato yellow leaf curl virus (TYLCV),transmitted by B. tabaci in a circulative manner, caused disease epidemics in many regions of China. Inthis study, we used the electrical penetration graph (EPG) toolkit to look at elements from both theinsect (feeding behavior) and plant (host suitability) perspective to better understand the factorsinvolved in the competitive displacement of B. tabaci biotype B by Q. The main findings are listed asfollows.
(1) Pair-wise comparisons of B. tabaci biotypes B and Q feeding on each of the five host plantsindicates that Q feeds better on tomato, cotton, and poinsettia, while B thrives on cabbage and cucumber.Our current results suggest that tomato, cotton and poinsettia are more suited for Q, while cabbage andcucumber are more suited for B. combined results from both insect and host plants suggest that hostsuitability and feeding behaviors displayed by adult whiteflies can affect their interaction and maycontribute to the rapid competitive displacement observed in China over the past6years.
(2) The indirect (plant infection) and direct (whitefly infection) effects of TYLCV on the feedingbehaviors of B. tabaci biotypes B and Q were evaluated using EPG technique. When non-viruliferouswhiteflies fed on healthy and TYLCV-infected plants, respectively, B. tabaci Q engaged in more phloemsalivation and phloem sap ingestion than B. Plant infection by virus significantly affected10of24feeding behavior parameters. When non-viruliferous and viruliferous whiteflies infested healthy plants,B. tabaci Q fed more readily than B on tomatoes, regardless of the infection status of whitefly. Whiteflyinfection significantly affected9of24feeding parameters. Viruliferous whiteflies engaged insignificantly more phloem salivation, which associated with inoculation of TYLCV. These combinedresults support the hypothesis that B. tabaci Q is a more efficient vector than B, and that whiteflyinfection by the virus directly modifies the vector’s feeding behavior in a manner which enhances viraltransmission.
(3) Pair-wise comparisons of the feeding behaviors between B. tabaci biotypes B and Q onthiamethoxam treated and untreated TYLCV-infected tomatoes were carried out. The results showedthat biotype B probed more rapidly, while biotype Q fed more continuously and secreted more saliva on thiamethoxam treated tomatoes than on control ones. This indicated that thiamethoxam brought aboutgreater modification on the feeding behaviors for biotype Q than B, and biotype Q has more suitabilityto thiamethoxam treatment. The whitefly salivation is closely associated with TYLCV transmission.Thus the thiamethoxam treatment improved the TYLCV transmission in this study.
(4) To evaluate the effects of endosymbiont, Hamiltonella, on feeding behaviors of B. tabaci, twoB. tabaci biotype Q colonies with the same genetic background, Hamiltonella-infeced (H+) anduninfected (H-), were established by antibiotic treatments and introgression. Comparisons of the feedingbehaviors on cucumber between the two colonies revealed that the Hamiltonella-infeced biotype Q(H+Q) reach the phloem phase more rapidly and easily than Hamiltonella-uninfeced biotype Q (H-Q).Obviously, the infection of Hamiltonella improved the fitness of whitefly. Meanwhile, the H+Q havehigher salivation frequency than H-Q, which indicated that the Hamiltonella is likely to improve theTYLCV transmission.
(5) Feeding behavioral characters of long-term host isolated B. tabaci were examined. Three hostpopulations of B. tabaci biotype B with the same genetic background were established through rearingthe whiteflies segregated on cabbage, tomato, and cucumber for6years (hereafter referred to asCa-population, To-population, and Cu-population, respectively). Feeding behavior comparisons amongthe three host populations were carried out on each of the cabbage, tomato and cucumber plant,respectively. The results showed that Ca-population reached to the phloem phase more readily, secretemore saliva, and feed better than the other two host populations on any of the3host plants. Evidently,the feeding behaviors of B biotype were domesticated by adaptation for a long time on a specific hostplant, and the domestication degree varied with the host plants species.
(6) Understanding the feeding behavior differences of the host specific B. tabaci biotype Bpopulations, the genetic differentiations of the3B biotype populations above and1additional Q biotypepopulation were further studied through AFLP and ISSR markers. The results indicated that thelong-term isolation of host plants induced a significant genetic differentiation for B biotype. However,the genetic distances among the three B biotype host populations were all lower than that between Band Q biotypes.
In conclusion, B. tabaci biotype Q exhibited stronger host suitability, higher virus transmissionefficiency, and more adaption to the pesticide than B biotype. These factors may contribute to thecompetitive displacement of biotype B by Q.
（1）B型和Q型烟粉虱在不同寄主植物上具有不同的取食优势。B型烟粉虱在甘蓝和黄瓜上的取食能力显著强于Q型，而Q型烟粉虱在番茄、棉花和一品红上的取食能力显著强于B型。B型和Q型烟粉虱取食行为的差异表明二者具有寄主适应性差异。
（2）TYLCV能以间接（植物感染）和直接（烟粉虱感染）2种方式修饰B型和Q型烟粉虱的取食行为。无论是B型还是Q型，TYLCV感染均能促进烟粉虱唾液分泌来促进病毒自身的快速传播。无论是植物感染还是烟粉虱感染TYLCV，Q型烟粉虱较B型具有适应性和传毒优势。
（3）在感染TYLCV的番茄苗上研究噻虫嗪对B型和Q型烟粉虱取食行为的影响发现，噻虫嗪处理后，B型烟粉虱刺吸起始时间提前，而Q型烟粉虱则表现了更强的连续性刺探、分泌了更多的唾液。可见噻虫嗪处理对Q型烟粉虱表现了更强的行为修饰作用，Q型烟粉虱对药剂处理表现了更强的适应性。烟粉虱的唾液分泌行为与TYLCV传播密切相关，显然噻虫嗪处理促进了Q型烟粉虱对TYLCV的传播。
（4）经抗生素处理和基因融合获得具有相同遗传背景，分别携带内共生菌Hamiltonella和不含Hamiltonella的两个Q型烟粉虱种群。两种群在黄瓜上的取食行为研究发现，与去除Hamiltonella的Q型烟粉虱相比，携带Hamiltonella的Q型烟粉虱能够较快、较容易的刺探到寄主植物韧皮部，并可显著提高烟粉虱唾液分泌频率。携带Hamiltonella提高Q型烟粉虱对寄主的适应性，同样Hamiltonella有可能通过修饰取食行为来促进TYLCV病毒的传播。
（5）将B型烟粉虱分别用甘蓝、番茄和黄瓜隔离饲养6年形成烟粉虱甘蓝种群、番茄种群和黄瓜种群。比较3个寄主种群B型烟粉虱分别在甘蓝、番茄和黄瓜上的取食行为发现，与番茄和黄瓜种群相比，甘蓝种群无论是在其原始寄主甘蓝上、还是在转移寄主番茄和黄瓜上均能较容易地刺探到韧皮部、分泌较多的唾液并具有较强的韧皮部取食能力。显然，长期在单一的寄主植物上取食可以对B型烟粉虱取食行为产生驯化作用，并且这种驯化作用强弱依寄主植物而异。
（6）对以上长期寄主隔离的甘蓝种群、番茄种群和黄瓜种群分子标记研究发现，寄主植物的长期隔离可以诱导B型烟粉虱种群发生明显的遗传分化，但是与Q型烟粉虱种群相比，B型烟粉虱不同寄主种群间的遗传距离均小于B和Q型烟粉虱生物型间的遗传距离。
综上所述，与B型烟粉虱相比，Q型烟粉虱具有较好的寄主适应性、较高的传毒效率、对化学农药适应性更强等行为优势，这可能是Q型烟粉虱取代B型烟粉虱的重要原因之一。
The whitefly Bemisia tabaci (Gennadius)(Hemiptera: Aleyrodidae) is a complex speciescomposed of at least24morphologically indistinguishable biotypes. Among them, biotypes B and Q areregarded as the two most invasive and destructive whiteflies. In China, B biotype was first detected inthe mid-1990s. Since then, B has gradually replaced the indigenous species, and has been the majorwhitefly pest till2007. The situation started to change in2003when Q arrived in Yunnan Province,China. In the following years, biotype Q has rapidly displaced B in most part of China and has becomethe most dominant whitefly. In recent years, B. tabaci causes excessive crop losses through directfeeding and transmission of plant viruses. For example, Tomato yellow leaf curl virus (TYLCV),transmitted by B. tabaci in a circulative manner, caused disease epidemics in many regions of China. Inthis study, we used the electrical penetration graph (EPG) toolkit to look at elements from both theinsect (feeding behavior) and plant (host suitability) perspective to better understand the factorsinvolved in the competitive displacement of B. tabaci biotype B by Q. The main findings are listed asfollows.
(1) Pair-wise comparisons of B. tabaci biotypes B and Q feeding on each of the five host plantsindicates that Q feeds better on tomato, cotton, and poinsettia, while B thrives on cabbage and cucumber.Our current results suggest that tomato, cotton and poinsettia are more suited for Q, while cabbage andcucumber are more suited for B. combined results from both insect and host plants suggest that hostsuitability and feeding behaviors displayed by adult whiteflies can affect their interaction and maycontribute to the rapid competitive displacement observed in China over the past6years.
(2) The indirect (plant infection) and direct (whitefly infection) effects of TYLCV on the feedingbehaviors of B. tabaci biotypes B and Q were evaluated using EPG technique. When non-viruliferouswhiteflies fed on healthy and TYLCV-infected plants, respectively, B. tabaci Q engaged in more phloemsalivation and phloem sap ingestion than B. Plant infection by virus significantly affected10of24feeding behavior parameters. When non-viruliferous and viruliferous whiteflies infested healthy plants,B. tabaci Q fed more readily than B on tomatoes, regardless of the infection status of whitefly. Whiteflyinfection significantly affected9of24feeding parameters. Viruliferous whiteflies engaged insignificantly more phloem salivation, which associated with inoculation of TYLCV. These combinedresults support the hypothesis that B. tabaci Q is a more efficient vector than B, and that whiteflyinfection by the virus directly modifies the vector’s feeding behavior in a manner which enhances viraltransmission.
(3) Pair-wise comparisons of the feeding behaviors between B. tabaci biotypes B and Q onthiamethoxam treated and untreated TYLCV-infected tomatoes were carried out. The results showedthat biotype B probed more rapidly, while biotype Q fed more continuously and secreted more saliva on thiamethoxam treated tomatoes than on control ones. This indicated that thiamethoxam brought aboutgreater modification on the feeding behaviors for biotype Q than B, and biotype Q has more suitabilityto thiamethoxam treatment. The whitefly salivation is closely associated with TYLCV transmission.Thus the thiamethoxam treatment improved the TYLCV transmission in this study.
(4) To evaluate the effects of endosymbiont, Hamiltonella, on feeding behaviors of B. tabaci, twoB. tabaci biotype Q colonies with the same genetic background, Hamiltonella-infeced (H+) anduninfected (H-), were established by antibiotic treatments and introgression. Comparisons of the feedingbehaviors on cucumber between the two colonies revealed that the Hamiltonella-infeced biotype Q(H+Q) reach the phloem phase more rapidly and easily than Hamiltonella-uninfeced biotype Q (H-Q).Obviously, the infection of Hamiltonella improved the fitness of whitefly. Meanwhile, the H+Q havehigher salivation frequency than H-Q, which indicated that the Hamiltonella is likely to improve theTYLCV transmission.
(5) Feeding behavioral characters of long-term host isolated B. tabaci were examined. Three hostpopulations of B. tabaci biotype B with the same genetic background were established through rearingthe whiteflies segregated on cabbage, tomato, and cucumber for6years (hereafter referred to asCa-population, To-population, and Cu-population, respectively). Feeding behavior comparisons amongthe three host populations were carried out on each of the cabbage, tomato and cucumber plant,respectively. The results showed that Ca-population reached to the phloem phase more readily, secretemore saliva, and feed better than the other two host populations on any of the3host plants. Evidently,the feeding behaviors of B biotype were domesticated by adaptation for a long time on a specific hostplant, and the domestication degree varied with the host plants species.
(6) Understanding the feeding behavior differences of the host specific B. tabaci biotype Bpopulations, the genetic differentiations of the3B biotype populations above and1additional Q biotypepopulation were further studied through AFLP and ISSR markers. The results indicated that thelong-term isolation of host plants induced a significant genetic differentiation for B biotype. However,the genetic distances among the three B biotype host populations were all lower than that between Band Q biotypes.
In conclusion, B. tabaci biotype Q exhibited stronger host suitability, higher virus transmissionefficiency, and more adaption to the pesticide than B biotype. These factors may contribute to thecompetitive displacement of biotype B by Q.
引文
1.安瑞生,谭声江,陈晓峰.微卫星DNA在分子遗传标记研究中的应用.昆虫知识,2002,39(3):165~172.
2.褚栋,陈国发,徐宝云,吴青君,张友军.烟粉虱B型和Q型群体遗传结构的RAPD分析.昆虫学报,2007,50(3):264~270.
3.罗晨,姚远,王戎疆,阎凤鸣,胡敦孝,张芝利.利用mtDNA COI基因序列鉴定我国烟粉虱的生物型.昆虫学报,2002,45(6):759~763.
4.罗晨,张芝利.烟粉虱Bemisia tabaci (Gennadius)研究概述.北京农业科学,2000,18（增刊）:4~13.
5.张芝利.关于烟粉虱大发生的思考.北京农业科学（烟粉虱专辑）,2000:1~3.
6.周尧.中国粉虱名录.中国昆虫学,1949,3(4):1~18.
7. Alvarez AE, Garzo E, Verbeek M, Vosman B, Dicke M, Tjallingii WF. Infection of potato plantswith Potato Leafroll Virus changes attraction and feeding behaviour of Myzus persicae.Entomologia Experimentalis et Applicata2007,125:135~144.
8. Alvarez AE, Tjallingii WF, Garzo E, Vleeshouwers V, Dicke M, Vosman B. Location of resistancefactors in the leaves of potato and wild tuber-bearing Solanum species to the aphid Myzus persicae.Entomologia Experimentalis et Applicata2006,121(2):145~157.
9. Annan IB, Schaefers GA, Tingey WM, Tjallingii WF. Effects of treatments for electricalpenetration graph recordings on behaviour and biology of Aphis craccivora (Aphididae).Physiological Entomology1997,22(2):95~101.
10. Backus EA, Bennett WH. The AC-DC Correlation Monitor: New EPG design with flexible inputresistors to detect both R and emf components for any piercing-sucking hemipteran. Journal ofInsect Physilology2009,55(10):869~884.
11. Backus EA, Serrano MS, Ranger CM. Mechanisms of hopperburn: An overview of insecttaxonomy, behavior, and physiology. Annual Review of Entomology2005,50:125~151.
12. Backus EA. Sensory systems and behaviours which mediate hemipteran plant-feeding: ataxonomic overview. Journal of Insect Physiology1988,34:151~165.
13. Baumann P. Biology bacteriocyte-associated endosymbionts of plant sap-sucking insects. AnnualReview of Microbiology2005,59:155~189.
14. Bedford ID, Briddon RW, Brown JK, Rosell RC, Markham PG. Geminivirus transmission andbiological characterization of Bemisia tabaci (Gennadius) from different geographic regions.Annals of Applied Biology1994,125:311~325.
15. Bedford ID, Briddon RW, Markham PG, Brown JK, Rosell RC. A new species of Bemisia orbiotype of Bemisia tabaci (Genn.), as a future pest of European agriculture.(in) Plant Health andEuropean Single Market, D. Ebbles (eds). BCPC Monograph1993,54:381~386.
16. Berry SD, FondongVN, Rey C, Rogan D, Fauquet CM, Brown JK. Molecular evidence for fivedistinct Bemisia tabaci (Homoptera: Aleyrodidae) geographic haplotypes associated with cassavaplants in Sub-Saharan Africa. Annals of the Entomological Society of America2004,97:852~859.
17. Bird J. A whitefly-transmitted mosaic of Jatropha gossypifolia. Technical Paper, AgriculturalExperimental Station, University of Puerto Rico1957,22:1~35.
18. Blackmer JL, Byrne DN. Environmental and physiological factors influencing phototactic flight ofBemisia tabaci. Physiology Entomology1993b,18:336~342.
19. Blackmer JL, Byrne DN. Flight behaviour of Bemisia tabaci in a vertical flight chamber: effect oftime of day, sex, age and host quality. Physiology Entomology1993a,18:223~232.
20. Bonani JP, Fereres A, Garzo E, Miranda MP, Appezzato-Da-Gloria B, Lopes JRS. Characterizationof electrical penetration graphs of the Asian citrus psyllid, Diaphorina citri, in sweet orangeseedlings. Entomologia Experimentalis et Applicata2010,134:35~49.
21. Boykin LM, Shatters RG, Rosell RC, McKenzie CL, Bagnall RA, De Barro PJ, Frohlich DR.Global relationships of Bemisia tabaci (Hemiptera: Aleyrodidae) revealed using Bayesian analysisof mitochondrial COI DNA sequences. Molecular Phylogenetics and Evolution2007,44:1306~1319.
22. Bradley RHE, Sylvester ES, Wade CV. Note on the movements of the mandibular and maxillarystylets of the aphid, Myzus persicae (Sulzer). The Canadian Entomologist1962,94:653~654.
23. Briddon RW, Pinner MS, Stanley J, Markham PG. Geminivirus coat protein gene replacementalters insect specificity. Virology1990,177:85~94.
24. Brown CM, Holbrook FR. An improved electronic system for monitoring feeding of Aphids.American Journal of Potato Research1976,53:457~462.
25. Brown JK, Bird J. Variability within the Bemisia tabaci species complex and its relation to newepidemics caused by geminiviruses. CEIBA1995,36:73~80.
26. Brown JK, Coats SA, Bedford ID, Markham PG, Bird J. Biotypic characterization of Bemisiatabaci populations based on esterase profiles, DNA fingerprinting, virus transmission, and bioassayto key host plant species. Phytopathology1992,82:1104.
27. Brown JK, Frohlich DR, Rosell RC. The sweetpotato or silverleaf whiteflies: biotypes of Bemisiatabaci or a species complex? Annual Review of Entomology1995,40:511-534.
28. Brown JK. Current status of Bemisia tabaci as a plant pest and virus vector in agro-ecosystemsworldwide. Bulletin Phytosanitaire de la FAO1994,42:3~32.
29. Brown JK. Molecular markers for the identification and global tracking of whitefly vector-Begomovirus complexes. Virus Research2000,71:233~260.
30. Brown JK. Phylogenetic biology of the Bemisia tabaci sibling species group.(in) PA Stansly andSE Naranjo (eds). Bionomics and Management of a Global Pest. Springer, Amsterdam2010,31~67.
31. Brown JK. The Bemisia tabaci complex: genetic and phenotypic variation and relevance toTYLCV-vector interactions.(in) Czosnek H (eds). Tomato yellow leaf curl virus disease:management, molecular biology, breeding for resistance. Springer, Dordrecht2007,25~56.
32. Burban C, Fishpool LDC, Fauquet C, Fargette D, Thouvenel JC. Host-associated biotypes withinWest African populations of the whitefly Bemisia tabaci (Genn.),(Hom., Aleyrodidae). Journal ofApplied Entomology1992,113:416~423.
33. Butler CG. On the ecology of Aleurodes brassicae Walk.(Hemiptera). Transactions of the RoyalEntomological Society of London1938,87:291~311.
34. Calatayud PA, Rahb Y, Tjallingii WF, Tertuliano M, Le Rü B. Electrically recorded feedingbehaviour of cassava mealybug on host and non-host plants. Entomologia experimentalis etapplicata1994,72:219~232.
35. Campbell BC, Jones KC, Dreyer DL. Discriminative behavioral responses by aphids to variousplant matrix Polysaccharides. Entomologia Experimentalis et Applicata1986,41(1):17~24.
36. Casta eda LE, Sandrock C, Vorburger C. Variation and covariation of life history traits in aphidsare related to infection with the facultative bacterial endosymbiont Hamiltonella defensa.Biological Journal of the Linnean Society2010,100(1):273~247.
37. Cervera MT, Cabezas JA, Simón B, Martínez-Zapater JM, Beitia F, Cenis JL. Genetic relationshipsamong biotypes of Bemisia tabaci (Hemiptera: Aleyrodidae) based on AFLP analysis. Bulletin ofEntomological Research2000,90:391~396.
38. Cherqui A, Tjallingii WF. Salivary proteins of aphids, a pilot study on identification, separation andimmunolocalisation. Journal of Insect Physiology2000,46:1177~1186.
39. Chiel E, Gottlieb Y, Zchori-Fein E, Mozes-Daube N, Katzir N, Inbar M, Ghanim M.Biotype-dependent secondary symbiont communities in sympatric populations of Bemisia tabaci.Bulletin of Entomological Research2007,97:407~413.
40. Chiel E, Inbar M, Mozes-Daube N, White JA, Hunter MS, Zchori-Fein E. Assessments of FitnessEffects by the Facultative Symbiont Rickettsia in the Sweetpotato Whitefly (Hemiptera:Aleyrodidae). Annals of the Entomological Society of America2009,102(3):413~418.
41. Chu D, Gao CS, De Barro P, Zhang YJ, Wan FH, Khan IA. Further insights into the strange role ofbacterial endosymbionts in whitefly, Bemisia tabaci: comparison of secondary symbionts frombiotypes B and Q in China. Bulletin of Entomological Research2011,101(4):477~486.
42. Chu D, Gao CS, De Barro PJ, Wan FH, Zhang YJ. Investigation of the genetic diversity of aninvasive whitefly (Bemisia tabaci) in China using both mitochondrial and nuclear DNA markers.Bulletin of Entomological Research2011a,101(4):467~475.
43. Chu D, Wan FH, Xu BY, Wu QJ, Zhang YJ. Comparative analysis of population genetic structurein Bemisia tabaci (Gennadius) biotypes B and Q based on ISSR marker. Agricultural Sciences inChina2008,7(11):1348~1354.
44. Chu D, Wan FH, Zhang YJ, Brown JK. Change in the biotype composition of Bemisia tabaci inShandong province of China from2005to2008. Environmental Entomology2010a,39(3):1028~1036.
45. Chu D, Zhang YJ, Brown JK, Cong B, Xu BY, Wu QJ, Zhu GR. The introduction of the exotic Qbiotype of Bemisia tabaci (Gennadius) from the Mediterranean region into China on ornamentalcrops. Florida Entomologist2006,89(2):168~174.
46. Chu D, Zhang YJ, Wan FH. Cryptic invasion of the exotic Bemisia tabaci biotype Q occurredwidespread in Shandong Province of China. Florida Entomologist2010b,93:203~207.
47. Cohen AC, Hendrix DL. Demonstration and preliminary characterization of α-amylase in thesweetpotato whitefly, Bemisia tabaci (Aleyrodidae: Homoptera). Comparative Biochemistry andPhysiology Part B1994,109:593~601.
48. Cohen AC, Henneberry TJ, Chu CC. Geometric relationships between whitefly feeding behaviorand vascular bundle arrangements. Entomologia Experimentalis et Applicata1996,78:135~142.
49. Cohen S, Lapidot M. Appearance and expansion of TYLCV: a historical point of view.(in) HCzosnek (eds). Tomato Yellow Leaf Curl Virus Disease. Springer,2007,3~12.
50. Costa HS, Brown JK. Variation in biological characteristics and esterase patterns amongpopulations of Bemisia tabaci, and the association of one population with silverleaf symptominduction. Entomologia Experimentalis et Applicata1991,61:211~219.
51. Crowder DW, Ellers-Kirk C, Yafuso C, Dennehy TJ, Degain BA, Harpold VS, Tabashnik BE,Carrière Y. Inheritance of resistance to pyriproxyfen in Bemisia tabaci (Hemiptera: Aleyrodidae)males and females (B biotype). Journal of Economic Entomology2008,101:927~932.
52. Crowder DW, Horowitz AR, De Barro PJ, Liu SS, Showalter AW, Kontsedalov S, Khasdan V,Shargal A, Liu J, Carrière Y. Mating behaviour, life history and adaptation to insecticides determinespecies exclusion between whiteflies. Journal of Animal Ecology2010a,79:563~570.
53. Crowder DW, Sitvarin MI, Carrière Y. Plasticity in mating behaviour drives asymmetricreproductive interference in whiteflies. Animal Behaviour2010b,79:579~587.
54. Czosnek H, Ghanim M. Bemisia tabaci-Tomato Yellow Leaf Curl Virus interaction causingworldwide epidemics.(in) WMO Thompson (eds). the whitefly, Bemisia tabaci (Homoptera:Aleyrodidae) interaction with Geminivirus-infected host plants. Springer2011,51~67.
55. De Barro PJ, Driver F. Use of RAPD PCR to distinguish the B biotype from other biotypes ofBemisia tabaci (Gennadius)(Hemiptera, Aleyrodidae). Australian Journal of Entomology1997,36:149~152.
56. De Barro PJ, Liu SS, Boykin LM, Dinsdale A. Bemisia tabaci: a statement of species status.Annual Review Entomology2011,56:1~19.
57. De Barro PJ. Genetic structure of the whitefly Bemisia tabaci in the Asia–Pacific region revealedusing microsatellite markers. Molecular Ecology2005,14:3695~3718.
58. Dinsdale A, Cook L, Riginos C, Buckley YM, De Barro PJ. Refined global analysis of Bemisiatabaci (Hemiptera: Sternorrhyncha: Aleyrodoidea: Aleyrodidae) mitochondrial cytochrome oxidase1to identify species level genetic boundaries. Annals of the Entomological Society of America2010,103:196~208.
59. Dreyer DL, Campbell BC. Chemical basis of host-plant resistance to aphids. Plant CellEnvironment1987,10:353~361.
60. Dugravot S, Backus EA, Reardon BJ, Miller TA. Correlations of cibarial muscle activities ofHomalodisca spp. Sharpshooters (Hemiptera: Cicadellidae) with EPG ingestion waveform andexcretion. Journal of Insect Physiology2008,54:1467~1478.
61. Fauquet CM, Maxwell DP, Gronenborn B, Stanley J. Revised proposal for naming geminiviruses.Archives of Virology2000,145:1743~1761.
62. Fereres A, Moreno A. Behavioural aspects influencing plant virus transmission by homopteraninsects. Virus Research2009,141:158~168.
63. Freeman TP, Buckner JS, Nelson DR, Chu CC, Henneberry TJ. Stylet penetration by Bemisiaargentifolii (Homoptera: Aleyrodidae) into host leaf tissue. Annals of the Entomological Society ofAmerica2001,94:761~768.
64. Frohlich DR, Torres-Jerez I, Bedford ID, Markham PG, Brown JK. A phylogeographical analysis ofthe Bemisia tabaci species complex based on mitochondrial DNA markers. Molecular Ecology1999,8:1683~1691.
65. Funk CJ. Alkaline phosphatase activity in whitefly salivary glands and saliva. Archieves of InsectBiochemistry and Physiology2001,46:165~174.
66. Gabrys B, Tjallingii WF, van Beek TA. Analysis of EPG recorded probing by cabbage aphid onhost plant parts with different glucosinolate contents. Journal of Chemical Ecology1997,23(7):1661~1673.
67. Garzo E, Soria C, Gómez-Guillamón ML, Alberto F. Feeding behavior of Aphis gossypii onresistant accessions of different melon genotypes (Cucumis melo). Phytoparasitica2002,30(2):129~140.
68. Gelman DB, Blackburn MB, Hu JS, Gerling D. The nymphal-adult molt of the silverleaf whitefly(Bemisia argentifolii): timing, regulation, and progress. Archieves of Insect Biochemistry andPhysiology2002,51:67~79.
69. Gennadius P. Disease of tobacco plantations in Trikonia: the aleurodid of tobacco. EllennikeGeorgia1889,6(5): l~3.
70. Ghanim M, Rosell RC, Campbell LR, Czosnek H, Brown JK, Ullman DE. Digestive, salivary andreproductive organs of Bemisia tabaci (Gennadius)(Hemiptera: Aleyrodidae) type B. Journal ofMorphology2001,248:22~40.
71. Ghanim M, Sobol I, Ghanim M, Czosnek H. Horizontal transmission of begomoviruses betweenBemisia tabaci biotypes. Arthropod-Plant Interaction2007,1(3):195~204.
72. Gill RJ. A review of the sweetpotato whitefly in Southern California. The Pan-Pacific Entomologist1992,68:144~152.
73. Gottieb Y, Zchori-Fein E, Mozes-Daube N, Kontsedalov S, Skaljac M, Brumin I, Czosnek H,Vavre F, Fleury F, Ghanim M. The Transmission efficiency of Tomato Yellow Leaf Curl Virus bythe whitefly Bemisia tabaci is correlated with the presence of a specific symbiotic bacteriumspecies. Journal of Virology2010,84(18):9310~9317.
74. Gottlieb Y, Ghanim M, Chiel Eald, Gerling D, Portnoy V, Steinberg S, Tzuri G, Horowitz AR,Belausov E, Mozes-Daube N, Kontsedalov S, Gershon M, Shunit G, Katzir N, Zchori-Fein E.Identification and localization of a Rickettsia sp. in Bemisia tabaci (Homoptera: Aleyrodidae).Applied and Environmental Microbiology2006,72:3646~3652.
75. Gottlieb Y, Ghanim M, Gueguen G, Kontsedalov S, Vavre F, Fleury F, Zchori-Fein E. Inheritedintracellular ecosystem: symbiotic bacteria share bacteriocytes in whiteflies. The FASEB Journal2008,22:2591~2599.
76. Gueguen G, Vavre F, Gnankine O, Peterschmitt M, Charif D, Chiel E, Gottlieb Y, Ghanim M,Zchori-Fein E, Fleury F. Endosymbiont metacommunities, mtDNA diversity and the evolution ofthe Bemisia tabaci (Hemiptera: Aleyrodidae) species complex. Molecular Ecology2010,19(19):4365~4376.
77. Guershon M, Gerling D. Effect of foliar tomentosity on phenotypic plasticity in Bemisia tabaci(Hom., Aleyrodidae). Journal of Applied Entomology2001,125:449~453.
78. Guershon M, Gerling D. Predatory behavior of Delphastus pusillus in relation to the phenotypicplasticity of Bemisia tabaci nymphs. Entomologia Experimentalis et Applicata1999,92:239~248.
79. Guirao P, Beitia F, Cenis JL. Biotype determination of Spanish populations of Bemisia tabaci(Hemiptera: Aleyrodidae). Bulletin of Entomological Research1997,87:587~593.
80. Harrewijn P, Piron PGM, Mollema C. Electrically recorded probing behaviour of thrips species onoptimal and suboptimal hosts. Entomologia Experimentalis et Applicata,1996,80(1):43~45.
81. Harris KF, Esbroeck ZP-V, Duffus JE. Morphology of the sweet potato whitefly, Bemisia tabaci(Homoptera, Aleyrodidae) relative to virus transmission. Zoomorphology1996,116:143~156.
82. Harrison BD, Robinson DJ. Natural genomic and antigenic variation in whitefly transmittedgeminiviruses (Begomoviruses). Annual Review of Phytopathology1999,37:369~398.
83. Hedges LM, Brownlie JC, O'Neill SL, Johnson KN. Wolbachia and virus protection in insects.Science2008,322(5902):702.
84. Helmi A. Host-associated population variations of Bemisia tabaci (Genn.)(Hemiptera:Sternorrhyncha: Aleyrodidae) characterized with random DNA markers. International Journal ofZoological Research2011,7(1):77~84.
85. Hilker M, Meiners T. Early herbivore alert: insect eggs induce plant defense. Journal of ChemicalEcology2006,32(7):1379~1397.
86. Himler AG, Adachi-Hagimori T, Bergen JE, Kozuch A, Kelly SE, Tabashnik BE, Chiel E,Duckworth VE, Dennehy TJ, Zchori-Fein E, Hunter MS. Rapid spread of a bacterial symbiont inan Invasive Whitefly is driven by fitness benefits and female bias. Science2011,332(6026):254~256.
87. Horowitz, AR, Kontsedalov S, Khasdan V, Ishaaya I. Biotypes B and Q of Bemisia tabaci and theirrelevance to neonicotinoid and pyriproxyfen resistance. Archives of Insect Biochemistry andPhysiology2005,58:216~225.
88. Hu J, De Barro P, Zhao H, Wang J, Nardi F, Liu SS. An extensive field survey combined with aphylogenetic analysis reveals rapid and widespread invasion of two alien whiteflies in China. PLoSONE2011,6(1): e16061.
89. Jaenike J, Papaj DR. Behavioural plasticity and pattern of host use by insect.(in) Roitberg BD, andIsman MB (eds). Insect chemical ecology, an evolutionary approach, Chapman and Hall. New York,1992,245~264.
90. Janssen JAM, Tjallingii WF, van Lenteren JC. Electrical recording and ultrastructure of styletpenetration by the greenhouse whitefly. Entomologia Experimentalis et Applicata1989,52:69~81.
91. Ji YH, Xiong RY, Cheng ZB, Zhou T, Zhao TM, Yu WG, Fan YJ, Zhou YJ. Molecular diagnosis ofTomato yellow leaf curl disease in Jiangsu Province. Acta Horticulturae Sinica2008,35:1815~1818.
92. Jiang YX, Blas C, Barrios L, Fereres A. Correlation between whitefly (Homoptera: Aleyrodidae)feeding behavior and transmission of tomato yellow leaf curl virus. Annals of EntomologicalSociety of America2000,93:573~579.
93. Jiang YX, Lei H, Collar JL, Martin B, Muniz M, Fereres A. Probing and feeding behavior of twodistinct biotypes of Bemisia tabaci (Homoptera: Aleyrodidae) on tomato plants. Journal ofEconomic Entomology1999,92:357~366.
94. Jiang YX, Nombela G, Muniz M. Analysis by DC-EPG of the resistance to Bemisia tabaci on anMi-tomato line. Entomologia Experimentalis et Applicata2001,99:295~302.
95. Jiang YX, Walker GP. Identification of phloem sieve elements as the site of resistance to silverleafwhitefly in resistant alfalfa genotypes. Entomologia Experimentalis et Applicata2007,125:307~320.
96. Jiang YX, Walker GP. Pathway phase waveform characteristics correlated with length and rate ofstylet advancement and partial stylet withdrawal in AC electrical penetration graphs of adultwhiteflies. Entomologia Experimentalis et Applicata2001,101:233~246.
97. Johnson DD, Walker GP, Creamer R. Stylet penetration behavior resulting in inoculation of asemipersistently transmitted closterovirus by the whitefly Bemisia argentifolii. EntomologiaExperimentalis et Applicata2002,102:115~123.
98. Johnson DD, Walker GP. Intracellular punctures by the adult whitefly Bemisia argentifolii on DCand AC electronic feeding monitors. Entomologia Experimentalis et Applicata1999,92:257~270.
99. Jones DR. Plant viruses transmitted by whiteflies. European Journal of Plant Pathology2003,109:195~219.
100. Joosta PH, Backus EA, Morgan D, Yan FM. Correlation of stylet activities by the glassy-wingedsharpshooter, Homalodisca coagulata (Say), with electrical penetration graph (EPG) waveforms.Journal of Insect Physiology2006,52:327~337.
101. Kimmins FM, Tjallingii WF. Ultrastucture of sieve element penetration by aphid stylets duringelectrical recording. Entomologia Experimentalis et Applicata1985,39:135~141.
102. Kindt F, Joosten NN, Tjallingii WF. Electrical penetration graphs of thrips revised: combining DC-and AC-EPG signals. Journal of Insect Physilology2006,52(1):1~10.
103. Knoblauch M, van Bel AJE. Sieve tubes in action. Plant Cell1998,10:35~50.
104. Lazarowitz SG, Shepherd DRJ. Geminiviruses: genome structure and gene function. CriticalReviews in Plant Sciences1992,11(4):327~349.
105. Lei H, Tjallingii WF, van Lenteren JC, Xu RM. Stylet penetration by larvae of the greenhousewhitefly on cucumber. Entomologia Experimentalis et Applicata1996,79:77~84.
106. Lei H, Tjallingii WF, van Lenteren JC. Probing and feeding characteristics of the greenhousewhitefly in association with host-plant acceptance and whitefly strains. EntomologiaExperimentalis et Applicata1998,88:73~80.
107. Lei H, Tjallingii WF, van Lenteren JC. Effect of tethering during EPG recorded probing by adultsof the greenhouse whitefly. Journal of Applied Entomology1997,121:211~217.
108. Lei H, van Lenteren JC, Tjallingii WF. Analysis of resistance in tomato and sweet pepper againstthe greenhouse whitefly using electrically monitored and visually observed probing and feedingbehaviour. Entomologia Experimentalis et Applicata1999,92:299~309.
109. Lei H, van Lenteren JC, Xu RM. Effects of plant tissue factors on the acceptance of fourgreenhouse vegetable host plants by the greenhouse whitefly: an electrical penetration graph (EPG)study. European Journal of Entomolomogy2001,98:31~36.
110. Leopolda RA, Freemanb TP, Bucknera JS, Nelsona DR. Mouthpart morphology and styletpenetration of host plants by the glassywinged sharpshooter, Homalodisca coagulata,(Homoptera:Cicadellidae). Arthropod Structure&Development2003,32:189~199.
111. Leszczynski B, Tjallingii WF, Dixon AFG, Swiderski R. Effect of methoxyphenols on grain aphidfeeding behavior. Enwmologia Experimentalis etApplicata1995,76:157~162.
112. Liu SS, De Barro PJ, Xu J, Luan JB, Zang LS, Ruan YM, Wan FH. Asymmetric matinginteractions drive widespread invasion and displacement in a whitefly. Science2007,318:1769~1772.
113. Luan JB, Li JM, Varela N, Wang YL, Li FF, Bao YY, Zhang CX, Liu SS, Wang XW. Globalanalysis of the transcriptional response of whitefly to Tomato yellow leaf curl China virus revealsthe relationship of coevolved adaptations. Journal of Virology2011,85:3330~3340.
114. Marchetti E, Civolani S, Leis M, Chicca M, Tjallingii WF, Pasqualini E, Baronio P. Tissue locationof resistance in apple to the rosy apple aphid established by electrical penetration graphs. Bulletinof Insectology2009,62(2):203~208.
115. Martín B, Collar JL, Tjallingii WF, Fereres A. Intracellular ingestion and salivation by aphids maycause the acquisition and inoculation of non-persistently transmitted plant viruses Journal ofGeneral Virology1997,78:2701~2705.
116. Mayoral AM, Tjallingii WF, Casta era P. Probing behaviour of Diuraphis noxia on five cerealspecies with different hydroxamic acid levels. Entomologia Experimentalis et Applicata1996,78:341~348.
117. McKenzie CL, Hodges G, Osborne LS, Byrne FJ, Shatters Jr. RG. Distribution of Bemisia tabaci(Hemiptera: Aleyrodidae) biotypes in Florida-investigating the Q invasion. Journal of EconomicEntomology2009,102:670~676.
118. McLean DL, Kinsey MG. A technique for electronically recording aphid feeding and salivation.Nature1964,202:1358~1359.
119. McLean DL, Kinsey MG. Identification of electrically recorded curve patterns associated withaphid salivation and ingestion. Nature1965,205(4976):1130~1131.
120. McLean DL, Kinsey MG. Probing behavior of the pea aphid, Acyrthosiphon pisum. IV. Effects ofstarvation on certain probing activities.1969,62(5):987~994.
121. McLean DL, Kinsey MG. Probing behavior of the pea aphid, Acyrthosiphon pisum. I. Definitivecorrelation of electronically recorded waveforms with aphid probing activities.1967,(2):400~405.
122. McLean DL, Kinsey MG. Probing behavior of the pea aphid, Acyrthosiphon pisum. II.Comparisons of salivation and ingestion in host and non-host plant leaves.1968a,61(3):730~739.
123. McLean DL, Kinsey MG. Probing behavior of the pea aphid, Acyrthosiphon pisum. III. Effect oftemperature differences on certain probing activities.1968b,61(4):927~933.
124. McLean DL. Probing behavior of the pea aphid, Acyrthosiphon pisum. V. Comparison of Vicia faba,Pisum sativum, and a chemically defined diet as food sources.1971,64(2):499~503.
125. Miles PW. Aphid saliva. Biological Reviews1999,74:41~85.
126. Miller WJ, Ehrman L, Schneider D. Infectious speciation revisited: impact of symbiont-Depletionon female fitness and mating behavior of drosophila paulistorum. PLoS Pathogens2010,6(12):e1001214.
127. Montllor CB, Gildow FE. Feeding responses of two grain aphids to barley yellow dwarfvirus-infected oats. Entomologia Experimentalis et Applicata1986,42:63~69.
128. Montllor CB, Tjallingii WF. Stylet penetration by two aphid species on susceptible and resistantlettuce. Entomologia Experimentalis et Applicata1989,52:103~111.
129. Moreira LA, Saig E, Turley AP, Ribeiro JMC, O'Neill SL, McGraw EA. Human probing behaviorof Aedes aegypti when Infected with a life-shortening strain of Wolbachia. PLoS Neglectedtropical Diseases2009,3(12): e568.
130. Mugiira RB, Liu SS, Zhou XP. Tomato yellow leaf curl virus and Tomato leaf curl Taiwan virusinvade south-east coast of China. Journal of Phytopathology2008,156:217~221.
131. Nauen R. Behaviour modifying effects of low systemic concentrations of imidacloprid on Myzuspersicae with special reference to an antifeeding response. Pesticide Science1995,44(2):145~153.
132. Niemeyer HM. Hydroxamic acid (4-hydroxy-1,4-benzoxazin-3-ones), defence chemicals in theGramineae. Phytochemistry1988,27:3349~3358.
133. Oliver KM, Degnan PH, Hunter MS, Moran NA. Bacteriophages encode factors required forprotection in a symbiotic mutualism. Science2009,325(5943):992~994.
134. Oliver KM, Moran NA, Hunter MS. Variation in resistance to parasitism in aphids is due tosymbionts not host genotype. Proceedings of the National Academy of Sciences of the UnitedStates of America2005,102:12795~12800.
135. Oliver KM, Russell JA, Moran AN, Hunter MS. Facultative bacterial symbionts in aphids conferresistance to parasitic wasps. Proceedings of the National Academy of Sciences of the UnitedStates of America2002,100:1803~1807.
136. Oliver KM, Russell JA, Moran NA, Hunter MS. Facultative bacterial symbionts in aphids conferresistance to parasitic wasps. Proceedings of the National Academy of Sciences of the UnitedStates of America2003,100:1803~1807.
137. Pan HP, Chu D, Ge DQ, Wang SL, Wu QJ, Xie W, Jiao XG, Liu BM, Yang X, Yang NN, Su Q, XuBY, Zhang YJ. Further spread of and domination by Bemisia tabaci (Hemiptera: Aleyrodidae)biotype Q on field crops in China. Journal of Economic Entomology2011,104(3):978~985.
138. Pan HP, Chu D, Yan WQ, Su Q, Liu BM, Wang SL, Wu QJ, Xie W, Jiao XG, Li RM, Yang NN,Yang X, Xu BY, Brown JK, Zhou XG, Zhang YJ. Rapid spread of Tomato yellow leaf curl virus inChina is aided differentially by two invasive whiteflies. PLoS ONE2012a,7: e34817.
139. Pan HP, Li XC, Ge DQ, Wang SL, Wu QJ, Xie W, Jiao XG, Chu D, Liu BM, Xu BY, Zhang YJ.Factors affecting population dynamics of maternally transmitted endosymbionts in Bemisia tabaci.PLoS ONE2012b,7: e30760.
140. Park J, Jahan SM, Song WG, Lee H, Lee YS, Choi HS, Lee KS, Kim CS, Lee S, Lee KY.Identification of biotypes and secondary endosymbionts of Bemisia tabaci in Korea andrelationships with the occurrence of TYLCV disease. Journal of Asia-Pacific Entomology2012,15(1):186~191.
141. Perring TM, Cooper AD, Rodriguez RJ, Farrar CA, Bellows TS. Identification of a whiteflyspecies by genomic and behavioral studies. Science1993,259:74~77.
142. Perring TM. Biological differences of two species of Bemisia that contribute to adaptive advantage.(in) Gerling D and Mayer RT (eds). Bemisia1995: Taxonomy, Biology, Damage Control andManagement. Intercept, Andover, UK,1996,3~16.
143. Perring TM. The Bemisia tabaci species complex. Crop Protection2001,20:725~737.
144. Polston JE, McGovern RJ, Brown LG. Introduction of tomato yellow leaf curl virus in Florida andimplications for the spread of this and other geminiviruses of tomato. Plant Disease1999,83:984~988.
145. Prado E, Tjallingii WF. Aphid activities during sieve element punctures. EntomologiaExperimentalis Applicata1994,72:157~165.
146. Prado E, Tjallingii WF. Behavioral dvidence for local reduction of aphid-induced resistance.Journal of Insect Science2007,7(8):1~8.
147. Prado E, Tjallingii WF. Effects of previous plant infestation on sieve element acceptance by twoaphids. Entomologia Experimentalis et Applicata1997,82:189~200.
148. Ramírez CC, Niemeyer HM. The influence of previous experience and starvation on aphid feedingbehavior. Journal of Insect Behavior2000,13(5):699~710.
149. Rao Q, Luo C, Zhang H, Guo X, Devine GJ. Distribution and dynamics so Bemisia tabaci invasivebiotypes in central China. Bulletin of Entomological Research2011,101(1):81~88.
150. Rosell RC, Bedford ID, Frohlich DR, Gill RJ, Brown JK. Analysis of morphological variation indistinct populations of Bemisia tabaci (Homoptera: Aleyrodidae). Annals of Entomological Societyof America1997,90:575~589.
151. Rubinstein G, Czosnek H. Long-term association of tomato yellow leaf curl virus (TYLCV) withits whitefly vector Bemisia tabaci: Effect on the insect transmission capacity, longevity andfecundity. Journal of General Virology1997,78:2683~2689.
152. Sarria E, Cidb M, Garzob E, Fereresb A. Excel Workbook for automatic parameter calculation ofEPG data. Computers and Electronics in Agriculture2009,67:35~42.
153. Schaefers GA. The use of direct current for electronically recording of aphid feeding and salivation.Annuals of Entomological Society of America1966,59:1022~1029.
154. Seal SE, vanden Bosch F, Jeger MJ. Factors influencing begomovirus evolution and theirincreasing global significance: implications for sustainable control. Critical Review in PlantScience2006,25:23~46.
155. Simon JC, Carré S, Boutin M, Prunier-Leterme N, Sabater-Mun B, Latorre A, Bournoville R.Host-based divergence in populations of the pea aphid: insights from nuclear markers and theprevalence of facultative symbionts. Proceedings of the Royal Society B2003,270:1703~1712.
156. Spiller NJ, Koenders L, Tjallingii WF. Xylem ingestion by aphids-a strategy for maintainingwater balance. Entomologia Experimentalis et Applicata1990,55(2):101~104.
157. Stafford CA, Walker GP, Ullman D. Hitching a ride: Vector feeding and virus transmission.Communicative&Integrative Biology2012,5(1):43~49.
158. Stafford CA, Walker GP, Ullman D. Infection with a plant virus modifies vector feeding behavior.Proceedings of the National Academy of Sciences of the United States of America2011,108:9350~9355.
159. Sun ZW, Yang JX, Zhang MZ, Li XD. Occurrence and control of Tomato yellow leaf curl virusdisease in Shandong Province. China Vegetables2009,21:5~6.
160. Teng X, Wan FH, Chu D. Bemisia tabaci biotype Q dominates other biotypes across China. FloridaEntomology2010,93(3):363~368.
161. Tjallingii WF, Gabry B. Anomalous stylet punctures of phloem sieve elements by aphids.Entomologia Experimentalis et Applicata1999,91:97~103.
162. Tjallingii WF. Electrical nature of recorded signals during stylet penetration by aphids.Entomologia experimentalis et applicata1985,38:177~186.
163. Tjallingii WF. Electrical recording of stylet penetration activities.(in) Minks AK and Harrewijn P(Eds). Aphids, their biology, natural enemies and control. Elsevier, Amsterdam1988,95~108.
164. Tjallingii WF. Electronic recording of penetration behaviour by aphids. EntomologiaExperimentalis et Applicata1978,24:521~530.
165. Tjallingii WF. Mechanoreceptors of the aphid labium. Entomologia Experimentalis et Applicata1978,24(3):731~737.
166. Tjallingii WF. Salivary secretions by aphids interacting with proteins of phloem wound responses.Journal of Experimental Botany2006,57:739~745.
167. Tjallingii WF. Sieve element acceptance by aphids. European Journal of Entomology1994,91:47~52.
168. Turley AP, Moreira LA, O'Neill SL McGraw EA. Wolbachia infection reduces blood-feedingsuccess in the dengue fever mosquito, Aedes aegypti. PLoS Neglected Tropical Diseases2009,39:1~6.
169. Urbanska A, Tjallingii WF, Dixon AFG, Leszczynski B. Phenol oxidising enzymes in the grainaphid’s saliva. Entomologia Experimentalis et Applicata1998,86(2):197~203.
170. van Helden M, Tjallingii WF. Tissue localisation of lettuce resistance to the aphid Nasonoviaribisnigri using electrical penetration graphs. Entomologia Experimentalis et Applicata1993,68:269~278.
171. van Helden M, Tjallingii WF. Experimental design and analysis in EPG experiments with emphasison plant resistance research.(in) Walker GP and Backus EA (eds). Principles and applications ofelectronic monitoring and other techniques in the study of Homopteran Feeding Behavior. ThomasSay Publications in Entomology, Entomological Society of America2000,144~171.
172. van Helden M, Tjallingii WF. Phloem sap collection from lettuce (Lactuca sativa L.): Chemicalcomparison among collection methods. Journal of Chemical Ecology1994,20:3191~3206.
173. Villada ES, González EG, López-Sesé AI, Castiel AF, Gómez-Guillamón ML. Hypersensitiveresponse to Aphis gossypii clover in melon genotypes carrying the Vat gene. Journal ofExperimental Botany2009,(60)11:3269~3277.
174. Vos P, Hogers R, Bleeker M, Reijans M, van de Lee T, Hornes M, Frijters A, Pot J, Peleman J,Kuiper M, Zabeau M. AFLP: a new technique for DNA fingerprinting. Nucleic Acids Research1995,23:4407~4414.
175. Walker GP, Gordh G. The occurrence of apical labial sensilla in the Aleyrodidae and evidence for acontact chemosensory function. Entomologia Experimentalis et Applicata1989,51:215~224.
176. Walker GP, Perring TM. Feeding and oviposition behavior of whiteflies (Homoptera, Aleyrodidae)interpreted from AC electronic feeding monitor waveforms. Annuals of Entomological Society ofAmerica1994,87:363~374.
177. Walker GP. Stylet penetration by the bayberry whitefly, as affected by leaf age in lemon, Citruslimon. Entomologia Experimentalis et Applicata1985,39:115~121.
178. Walker GP. The role of leaf cuticle in leaf age preference by bayberry whitefly (Homoptera:Aleyrodidae) on Lemon. Annals of the Entomological Society of America1988,81(2):365~369.
179. Werren JH, Baldo L, Clark ME. Wolbachia: master manipulators of invertebrate biology. NatureReviews Microbiology2008,6:741~751.
180. Wilkinson TL, Douglas AE. Aphid feeding, as influenced by disruption of the symbiotic bacteria:an analysis of the pea aphid (Acyrthosiphon pisum). Journal of Insect Phyiology1995,41(8):635~640.
181. Will T, Tjallingii WF, Th nnessen A, van Bel AJE. Molecular sabotage of plant defense by aphidsaliva. Proceedings of National Academy Science of the United States of America2007,104(25):10536~10541.
182. Wu JB, Dai FM, Zhou XP. First report of Tomato yellow leaf curl virus in China. Annals of AppliedBiology2006,155:439~448.
183. Xu J, Lin KK, Liu SS. Performance on different host plants of an alien and an indigenous Bemisiatabaci from China. Journal of Applied Entomology2011,135(10):771~779.
184. Xue K, Deng S, Wang RJ, Yan FM, Xu CR. Leaf surface factors of transgenic Bt cotton associatedwith the feeding behaviors of cotton aphids: A case study on non-target effects. Science in ChinaSeries C: Life Sciences2008,51(2):145~156.
185. Xue K, Wang XY, Huang CH, Wang RJ, Liu B, Yan FM, Xu CR. Stylet penetration behaviors ofthe cotton aphid Aphis gossypii on transgenic Bt cotton. Insect Science2008,16:137~146.
186. Yin HD, Wang XY, Xue K, Huang CH, Wang RJ, Yan FM, Xu CR. Impacts of transgenic Bt cottonon the stylet penetration behaviors of Bemisia tabaci biotype B: Evidence from laboratoryexperiments. Insect Science2010,17(4):344~352.
187. Zhang AH, Zhang SM, Liu S, Di DP, Miao HQ. Occurrence and distribution of tomato yellow leafcurl disease in Hebei Province. Plant Protection2010,36:127~129.
188. Zhang LP, Zhang YJ, Zhang WJ, Wu QJ, Xu BY, Chu D. Analysis of genetic diversity amongdifferent geographical populations and determination of biotypes of Bemisia tabaci in China.Journal of Applied Entomology2005129(3):121~128.
189. Zhou T, Shi YC, Chen XY, Fan ZF. Identification and control of Tomato yellow leaf curl virusdisease in Beijing. Plant Protection2010,36:116~118.
2.褚栋,陈国发,徐宝云,吴青君,张友军.烟粉虱B型和Q型群体遗传结构的RAPD分析.昆虫学报,2007,50(3):264~270.
3.罗晨,姚远,王戎疆,阎凤鸣,胡敦孝,张芝利.利用mtDNA COI基因序列鉴定我国烟粉虱的生物型.昆虫学报,2002,45(6):759~763.
4.罗晨,张芝利.烟粉虱Bemisia tabaci (Gennadius)研究概述.北京农业科学,2000,18（增刊）:4~13.
5.张芝利.关于烟粉虱大发生的思考.北京农业科学（烟粉虱专辑）,2000:1~3.
6.周尧.中国粉虱名录.中国昆虫学,1949,3(4):1~18.
7. Alvarez AE, Garzo E, Verbeek M, Vosman B, Dicke M, Tjallingii WF. Infection of potato plantswith Potato Leafroll Virus changes attraction and feeding behaviour of Myzus persicae.Entomologia Experimentalis et Applicata2007,125:135~144.
8. Alvarez AE, Tjallingii WF, Garzo E, Vleeshouwers V, Dicke M, Vosman B. Location of resistancefactors in the leaves of potato and wild tuber-bearing Solanum species to the aphid Myzus persicae.Entomologia Experimentalis et Applicata2006,121(2):145~157.
9. Annan IB, Schaefers GA, Tingey WM, Tjallingii WF. Effects of treatments for electricalpenetration graph recordings on behaviour and biology of Aphis craccivora (Aphididae).Physiological Entomology1997,22(2):95~101.
10. Backus EA, Bennett WH. The AC-DC Correlation Monitor: New EPG design with flexible inputresistors to detect both R and emf components for any piercing-sucking hemipteran. Journal ofInsect Physilology2009,55(10):869~884.
11. Backus EA, Serrano MS, Ranger CM. Mechanisms of hopperburn: An overview of insecttaxonomy, behavior, and physiology. Annual Review of Entomology2005,50:125~151.
12. Backus EA. Sensory systems and behaviours which mediate hemipteran plant-feeding: ataxonomic overview. Journal of Insect Physiology1988,34:151~165.
13. Baumann P. Biology bacteriocyte-associated endosymbionts of plant sap-sucking insects. AnnualReview of Microbiology2005,59:155~189.
14. Bedford ID, Briddon RW, Brown JK, Rosell RC, Markham PG. Geminivirus transmission andbiological characterization of Bemisia tabaci (Gennadius) from different geographic regions.Annals of Applied Biology1994,125:311~325.
15. Bedford ID, Briddon RW, Markham PG, Brown JK, Rosell RC. A new species of Bemisia orbiotype of Bemisia tabaci (Genn.), as a future pest of European agriculture.(in) Plant Health andEuropean Single Market, D. Ebbles (eds). BCPC Monograph1993,54:381~386.
16. Berry SD, FondongVN, Rey C, Rogan D, Fauquet CM, Brown JK. Molecular evidence for fivedistinct Bemisia tabaci (Homoptera: Aleyrodidae) geographic haplotypes associated with cassavaplants in Sub-Saharan Africa. Annals of the Entomological Society of America2004,97:852~859.
17. Bird J. A whitefly-transmitted mosaic of Jatropha gossypifolia. Technical Paper, AgriculturalExperimental Station, University of Puerto Rico1957,22:1~35.
18. Blackmer JL, Byrne DN. Environmental and physiological factors influencing phototactic flight ofBemisia tabaci. Physiology Entomology1993b,18:336~342.
19. Blackmer JL, Byrne DN. Flight behaviour of Bemisia tabaci in a vertical flight chamber: effect oftime of day, sex, age and host quality. Physiology Entomology1993a,18:223~232.
20. Bonani JP, Fereres A, Garzo E, Miranda MP, Appezzato-Da-Gloria B, Lopes JRS. Characterizationof electrical penetration graphs of the Asian citrus psyllid, Diaphorina citri, in sweet orangeseedlings. Entomologia Experimentalis et Applicata2010,134:35~49.
21. Boykin LM, Shatters RG, Rosell RC, McKenzie CL, Bagnall RA, De Barro PJ, Frohlich DR.Global relationships of Bemisia tabaci (Hemiptera: Aleyrodidae) revealed using Bayesian analysisof mitochondrial COI DNA sequences. Molecular Phylogenetics and Evolution2007,44:1306~1319.
22. Bradley RHE, Sylvester ES, Wade CV. Note on the movements of the mandibular and maxillarystylets of the aphid, Myzus persicae (Sulzer). The Canadian Entomologist1962,94:653~654.
23. Briddon RW, Pinner MS, Stanley J, Markham PG. Geminivirus coat protein gene replacementalters insect specificity. Virology1990,177:85~94.
24. Brown CM, Holbrook FR. An improved electronic system for monitoring feeding of Aphids.American Journal of Potato Research1976,53:457~462.
25. Brown JK, Bird J. Variability within the Bemisia tabaci species complex and its relation to newepidemics caused by geminiviruses. CEIBA1995,36:73~80.
26. Brown JK, Coats SA, Bedford ID, Markham PG, Bird J. Biotypic characterization of Bemisiatabaci populations based on esterase profiles, DNA fingerprinting, virus transmission, and bioassayto key host plant species. Phytopathology1992,82:1104.
27. Brown JK, Frohlich DR, Rosell RC. The sweetpotato or silverleaf whiteflies: biotypes of Bemisiatabaci or a species complex? Annual Review of Entomology1995,40:511-534.
28. Brown JK. Current status of Bemisia tabaci as a plant pest and virus vector in agro-ecosystemsworldwide. Bulletin Phytosanitaire de la FAO1994,42:3~32.
29. Brown JK. Molecular markers for the identification and global tracking of whitefly vector-Begomovirus complexes. Virus Research2000,71:233~260.
30. Brown JK. Phylogenetic biology of the Bemisia tabaci sibling species group.(in) PA Stansly andSE Naranjo (eds). Bionomics and Management of a Global Pest. Springer, Amsterdam2010,31~67.
31. Brown JK. The Bemisia tabaci complex: genetic and phenotypic variation and relevance toTYLCV-vector interactions.(in) Czosnek H (eds). Tomato yellow leaf curl virus disease:management, molecular biology, breeding for resistance. Springer, Dordrecht2007,25~56.
32. Burban C, Fishpool LDC, Fauquet C, Fargette D, Thouvenel JC. Host-associated biotypes withinWest African populations of the whitefly Bemisia tabaci (Genn.),(Hom., Aleyrodidae). Journal ofApplied Entomology1992,113:416~423.
33. Butler CG. On the ecology of Aleurodes brassicae Walk.(Hemiptera). Transactions of the RoyalEntomological Society of London1938,87:291~311.
34. Calatayud PA, Rahb Y, Tjallingii WF, Tertuliano M, Le Rü B. Electrically recorded feedingbehaviour of cassava mealybug on host and non-host plants. Entomologia experimentalis etapplicata1994,72:219~232.
35. Campbell BC, Jones KC, Dreyer DL. Discriminative behavioral responses by aphids to variousplant matrix Polysaccharides. Entomologia Experimentalis et Applicata1986,41(1):17~24.
36. Casta eda LE, Sandrock C, Vorburger C. Variation and covariation of life history traits in aphidsare related to infection with the facultative bacterial endosymbiont Hamiltonella defensa.Biological Journal of the Linnean Society2010,100(1):273~247.
37. Cervera MT, Cabezas JA, Simón B, Martínez-Zapater JM, Beitia F, Cenis JL. Genetic relationshipsamong biotypes of Bemisia tabaci (Hemiptera: Aleyrodidae) based on AFLP analysis. Bulletin ofEntomological Research2000,90:391~396.
38. Cherqui A, Tjallingii WF. Salivary proteins of aphids, a pilot study on identification, separation andimmunolocalisation. Journal of Insect Physiology2000,46:1177~1186.
39. Chiel E, Gottlieb Y, Zchori-Fein E, Mozes-Daube N, Katzir N, Inbar M, Ghanim M.Biotype-dependent secondary symbiont communities in sympatric populations of Bemisia tabaci.Bulletin of Entomological Research2007,97:407~413.
40. Chiel E, Inbar M, Mozes-Daube N, White JA, Hunter MS, Zchori-Fein E. Assessments of FitnessEffects by the Facultative Symbiont Rickettsia in the Sweetpotato Whitefly (Hemiptera:Aleyrodidae). Annals of the Entomological Society of America2009,102(3):413~418.
41. Chu D, Gao CS, De Barro P, Zhang YJ, Wan FH, Khan IA. Further insights into the strange role ofbacterial endosymbionts in whitefly, Bemisia tabaci: comparison of secondary symbionts frombiotypes B and Q in China. Bulletin of Entomological Research2011,101(4):477~486.
42. Chu D, Gao CS, De Barro PJ, Wan FH, Zhang YJ. Investigation of the genetic diversity of aninvasive whitefly (Bemisia tabaci) in China using both mitochondrial and nuclear DNA markers.Bulletin of Entomological Research2011a,101(4):467~475.
43. Chu D, Wan FH, Xu BY, Wu QJ, Zhang YJ. Comparative analysis of population genetic structurein Bemisia tabaci (Gennadius) biotypes B and Q based on ISSR marker. Agricultural Sciences inChina2008,7(11):1348~1354.
44. Chu D, Wan FH, Zhang YJ, Brown JK. Change in the biotype composition of Bemisia tabaci inShandong province of China from2005to2008. Environmental Entomology2010a,39(3):1028~1036.
45. Chu D, Zhang YJ, Brown JK, Cong B, Xu BY, Wu QJ, Zhu GR. The introduction of the exotic Qbiotype of Bemisia tabaci (Gennadius) from the Mediterranean region into China on ornamentalcrops. Florida Entomologist2006,89(2):168~174.
46. Chu D, Zhang YJ, Wan FH. Cryptic invasion of the exotic Bemisia tabaci biotype Q occurredwidespread in Shandong Province of China. Florida Entomologist2010b,93:203~207.
47. Cohen AC, Hendrix DL. Demonstration and preliminary characterization of α-amylase in thesweetpotato whitefly, Bemisia tabaci (Aleyrodidae: Homoptera). Comparative Biochemistry andPhysiology Part B1994,109:593~601.
48. Cohen AC, Henneberry TJ, Chu CC. Geometric relationships between whitefly feeding behaviorand vascular bundle arrangements. Entomologia Experimentalis et Applicata1996,78:135~142.
49. Cohen S, Lapidot M. Appearance and expansion of TYLCV: a historical point of view.(in) HCzosnek (eds). Tomato Yellow Leaf Curl Virus Disease. Springer,2007,3~12.
50. Costa HS, Brown JK. Variation in biological characteristics and esterase patterns amongpopulations of Bemisia tabaci, and the association of one population with silverleaf symptominduction. Entomologia Experimentalis et Applicata1991,61:211~219.
51. Crowder DW, Ellers-Kirk C, Yafuso C, Dennehy TJ, Degain BA, Harpold VS, Tabashnik BE,Carrière Y. Inheritance of resistance to pyriproxyfen in Bemisia tabaci (Hemiptera: Aleyrodidae)males and females (B biotype). Journal of Economic Entomology2008,101:927~932.
52. Crowder DW, Horowitz AR, De Barro PJ, Liu SS, Showalter AW, Kontsedalov S, Khasdan V,Shargal A, Liu J, Carrière Y. Mating behaviour, life history and adaptation to insecticides determinespecies exclusion between whiteflies. Journal of Animal Ecology2010a,79:563~570.
53. Crowder DW, Sitvarin MI, Carrière Y. Plasticity in mating behaviour drives asymmetricreproductive interference in whiteflies. Animal Behaviour2010b,79:579~587.
54. Czosnek H, Ghanim M. Bemisia tabaci-Tomato Yellow Leaf Curl Virus interaction causingworldwide epidemics.(in) WMO Thompson (eds). the whitefly, Bemisia tabaci (Homoptera:Aleyrodidae) interaction with Geminivirus-infected host plants. Springer2011,51~67.
55. De Barro PJ, Driver F. Use of RAPD PCR to distinguish the B biotype from other biotypes ofBemisia tabaci (Gennadius)(Hemiptera, Aleyrodidae). Australian Journal of Entomology1997,36:149~152.
56. De Barro PJ, Liu SS, Boykin LM, Dinsdale A. Bemisia tabaci: a statement of species status.Annual Review Entomology2011,56:1~19.
57. De Barro PJ. Genetic structure of the whitefly Bemisia tabaci in the Asia–Pacific region revealedusing microsatellite markers. Molecular Ecology2005,14:3695~3718.
58. Dinsdale A, Cook L, Riginos C, Buckley YM, De Barro PJ. Refined global analysis of Bemisiatabaci (Hemiptera: Sternorrhyncha: Aleyrodoidea: Aleyrodidae) mitochondrial cytochrome oxidase1to identify species level genetic boundaries. Annals of the Entomological Society of America2010,103:196~208.
59. Dreyer DL, Campbell BC. Chemical basis of host-plant resistance to aphids. Plant CellEnvironment1987,10:353~361.
60. Dugravot S, Backus EA, Reardon BJ, Miller TA. Correlations of cibarial muscle activities ofHomalodisca spp. Sharpshooters (Hemiptera: Cicadellidae) with EPG ingestion waveform andexcretion. Journal of Insect Physiology2008,54:1467~1478.
61. Fauquet CM, Maxwell DP, Gronenborn B, Stanley J. Revised proposal for naming geminiviruses.Archives of Virology2000,145:1743~1761.
62. Fereres A, Moreno A. Behavioural aspects influencing plant virus transmission by homopteraninsects. Virus Research2009,141:158~168.
63. Freeman TP, Buckner JS, Nelson DR, Chu CC, Henneberry TJ. Stylet penetration by Bemisiaargentifolii (Homoptera: Aleyrodidae) into host leaf tissue. Annals of the Entomological Society ofAmerica2001,94:761~768.
64. Frohlich DR, Torres-Jerez I, Bedford ID, Markham PG, Brown JK. A phylogeographical analysis ofthe Bemisia tabaci species complex based on mitochondrial DNA markers. Molecular Ecology1999,8:1683~1691.
65. Funk CJ. Alkaline phosphatase activity in whitefly salivary glands and saliva. Archieves of InsectBiochemistry and Physiology2001,46:165~174.
66. Gabrys B, Tjallingii WF, van Beek TA. Analysis of EPG recorded probing by cabbage aphid onhost plant parts with different glucosinolate contents. Journal of Chemical Ecology1997,23(7):1661~1673.
67. Garzo E, Soria C, Gómez-Guillamón ML, Alberto F. Feeding behavior of Aphis gossypii onresistant accessions of different melon genotypes (Cucumis melo). Phytoparasitica2002,30(2):129~140.
68. Gelman DB, Blackburn MB, Hu JS, Gerling D. The nymphal-adult molt of the silverleaf whitefly(Bemisia argentifolii): timing, regulation, and progress. Archieves of Insect Biochemistry andPhysiology2002,51:67~79.
69. Gennadius P. Disease of tobacco plantations in Trikonia: the aleurodid of tobacco. EllennikeGeorgia1889,6(5): l~3.
70. Ghanim M, Rosell RC, Campbell LR, Czosnek H, Brown JK, Ullman DE. Digestive, salivary andreproductive organs of Bemisia tabaci (Gennadius)(Hemiptera: Aleyrodidae) type B. Journal ofMorphology2001,248:22~40.
71. Ghanim M, Sobol I, Ghanim M, Czosnek H. Horizontal transmission of begomoviruses betweenBemisia tabaci biotypes. Arthropod-Plant Interaction2007,1(3):195~204.
72. Gill RJ. A review of the sweetpotato whitefly in Southern California. The Pan-Pacific Entomologist1992,68:144~152.
73. Gottieb Y, Zchori-Fein E, Mozes-Daube N, Kontsedalov S, Skaljac M, Brumin I, Czosnek H,Vavre F, Fleury F, Ghanim M. The Transmission efficiency of Tomato Yellow Leaf Curl Virus bythe whitefly Bemisia tabaci is correlated with the presence of a specific symbiotic bacteriumspecies. Journal of Virology2010,84(18):9310~9317.
74. Gottlieb Y, Ghanim M, Chiel Eald, Gerling D, Portnoy V, Steinberg S, Tzuri G, Horowitz AR,Belausov E, Mozes-Daube N, Kontsedalov S, Gershon M, Shunit G, Katzir N, Zchori-Fein E.Identification and localization of a Rickettsia sp. in Bemisia tabaci (Homoptera: Aleyrodidae).Applied and Environmental Microbiology2006,72:3646~3652.
75. Gottlieb Y, Ghanim M, Gueguen G, Kontsedalov S, Vavre F, Fleury F, Zchori-Fein E. Inheritedintracellular ecosystem: symbiotic bacteria share bacteriocytes in whiteflies. The FASEB Journal2008,22:2591~2599.
76. Gueguen G, Vavre F, Gnankine O, Peterschmitt M, Charif D, Chiel E, Gottlieb Y, Ghanim M,Zchori-Fein E, Fleury F. Endosymbiont metacommunities, mtDNA diversity and the evolution ofthe Bemisia tabaci (Hemiptera: Aleyrodidae) species complex. Molecular Ecology2010,19(19):4365~4376.
77. Guershon M, Gerling D. Effect of foliar tomentosity on phenotypic plasticity in Bemisia tabaci(Hom., Aleyrodidae). Journal of Applied Entomology2001,125:449~453.
78. Guershon M, Gerling D. Predatory behavior of Delphastus pusillus in relation to the phenotypicplasticity of Bemisia tabaci nymphs. Entomologia Experimentalis et Applicata1999,92:239~248.
79. Guirao P, Beitia F, Cenis JL. Biotype determination of Spanish populations of Bemisia tabaci(Hemiptera: Aleyrodidae). Bulletin of Entomological Research1997,87:587~593.
80. Harrewijn P, Piron PGM, Mollema C. Electrically recorded probing behaviour of thrips species onoptimal and suboptimal hosts. Entomologia Experimentalis et Applicata,1996,80(1):43~45.
81. Harris KF, Esbroeck ZP-V, Duffus JE. Morphology of the sweet potato whitefly, Bemisia tabaci(Homoptera, Aleyrodidae) relative to virus transmission. Zoomorphology1996,116:143~156.
82. Harrison BD, Robinson DJ. Natural genomic and antigenic variation in whitefly transmittedgeminiviruses (Begomoviruses). Annual Review of Phytopathology1999,37:369~398.
83. Hedges LM, Brownlie JC, O'Neill SL, Johnson KN. Wolbachia and virus protection in insects.Science2008,322(5902):702.
84. Helmi A. Host-associated population variations of Bemisia tabaci (Genn.)(Hemiptera:Sternorrhyncha: Aleyrodidae) characterized with random DNA markers. International Journal ofZoological Research2011,7(1):77~84.
85. Hilker M, Meiners T. Early herbivore alert: insect eggs induce plant defense. Journal of ChemicalEcology2006,32(7):1379~1397.
86. Himler AG, Adachi-Hagimori T, Bergen JE, Kozuch A, Kelly SE, Tabashnik BE, Chiel E,Duckworth VE, Dennehy TJ, Zchori-Fein E, Hunter MS. Rapid spread of a bacterial symbiont inan Invasive Whitefly is driven by fitness benefits and female bias. Science2011,332(6026):254~256.
87. Horowitz, AR, Kontsedalov S, Khasdan V, Ishaaya I. Biotypes B and Q of Bemisia tabaci and theirrelevance to neonicotinoid and pyriproxyfen resistance. Archives of Insect Biochemistry andPhysiology2005,58:216~225.
88. Hu J, De Barro P, Zhao H, Wang J, Nardi F, Liu SS. An extensive field survey combined with aphylogenetic analysis reveals rapid and widespread invasion of two alien whiteflies in China. PLoSONE2011,6(1): e16061.
89. Jaenike J, Papaj DR. Behavioural plasticity and pattern of host use by insect.(in) Roitberg BD, andIsman MB (eds). Insect chemical ecology, an evolutionary approach, Chapman and Hall. New York,1992,245~264.
90. Janssen JAM, Tjallingii WF, van Lenteren JC. Electrical recording and ultrastructure of styletpenetration by the greenhouse whitefly. Entomologia Experimentalis et Applicata1989,52:69~81.
91. Ji YH, Xiong RY, Cheng ZB, Zhou T, Zhao TM, Yu WG, Fan YJ, Zhou YJ. Molecular diagnosis ofTomato yellow leaf curl disease in Jiangsu Province. Acta Horticulturae Sinica2008,35:1815~1818.
92. Jiang YX, Blas C, Barrios L, Fereres A. Correlation between whitefly (Homoptera: Aleyrodidae)feeding behavior and transmission of tomato yellow leaf curl virus. Annals of EntomologicalSociety of America2000,93:573~579.
93. Jiang YX, Lei H, Collar JL, Martin B, Muniz M, Fereres A. Probing and feeding behavior of twodistinct biotypes of Bemisia tabaci (Homoptera: Aleyrodidae) on tomato plants. Journal ofEconomic Entomology1999,92:357~366.
94. Jiang YX, Nombela G, Muniz M. Analysis by DC-EPG of the resistance to Bemisia tabaci on anMi-tomato line. Entomologia Experimentalis et Applicata2001,99:295~302.
95. Jiang YX, Walker GP. Identification of phloem sieve elements as the site of resistance to silverleafwhitefly in resistant alfalfa genotypes. Entomologia Experimentalis et Applicata2007,125:307~320.
96. Jiang YX, Walker GP. Pathway phase waveform characteristics correlated with length and rate ofstylet advancement and partial stylet withdrawal in AC electrical penetration graphs of adultwhiteflies. Entomologia Experimentalis et Applicata2001,101:233~246.
97. Johnson DD, Walker GP, Creamer R. Stylet penetration behavior resulting in inoculation of asemipersistently transmitted closterovirus by the whitefly Bemisia argentifolii. EntomologiaExperimentalis et Applicata2002,102:115~123.
98. Johnson DD, Walker GP. Intracellular punctures by the adult whitefly Bemisia argentifolii on DCand AC electronic feeding monitors. Entomologia Experimentalis et Applicata1999,92:257~270.
99. Jones DR. Plant viruses transmitted by whiteflies. European Journal of Plant Pathology2003,109:195~219.
100. Joosta PH, Backus EA, Morgan D, Yan FM. Correlation of stylet activities by the glassy-wingedsharpshooter, Homalodisca coagulata (Say), with electrical penetration graph (EPG) waveforms.Journal of Insect Physiology2006,52:327~337.
101. Kimmins FM, Tjallingii WF. Ultrastucture of sieve element penetration by aphid stylets duringelectrical recording. Entomologia Experimentalis et Applicata1985,39:135~141.
102. Kindt F, Joosten NN, Tjallingii WF. Electrical penetration graphs of thrips revised: combining DC-and AC-EPG signals. Journal of Insect Physilology2006,52(1):1~10.
103. Knoblauch M, van Bel AJE. Sieve tubes in action. Plant Cell1998,10:35~50.
104. Lazarowitz SG, Shepherd DRJ. Geminiviruses: genome structure and gene function. CriticalReviews in Plant Sciences1992,11(4):327~349.
105. Lei H, Tjallingii WF, van Lenteren JC, Xu RM. Stylet penetration by larvae of the greenhousewhitefly on cucumber. Entomologia Experimentalis et Applicata1996,79:77~84.
106. Lei H, Tjallingii WF, van Lenteren JC. Probing and feeding characteristics of the greenhousewhitefly in association with host-plant acceptance and whitefly strains. EntomologiaExperimentalis et Applicata1998,88:73~80.
107. Lei H, Tjallingii WF, van Lenteren JC. Effect of tethering during EPG recorded probing by adultsof the greenhouse whitefly. Journal of Applied Entomology1997,121:211~217.
108. Lei H, van Lenteren JC, Tjallingii WF. Analysis of resistance in tomato and sweet pepper againstthe greenhouse whitefly using electrically monitored and visually observed probing and feedingbehaviour. Entomologia Experimentalis et Applicata1999,92:299~309.
109. Lei H, van Lenteren JC, Xu RM. Effects of plant tissue factors on the acceptance of fourgreenhouse vegetable host plants by the greenhouse whitefly: an electrical penetration graph (EPG)study. European Journal of Entomolomogy2001,98:31~36.
110. Leopolda RA, Freemanb TP, Bucknera JS, Nelsona DR. Mouthpart morphology and styletpenetration of host plants by the glassywinged sharpshooter, Homalodisca coagulata,(Homoptera:Cicadellidae). Arthropod Structure&Development2003,32:189~199.
111. Leszczynski B, Tjallingii WF, Dixon AFG, Swiderski R. Effect of methoxyphenols on grain aphidfeeding behavior. Enwmologia Experimentalis etApplicata1995,76:157~162.
112. Liu SS, De Barro PJ, Xu J, Luan JB, Zang LS, Ruan YM, Wan FH. Asymmetric matinginteractions drive widespread invasion and displacement in a whitefly. Science2007,318:1769~1772.
113. Luan JB, Li JM, Varela N, Wang YL, Li FF, Bao YY, Zhang CX, Liu SS, Wang XW. Globalanalysis of the transcriptional response of whitefly to Tomato yellow leaf curl China virus revealsthe relationship of coevolved adaptations. Journal of Virology2011,85:3330~3340.
114. Marchetti E, Civolani S, Leis M, Chicca M, Tjallingii WF, Pasqualini E, Baronio P. Tissue locationof resistance in apple to the rosy apple aphid established by electrical penetration graphs. Bulletinof Insectology2009,62(2):203~208.
115. Martín B, Collar JL, Tjallingii WF, Fereres A. Intracellular ingestion and salivation by aphids maycause the acquisition and inoculation of non-persistently transmitted plant viruses Journal ofGeneral Virology1997,78:2701~2705.
116. Mayoral AM, Tjallingii WF, Casta era P. Probing behaviour of Diuraphis noxia on five cerealspecies with different hydroxamic acid levels. Entomologia Experimentalis et Applicata1996,78:341~348.
117. McKenzie CL, Hodges G, Osborne LS, Byrne FJ, Shatters Jr. RG. Distribution of Bemisia tabaci(Hemiptera: Aleyrodidae) biotypes in Florida-investigating the Q invasion. Journal of EconomicEntomology2009,102:670~676.
118. McLean DL, Kinsey MG. A technique for electronically recording aphid feeding and salivation.Nature1964,202:1358~1359.
119. McLean DL, Kinsey MG. Identification of electrically recorded curve patterns associated withaphid salivation and ingestion. Nature1965,205(4976):1130~1131.
120. McLean DL, Kinsey MG. Probing behavior of the pea aphid, Acyrthosiphon pisum. IV. Effects ofstarvation on certain probing activities.1969,62(5):987~994.
121. McLean DL, Kinsey MG. Probing behavior of the pea aphid, Acyrthosiphon pisum. I. Definitivecorrelation of electronically recorded waveforms with aphid probing activities.1967,(2):400~405.
122. McLean DL, Kinsey MG. Probing behavior of the pea aphid, Acyrthosiphon pisum. II.Comparisons of salivation and ingestion in host and non-host plant leaves.1968a,61(3):730~739.
123. McLean DL, Kinsey MG. Probing behavior of the pea aphid, Acyrthosiphon pisum. III. Effect oftemperature differences on certain probing activities.1968b,61(4):927~933.
124. McLean DL. Probing behavior of the pea aphid, Acyrthosiphon pisum. V. Comparison of Vicia faba,Pisum sativum, and a chemically defined diet as food sources.1971,64(2):499~503.
125. Miles PW. Aphid saliva. Biological Reviews1999,74:41~85.
126. Miller WJ, Ehrman L, Schneider D. Infectious speciation revisited: impact of symbiont-Depletionon female fitness and mating behavior of drosophila paulistorum. PLoS Pathogens2010,6(12):e1001214.
127. Montllor CB, Gildow FE. Feeding responses of two grain aphids to barley yellow dwarfvirus-infected oats. Entomologia Experimentalis et Applicata1986,42:63~69.
128. Montllor CB, Tjallingii WF. Stylet penetration by two aphid species on susceptible and resistantlettuce. Entomologia Experimentalis et Applicata1989,52:103~111.
129. Moreira LA, Saig E, Turley AP, Ribeiro JMC, O'Neill SL, McGraw EA. Human probing behaviorof Aedes aegypti when Infected with a life-shortening strain of Wolbachia. PLoS Neglectedtropical Diseases2009,3(12): e568.
130. Mugiira RB, Liu SS, Zhou XP. Tomato yellow leaf curl virus and Tomato leaf curl Taiwan virusinvade south-east coast of China. Journal of Phytopathology2008,156:217~221.
131. Nauen R. Behaviour modifying effects of low systemic concentrations of imidacloprid on Myzuspersicae with special reference to an antifeeding response. Pesticide Science1995,44(2):145~153.
132. Niemeyer HM. Hydroxamic acid (4-hydroxy-1,4-benzoxazin-3-ones), defence chemicals in theGramineae. Phytochemistry1988,27:3349~3358.
133. Oliver KM, Degnan PH, Hunter MS, Moran NA. Bacteriophages encode factors required forprotection in a symbiotic mutualism. Science2009,325(5943):992~994.
134. Oliver KM, Moran NA, Hunter MS. Variation in resistance to parasitism in aphids is due tosymbionts not host genotype. Proceedings of the National Academy of Sciences of the UnitedStates of America2005,102:12795~12800.
135. Oliver KM, Russell JA, Moran AN, Hunter MS. Facultative bacterial symbionts in aphids conferresistance to parasitic wasps. Proceedings of the National Academy of Sciences of the UnitedStates of America2002,100:1803~1807.
136. Oliver KM, Russell JA, Moran NA, Hunter MS. Facultative bacterial symbionts in aphids conferresistance to parasitic wasps. Proceedings of the National Academy of Sciences of the UnitedStates of America2003,100:1803~1807.
137. Pan HP, Chu D, Ge DQ, Wang SL, Wu QJ, Xie W, Jiao XG, Liu BM, Yang X, Yang NN, Su Q, XuBY, Zhang YJ. Further spread of and domination by Bemisia tabaci (Hemiptera: Aleyrodidae)biotype Q on field crops in China. Journal of Economic Entomology2011,104(3):978~985.
138. Pan HP, Chu D, Yan WQ, Su Q, Liu BM, Wang SL, Wu QJ, Xie W, Jiao XG, Li RM, Yang NN,Yang X, Xu BY, Brown JK, Zhou XG, Zhang YJ. Rapid spread of Tomato yellow leaf curl virus inChina is aided differentially by two invasive whiteflies. PLoS ONE2012a,7: e34817.
139. Pan HP, Li XC, Ge DQ, Wang SL, Wu QJ, Xie W, Jiao XG, Chu D, Liu BM, Xu BY, Zhang YJ.Factors affecting population dynamics of maternally transmitted endosymbionts in Bemisia tabaci.PLoS ONE2012b,7: e30760.
140. Park J, Jahan SM, Song WG, Lee H, Lee YS, Choi HS, Lee KS, Kim CS, Lee S, Lee KY.Identification of biotypes and secondary endosymbionts of Bemisia tabaci in Korea andrelationships with the occurrence of TYLCV disease. Journal of Asia-Pacific Entomology2012,15(1):186~191.
141. Perring TM, Cooper AD, Rodriguez RJ, Farrar CA, Bellows TS. Identification of a whiteflyspecies by genomic and behavioral studies. Science1993,259:74~77.
142. Perring TM. Biological differences of two species of Bemisia that contribute to adaptive advantage.(in) Gerling D and Mayer RT (eds). Bemisia1995: Taxonomy, Biology, Damage Control andManagement. Intercept, Andover, UK,1996,3~16.
143. Perring TM. The Bemisia tabaci species complex. Crop Protection2001,20:725~737.
144. Polston JE, McGovern RJ, Brown LG. Introduction of tomato yellow leaf curl virus in Florida andimplications for the spread of this and other geminiviruses of tomato. Plant Disease1999,83:984~988.
145. Prado E, Tjallingii WF. Aphid activities during sieve element punctures. EntomologiaExperimentalis Applicata1994,72:157~165.
146. Prado E, Tjallingii WF. Behavioral dvidence for local reduction of aphid-induced resistance.Journal of Insect Science2007,7(8):1~8.
147. Prado E, Tjallingii WF. Effects of previous plant infestation on sieve element acceptance by twoaphids. Entomologia Experimentalis et Applicata1997,82:189~200.
148. Ramírez CC, Niemeyer HM. The influence of previous experience and starvation on aphid feedingbehavior. Journal of Insect Behavior2000,13(5):699~710.
149. Rao Q, Luo C, Zhang H, Guo X, Devine GJ. Distribution and dynamics so Bemisia tabaci invasivebiotypes in central China. Bulletin of Entomological Research2011,101(1):81~88.
150. Rosell RC, Bedford ID, Frohlich DR, Gill RJ, Brown JK. Analysis of morphological variation indistinct populations of Bemisia tabaci (Homoptera: Aleyrodidae). Annals of Entomological Societyof America1997,90:575~589.
151. Rubinstein G, Czosnek H. Long-term association of tomato yellow leaf curl virus (TYLCV) withits whitefly vector Bemisia tabaci: Effect on the insect transmission capacity, longevity andfecundity. Journal of General Virology1997,78:2683~2689.
152. Sarria E, Cidb M, Garzob E, Fereresb A. Excel Workbook for automatic parameter calculation ofEPG data. Computers and Electronics in Agriculture2009,67:35~42.
153. Schaefers GA. The use of direct current for electronically recording of aphid feeding and salivation.Annuals of Entomological Society of America1966,59:1022~1029.
154. Seal SE, vanden Bosch F, Jeger MJ. Factors influencing begomovirus evolution and theirincreasing global significance: implications for sustainable control. Critical Review in PlantScience2006,25:23~46.
155. Simon JC, Carré S, Boutin M, Prunier-Leterme N, Sabater-Mun B, Latorre A, Bournoville R.Host-based divergence in populations of the pea aphid: insights from nuclear markers and theprevalence of facultative symbionts. Proceedings of the Royal Society B2003,270:1703~1712.
156. Spiller NJ, Koenders L, Tjallingii WF. Xylem ingestion by aphids-a strategy for maintainingwater balance. Entomologia Experimentalis et Applicata1990,55(2):101~104.
157. Stafford CA, Walker GP, Ullman D. Hitching a ride: Vector feeding and virus transmission.Communicative&Integrative Biology2012,5(1):43~49.
158. Stafford CA, Walker GP, Ullman D. Infection with a plant virus modifies vector feeding behavior.Proceedings of the National Academy of Sciences of the United States of America2011,108:9350~9355.
159. Sun ZW, Yang JX, Zhang MZ, Li XD. Occurrence and control of Tomato yellow leaf curl virusdisease in Shandong Province. China Vegetables2009,21:5~6.
160. Teng X, Wan FH, Chu D. Bemisia tabaci biotype Q dominates other biotypes across China. FloridaEntomology2010,93(3):363~368.
161. Tjallingii WF, Gabry B. Anomalous stylet punctures of phloem sieve elements by aphids.Entomologia Experimentalis et Applicata1999,91:97~103.
162. Tjallingii WF. Electrical nature of recorded signals during stylet penetration by aphids.Entomologia experimentalis et applicata1985,38:177~186.
163. Tjallingii WF. Electrical recording of stylet penetration activities.(in) Minks AK and Harrewijn P(Eds). Aphids, their biology, natural enemies and control. Elsevier, Amsterdam1988,95~108.
164. Tjallingii WF. Electronic recording of penetration behaviour by aphids. EntomologiaExperimentalis et Applicata1978,24:521~530.
165. Tjallingii WF. Mechanoreceptors of the aphid labium. Entomologia Experimentalis et Applicata1978,24(3):731~737.
166. Tjallingii WF. Salivary secretions by aphids interacting with proteins of phloem wound responses.Journal of Experimental Botany2006,57:739~745.
167. Tjallingii WF. Sieve element acceptance by aphids. European Journal of Entomology1994,91:47~52.
168. Turley AP, Moreira LA, O'Neill SL McGraw EA. Wolbachia infection reduces blood-feedingsuccess in the dengue fever mosquito, Aedes aegypti. PLoS Neglected Tropical Diseases2009,39:1~6.
169. Urbanska A, Tjallingii WF, Dixon AFG, Leszczynski B. Phenol oxidising enzymes in the grainaphid’s saliva. Entomologia Experimentalis et Applicata1998,86(2):197~203.
170. van Helden M, Tjallingii WF. Tissue localisation of lettuce resistance to the aphid Nasonoviaribisnigri using electrical penetration graphs. Entomologia Experimentalis et Applicata1993,68:269~278.
171. van Helden M, Tjallingii WF. Experimental design and analysis in EPG experiments with emphasison plant resistance research.(in) Walker GP and Backus EA (eds). Principles and applications ofelectronic monitoring and other techniques in the study of Homopteran Feeding Behavior. ThomasSay Publications in Entomology, Entomological Society of America2000,144~171.
172. van Helden M, Tjallingii WF. Phloem sap collection from lettuce (Lactuca sativa L.): Chemicalcomparison among collection methods. Journal of Chemical Ecology1994,20:3191~3206.
173. Villada ES, González EG, López-Sesé AI, Castiel AF, Gómez-Guillamón ML. Hypersensitiveresponse to Aphis gossypii clover in melon genotypes carrying the Vat gene. Journal ofExperimental Botany2009,(60)11:3269~3277.
174. Vos P, Hogers R, Bleeker M, Reijans M, van de Lee T, Hornes M, Frijters A, Pot J, Peleman J,Kuiper M, Zabeau M. AFLP: a new technique for DNA fingerprinting. Nucleic Acids Research1995,23:4407~4414.
175. Walker GP, Gordh G. The occurrence of apical labial sensilla in the Aleyrodidae and evidence for acontact chemosensory function. Entomologia Experimentalis et Applicata1989,51:215~224.
176. Walker GP, Perring TM. Feeding and oviposition behavior of whiteflies (Homoptera, Aleyrodidae)interpreted from AC electronic feeding monitor waveforms. Annuals of Entomological Society ofAmerica1994,87:363~374.
177. Walker GP. Stylet penetration by the bayberry whitefly, as affected by leaf age in lemon, Citruslimon. Entomologia Experimentalis et Applicata1985,39:115~121.
178. Walker GP. The role of leaf cuticle in leaf age preference by bayberry whitefly (Homoptera:Aleyrodidae) on Lemon. Annals of the Entomological Society of America1988,81(2):365~369.
179. Werren JH, Baldo L, Clark ME. Wolbachia: master manipulators of invertebrate biology. NatureReviews Microbiology2008,6:741~751.
180. Wilkinson TL, Douglas AE. Aphid feeding, as influenced by disruption of the symbiotic bacteria:an analysis of the pea aphid (Acyrthosiphon pisum). Journal of Insect Phyiology1995,41(8):635~640.
181. Will T, Tjallingii WF, Th nnessen A, van Bel AJE. Molecular sabotage of plant defense by aphidsaliva. Proceedings of National Academy Science of the United States of America2007,104(25):10536~10541.
182. Wu JB, Dai FM, Zhou XP. First report of Tomato yellow leaf curl virus in China. Annals of AppliedBiology2006,155:439~448.
183. Xu J, Lin KK, Liu SS. Performance on different host plants of an alien and an indigenous Bemisiatabaci from China. Journal of Applied Entomology2011,135(10):771~779.
184. Xue K, Deng S, Wang RJ, Yan FM, Xu CR. Leaf surface factors of transgenic Bt cotton associatedwith the feeding behaviors of cotton aphids: A case study on non-target effects. Science in ChinaSeries C: Life Sciences2008,51(2):145~156.
185. Xue K, Wang XY, Huang CH, Wang RJ, Liu B, Yan FM, Xu CR. Stylet penetration behaviors ofthe cotton aphid Aphis gossypii on transgenic Bt cotton. Insect Science2008,16:137~146.
186. Yin HD, Wang XY, Xue K, Huang CH, Wang RJ, Yan FM, Xu CR. Impacts of transgenic Bt cottonon the stylet penetration behaviors of Bemisia tabaci biotype B: Evidence from laboratoryexperiments. Insect Science2010,17(4):344~352.
187. Zhang AH, Zhang SM, Liu S, Di DP, Miao HQ. Occurrence and distribution of tomato yellow leafcurl disease in Hebei Province. Plant Protection2010,36:127~129.
188. Zhang LP, Zhang YJ, Zhang WJ, Wu QJ, Xu BY, Chu D. Analysis of genetic diversity amongdifferent geographical populations and determination of biotypes of Bemisia tabaci in China.Journal of Applied Entomology2005129(3):121~128.
189. Zhou T, Shi YC, Chen XY, Fan ZF. Identification and control of Tomato yellow leaf curl virusdisease in Beijing. Plant Protection2010,36:116~118.