烟粉虱Q型替代B型的生态学机制研究
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摘要
外来生物入侵危害国家的生物安全、生态安全和经济安全。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型是两种危害最大、入侵性最强的生物型。B型烟粉虱自20世纪90年代中期入侵我国，给我国农业生产、国民经济造成了巨大损失；Q型烟粉虱则于2003年首先在云南发现，并随后在我国华东、华北地区迅速扩散；同时，由烟粉虱传播的番茄黄化曲叶病毒也在各地暴发成灾。本论文通过对我国20多个省市多种作物上的烟粉虱种群的系统监测研究，发现烟粉虱入侵我国十多年后，其主要危害生物型已由B型转变为Q型；生物因子（寄主植物、内共生菌和植物病毒）与非生物因子（杀虫剂）对B型和Q型烟粉虱种群互作特性研究，证实杀虫剂的使用是导致B型烟粉虱被Q型烟粉虱替代的关键生态因子；进一步对B型与Q型烟粉虱获毒及传毒特性研究，发现烟粉虱Q型替代B型是导致TYLCV在我国暴发流行的机制。相关研究结果对入侵烟粉虱及其传播病毒的可持续控制具有重要指导意义。
主要研究内容和结论如下：
一、B型烟粉虱和Q型烟粉虱在我国的入侵分布动态
利用特异分子标记并结合mtCO1基因序列分析，分别于2009年和2011年对采自我国的19个省市和27个省市的烟粉虱种群的生物型进行了鉴定，结果发现，2009年的61个烟粉虱种群中有44个是Q型烟粉虱种群，17个是B型烟粉虱种群，Q型烟粉虱主要分布在长江流域及东部沿海地区；2011年的98个烟粉虱种群中有60个Q型烟粉虱种群，9个B型烟粉虱种群，29个混合种群中有20个种群Q型烟粉虱占有较高的比例（>50%），除了东南省份，Q型烟粉虱都占据主导优势。监测结果发现，烟粉虱入侵我国十多年后，其主要危害生物型已由早期入侵B型转变为Q型。
二、寄主植物对B型和Q型烟粉虱种群竞争取代的影响
将B型和Q型烟粉虱按各占50%的比例分别接种到一品红、辣椒、番茄和甘蓝四种寄主植物上，一定时间后随机采集寄主植物上一定数量的烟粉虱个体并鉴定其生物型，明确寄主植物对两种生物型烟粉虱种群间互作的影响。结果发现在一品红和辣椒植株上，分别经过8代和2代，Q型烟粉虱替代了B型烟粉虱；而在番茄和甘蓝植株上则刚好相反，分别经过5代和4代，B型烟粉虱替代了Q型烟粉虱。不同寄主植物对B型和Q型烟粉虱种群间的互作表现了不同的特性，导致了不同的替代方向和速度。
三、杀虫剂对B型和Q型烟粉虱种群竞争取代的影响
同样将B型和Q型烟粉虱按各占50%比例分别接种到棉花和黄瓜植物上，并分别设置一定浓度的杀虫剂处理和未处理的重复，一定时间后随机采集寄主植物上一定数量的烟粉虱个体并鉴定其生物型，明确杀虫药剂对两种生物型烟粉虱种群间互作的影响。结果发现，在未经杀虫剂处理的棉花植株上，经过6代Q型烟粉虱替代了B型烟粉虱，而在经一定剂量的噻虫嗪处理后的棉花植株上，Q型烟粉虱仅需要4代即替代了B型烟粉虱。在棉花植株上，杀虫剂的使用显著加快了Q型烟粉虱替代B型烟粉虱的速度。
而在黄瓜植株上，未用杀虫剂处理情况下，经过9代B型烟粉虱替代了Q型烟粉虱；但使用杀虫剂噻虫嗪和螺虫乙酯处理后，B型和Q型烟粉虱间的替代方向发生改变，经过5代后B型烟粉虱被Q型烟粉虱所替代，使用联苯菊酯处理时，仅需4代B型烟粉虱即被Q型烟粉虱替代。在黄瓜植株上，杀虫剂的使用完全改变了寄主植物对B型和Q型烟粉虱种群间的互作特性与替代方向，杀虫剂的使用导致了Q型烟粉虱替代B型烟粉虱。
显然，在Q型烟粉虱适宜的寄主植物上，杀虫剂的使用加速了Q型烟粉虱对B型烟粉虱的替代；在B型烟粉虱适宜的寄主植物上，杀虫剂的使用则逆转了B型烟粉虱对Q型烟粉虱的替代方向。毫无疑问，杀虫剂的使用在田间Q型烟粉虱替代B型烟粉虱的过程中起着关键性的作用。
四、烟粉虱体内共生菌的多样性与其对B型和Q型烟粉虱种群互作特性的影响
共生菌影响宿主昆虫的生长、发育与繁殖特性。本论文对2009年采集的61个烟粉虱田间种群（17个B型和44个Q型）和5种不同寄主植物饲养5年的B型烟粉虱实验室种群体内的原生共生菌Portiera和6种次生共生菌Hamiltonella、Arsenophonus、Cardinium、Wolbachia、Rickettsia和Fritschea进行了检测。结果表明，不同烟粉虱种群其体内共生菌的多样性与感染率呈现多样性特点，寄主植物种类、采集地点、烟粉虱生物型和性别这四个因子均显著影响其体内次生共生菌的多样性和感染率。田间B型和Q型烟粉虱共生菌的多样性和感染率与Q型烟粉虱替代B型烟粉虱没有明显的相关性。
进一步对3种不同寄主饲养6年的B型烟粉虱和噻虫嗪敏感和抗性B型烟粉虱体内共生菌密度研究后发现,寄主植物和杀虫剂是影响烟粉虱体内共生菌密度的重要因子。
五、番茄黄化曲叶病毒对B型和Q型烟粉虱竞争取代的影响
将携带和未携带TYLCV的B型和Q型烟粉虱分别接种在棉花植株上（TYLCV的非宿主植物），研究TYLCV对B型和Q型烟粉虱生长发育的直接作用。结果发现，携带TYLCV的B型烟粉虱较未携带TYLCV的B型烟粉虱在棉花上的存活率、产卵量均显著下降，雌成虫和雄成虫的体长明显变短，TYLCV对B型烟粉虱的生长发育呈现不利作用；而携带和未携带TYLCV的Q型烟粉虱在棉花上的生物学特性基本无差异，TYLCV对Q型烟粉虱的生长发育呈中性作用。
另外，将不带毒的B型和Q型烟粉虱分别接种在健康和感染TYLCV的番茄植株上，研究TYLCV对B型和Q型烟粉虱生长发育的间接作用。结果发现，在感染TYLCV的番茄植株上较健康的番茄植株上，Q型烟粉虱的发育历期缩短、寿命延长、存活率和产卵量提高及雌成虫体长变长，TYLCV显著促进了Q型烟粉虱的生长发育；而对B型烟粉虱，其作用刚好相反，在感染TYLCV的番茄植株上较健康的番茄植株上，B型烟粉虱的发育历期延长、寿命缩短、存活率和产卵量下降及雌成虫体长变短，TYLCV不利于B型烟粉虱的生长发育。
进一步将B型和Q型烟粉虱按各占50%比例分别接种到健康和感染TYLCV的番茄植株上，一定时间后随机采集寄主植物上一定数量的烟粉虱个体并鉴定其生物型，明确TYLCV对两种生物型烟粉虱种群间互作的影响。结果发现，未感染TYLCV的番茄上，B型烟粉虱经过5代替代了Q型烟粉虱，在感染TYLCV的番茄上，B型烟粉虱经过6代替代了Q型烟粉虱，TYLCV的感染不能逆转番茄植株上B型烟粉虱替代Q型烟粉虱的方向，但明显的减缓了B型烟粉虱替代Q型烟粉虱的速度，烟粉虱与TYLCV的互作一定意义上有助于田间Q型烟粉虱替代B型烟粉虱。
六、烟粉虱Q型替代B型促进中国TYLCV的暴发流行
对2009年采自我国18个省，55个烟粉虱种群（12个B型和43个Q型）的2750个烟粉虱个体进行TYLCV检测，结果发现，B型烟粉虱的带毒率为4.2%、Q型烟粉虱带毒率为24.4%，Q型烟粉虱的带毒率显著高于B型。
B型和Q型烟粉虱获取TYLCV的能力研究发现，相同时间内Q型烟粉虱获取病毒的量要显著的高于B型烟粉虱，B型烟粉虱需要48小时体内的病毒量才能达到饱和，而Q型烟粉虱仅需12小时体内的病毒量就能达到饱。
进一步的传毒特性研究发现，B型和Q型烟粉虱均不能垂直传播TYLCV，两者均仅能垂直传播TYLCV到卵和1龄若虫，而不能垂直传播到伪蛹和成虫。但两者均能水平传播TYLCV，Q型烟粉虱水平传播TYLCV的平均效率为52.2%，B型烟粉虱为10.5%，Q型烟粉虱水平传播TYLCV的能力显著高于B型烟粉虱。TYLCV在田间的暴发流行与Q型烟粉虱对B型烟粉虱的替代以及Q型烟粉虱水平传播TYLCV能力显著强于B型烟粉虱密切相关。
总之，本论文密切结合生产实际，针对重大外来入侵害虫—B型和Q型烟粉虱及其传播的TYLCV，通过系统监测，首次阐明了烟粉虱入侵我国十年来，田间危害优势生物型的变化动态；率先揭示了Q型烟粉虱替代B型烟粉虱的生态学机制和近年来TYLCV在我国暴发流行的机制。
Biological invasions endanger national security, ecological security, and economic security.Bemisia tabaci, which has been regarded as a species complex consisting of many biotypes, is aninvasive and destructive pest of numerous protected and field crops worldwide. Among the B.tabaci biotypes, the B biotype and Q biotype are two damaging and invasive crop pests. In themid-1990s, the B biotype was introduced into China and caused tremendous losses to agriculturalproduction and the national economy. After the Q biotype was first detected in Yunnan Provincein2003, it spread quickly into East China and North China. The invasion and spread of the Qbiotype was accompanied by outbreaks of a B. tabaci-transmitted virus called Tomato yellow leafcurl virus (TYLCV). In this dissertation, a systematic monitoring of B. tabaci on many crops inmore than20provinces in China was conducted. The results indicated that over the last10yearsthe Q biotype has been displacing the previously well-established B biotype as the predominantbiotype in China. Studies on the influence of biotic factors (host plant, plant virus, and symbioticbacteria) and an abiotic factor (insecticide application) on the interactions between B and Qbiotypes showed that insecticide application is the key ecological factor contributing to thedisplacement of the B biotype by the Q biotype. Controlled laboratory experiments wereconducted to compare the virus acquisition capability of the two biotypes and to investigate themode of viral transmission (horizontal and vertical). The results showed that the displacement ofthe B biotype by the Q biotype contributed to TYLCV epidemics in China. The research resultswill help guide the sustainable control of B. tabaci and TYLCV.
The main research contents and conclusions are as follows:
1. The distribution and population dynamics of B and Q biotypes in China
PCR primers specific for mtCO1of biotypes B and Q and gene sequencing were used in anextensive survey (covering19provinces in2009and27provinces in2011) to determine thebiotype status of B. tabaci in China. The results in2009showed that the Q biotype was dominantin44locations while B biotype was dominant in17locations and that the Q biotype was mainlydistributed in the Yangtze River and the eastern coastal areas. Among98populations examined in2011,57were pure Q biotype,12were pure B biotype, and29were mixtures of B and Q biotypes(with the Q biotype representing>50%of the individuals in20of the29); the Q biotype waspredominant across China except in southeast provinces. Overall, the results showed that over thelast10years, the main damaging biotype of B. tabaci in China has changed from the B biotype tothe Q biotype.
2. The impact of host plant on the competitive displacement between B and Q biotypes
To explore the impact of host plant on the competitive displacement between B and Qbiotypes, equal numbers of B and Q biotypes were added to the same poinsettia, pepper, tomato,and cabbage plants, and the biotypes of individuals in the developing populations wereperiodically determined. The results showed that the Q biotype could displace the B biotype on poinsettia and pepper after8and2successive generations, respectively. In contrast, the B biotypecould displace the Q biotype on tomato and cabbage after5and4successive generations,respectively. Different host plants had different effects on the interactions between B and Qbiotypes, resulting in the differences in the speed and direction of displacement of one biotype bythe other.
3. The impact of insecticide on the competitive displacement between B and Q biotypes
To explore the impact of insecticide on the competitive displacement between B and Qbiotypes, equal numbers of both B and Q biotypes were added to the same cotton and cucumberplants. The plants were either treated or not treated with an insecticide, and the biotypes ofindividuals in the developing populations were periodically determined. The results showed thatthe Q biotype displaced the B biotype on cotton after6successive generations without insecticidebut after only4successive generations with insecticide (Thiamethoxam). On cotton, the use ofinsecticide significantly increased the rate at which the Q biotype displaced the B biotype.
Although the B biotype displaced the Q biotype on cucumber after9successive generationswithout insecticide, the Q biotype replaced the B biotype when cucumber was treated withinsecticide. The Q biotype replaced the B biotype after5successive generations with theinsecticide Movento or Thiamethoxam, and after4successive generations with the insecticideBifenthrin. On cucumber, application of insecticide reversed the impact of host plant on theinteractions between B and Q biotypes, resulting in the displacement of the B biotype by the Qbiotype.
Overall, the use of insecticide enhanced displacement of the B biotype by the Q biotype inone of two ways depending on the host: if the untreated host plant was more suitable to the Qbiotype (as is cotton), an insecticide increased the rate at which the Q biotype displaced the Bbiotype; if the untreated host plant was more suitable to the B biotype (as is cucumber), aninsecticide reversed the direction of displacement. These results demonstrate that insecticideapplication plays a key role in the displacement of the B biotype by the Q biotype in the field.
4. Diversity of symbionts in Bemisia tabaci and its impact on the interactions between Band Q biotypes
Symbiotic bacteria can influence the growth, development, and reproduction of insect hosts.In this study, gene-specific PCR was used to screen for the presence of the primary symbiontPortiera and the six secondary symbionts (Hamiltonella, Arsenophonus, Cardinium, Wolbachia,Rickettsia, and Fritschea) in61field populations (17B and44Q biotypes) collected fromdifferent plant species and locations in China and in5laboratory populations that had similargenetic background but that were cultured on5different host plant species for5years. The resultsindicated that the diversity and infection frequency of secondary symbionts in B. tabaci wasinfluenced by B. tabaci biotype, B. tabaci sex, host plant, and geographical location. Nocorrelation was detected between the diversity and infection frequency of symbionts in B. tabaci and the displacement of the B biotype by the Q biotype.
An additional experiment was conducted to investigate the infection densities of symbionts in3laboratory populations that had similar genetic background but that were cultured on3differenthost plant species for6years of the B biotype and of a Thiamethoxam-resistant and-susceptible Bbiotype population. Overall, the results revealed that host plant and insecticide affected theinfection densities of symbionts in B. tabaci.
5. The impact of TYLCV on the competitive displacement between B and Q biotypes
The direct effect of TYLCV on the growth and development of B. tabaci was investigated bycomparing the performance of viruliferous and non-viruliferous B and Q biotypeson cotton (anon-host for TYLCV). The results showed that TYLCV significantly decreased the survival,fecundity, and adult body size of viruliferous individuals of the B biotype but not of the Q biotypefeeding on cotton.
The indirect effect of TYLCV on the growth and development of B. tabaci was investigatedby comparing the performance of non-viruliferous B and Q biotypes on healthy andTYLCV-infected tomato plants. The results showed that TYLCV-infected tomato plants improvedthe performance of the Q biotype but decreased the performance of the B biotype with respect tothe fecundity, survival rate, development time, and female body size.
The impact of TYLCV on the competitive displacement between B and Q biotypes wasfurther examined by adding equal numbers of both B and Q biotypes to the same healthy andTYLCV-infected tomato plants, respectively and periodically determining the biotypes ofindividuals in the developing populations. The results showed that the B biotype displaced the Qbiotype after5successive generations on healthy tomato plants but after6successive generationson TYLCV-infected tomato plants. Although TYLCV infection did not prevent the displacementof the Q biotype by the B biotype, it significantly delayed the rate of displacement. The overallresults indicate that the mutualistic relationship between TYLCV and the Q biotype contributes tothe displacement of the B biotype by the Q biotype in China.
6. The displacement of B biotype by Q biotype contribute to the epidemics of TYLCV inChina
A nation-wide field survey of18provinces that included55B. tabaci populations (12B and43Q biotypes) and2750individuals was conducted in2009to investigate the infection frequencyof TYLCV. The results showed that the percentage of TYLCV-infected individuals wassignificantly lower among B biotype (4.2%) than Q biotype (24.4%) individuals.
Virus acquisition experiments indicated that the relative viral gene expression was greater inthe Q biotype than in the B biotype after each of five acquisition access periods (AAPs) and thatthe B and Q biotypes required AAPs of48and12h, respectively, to reach their maximum viralloads.
Transmission experiments demonstrated that TYLCV was not transmitted vertically, i.e., was not transmitted to the next generation by the B and Q biotypes. TYLCV was detected in eggs andcrawlers but was not detected in the pupa or adult progeny from viruliferous B or Q whiteflies.TYLCV could be transmitted horizontally by both B and Q biotypes, and the overall percentage ofviruliferous individuals that transmitted TYLCV was52.2%for the Q biotype but only10.5%forthe B biotype, indicating that horizontal transmission of TYLCV was greater with the Q biotype.The epidemics of TYLCV are related to the displacement the B biotype by the Q biotype and thegreater horizontal transmission of TYLCV by the Q biotype than the B biotype.
In summary, in this dissertation, we found that the predominant biotype of B. tabaci haschanged during the past decade, the ecological mechanisms of displacement of B biotype by Qbiotype and the outbreaks of TYLCV in China were fist revealed.
主要研究内容和结论如下：
一、B型烟粉虱和Q型烟粉虱在我国的入侵分布动态
利用特异分子标记并结合mtCO1基因序列分析，分别于2009年和2011年对采自我国的19个省市和27个省市的烟粉虱种群的生物型进行了鉴定，结果发现，2009年的61个烟粉虱种群中有44个是Q型烟粉虱种群，17个是B型烟粉虱种群，Q型烟粉虱主要分布在长江流域及东部沿海地区；2011年的98个烟粉虱种群中有60个Q型烟粉虱种群，9个B型烟粉虱种群，29个混合种群中有20个种群Q型烟粉虱占有较高的比例（>50%），除了东南省份，Q型烟粉虱都占据主导优势。监测结果发现，烟粉虱入侵我国十多年后，其主要危害生物型已由早期入侵B型转变为Q型。
二、寄主植物对B型和Q型烟粉虱种群竞争取代的影响
将B型和Q型烟粉虱按各占50%的比例分别接种到一品红、辣椒、番茄和甘蓝四种寄主植物上，一定时间后随机采集寄主植物上一定数量的烟粉虱个体并鉴定其生物型，明确寄主植物对两种生物型烟粉虱种群间互作的影响。结果发现在一品红和辣椒植株上，分别经过8代和2代，Q型烟粉虱替代了B型烟粉虱；而在番茄和甘蓝植株上则刚好相反，分别经过5代和4代，B型烟粉虱替代了Q型烟粉虱。不同寄主植物对B型和Q型烟粉虱种群间的互作表现了不同的特性，导致了不同的替代方向和速度。
三、杀虫剂对B型和Q型烟粉虱种群竞争取代的影响
同样将B型和Q型烟粉虱按各占50%比例分别接种到棉花和黄瓜植物上，并分别设置一定浓度的杀虫剂处理和未处理的重复，一定时间后随机采集寄主植物上一定数量的烟粉虱个体并鉴定其生物型，明确杀虫药剂对两种生物型烟粉虱种群间互作的影响。结果发现，在未经杀虫剂处理的棉花植株上，经过6代Q型烟粉虱替代了B型烟粉虱，而在经一定剂量的噻虫嗪处理后的棉花植株上，Q型烟粉虱仅需要4代即替代了B型烟粉虱。在棉花植株上，杀虫剂的使用显著加快了Q型烟粉虱替代B型烟粉虱的速度。
而在黄瓜植株上，未用杀虫剂处理情况下，经过9代B型烟粉虱替代了Q型烟粉虱；但使用杀虫剂噻虫嗪和螺虫乙酯处理后，B型和Q型烟粉虱间的替代方向发生改变，经过5代后B型烟粉虱被Q型烟粉虱所替代，使用联苯菊酯处理时，仅需4代B型烟粉虱即被Q型烟粉虱替代。在黄瓜植株上，杀虫剂的使用完全改变了寄主植物对B型和Q型烟粉虱种群间的互作特性与替代方向，杀虫剂的使用导致了Q型烟粉虱替代B型烟粉虱。
显然，在Q型烟粉虱适宜的寄主植物上，杀虫剂的使用加速了Q型烟粉虱对B型烟粉虱的替代；在B型烟粉虱适宜的寄主植物上，杀虫剂的使用则逆转了B型烟粉虱对Q型烟粉虱的替代方向。毫无疑问，杀虫剂的使用在田间Q型烟粉虱替代B型烟粉虱的过程中起着关键性的作用。
四、烟粉虱体内共生菌的多样性与其对B型和Q型烟粉虱种群互作特性的影响
共生菌影响宿主昆虫的生长、发育与繁殖特性。本论文对2009年采集的61个烟粉虱田间种群（17个B型和44个Q型）和5种不同寄主植物饲养5年的B型烟粉虱实验室种群体内的原生共生菌Portiera和6种次生共生菌Hamiltonella、Arsenophonus、Cardinium、Wolbachia、Rickettsia和Fritschea进行了检测。结果表明，不同烟粉虱种群其体内共生菌的多样性与感染率呈现多样性特点，寄主植物种类、采集地点、烟粉虱生物型和性别这四个因子均显著影响其体内次生共生菌的多样性和感染率。田间B型和Q型烟粉虱共生菌的多样性和感染率与Q型烟粉虱替代B型烟粉虱没有明显的相关性。
进一步对3种不同寄主饲养6年的B型烟粉虱和噻虫嗪敏感和抗性B型烟粉虱体内共生菌密度研究后发现,寄主植物和杀虫剂是影响烟粉虱体内共生菌密度的重要因子。
五、番茄黄化曲叶病毒对B型和Q型烟粉虱竞争取代的影响
将携带和未携带TYLCV的B型和Q型烟粉虱分别接种在棉花植株上（TYLCV的非宿主植物），研究TYLCV对B型和Q型烟粉虱生长发育的直接作用。结果发现，携带TYLCV的B型烟粉虱较未携带TYLCV的B型烟粉虱在棉花上的存活率、产卵量均显著下降，雌成虫和雄成虫的体长明显变短，TYLCV对B型烟粉虱的生长发育呈现不利作用；而携带和未携带TYLCV的Q型烟粉虱在棉花上的生物学特性基本无差异，TYLCV对Q型烟粉虱的生长发育呈中性作用。
另外，将不带毒的B型和Q型烟粉虱分别接种在健康和感染TYLCV的番茄植株上，研究TYLCV对B型和Q型烟粉虱生长发育的间接作用。结果发现，在感染TYLCV的番茄植株上较健康的番茄植株上，Q型烟粉虱的发育历期缩短、寿命延长、存活率和产卵量提高及雌成虫体长变长，TYLCV显著促进了Q型烟粉虱的生长发育；而对B型烟粉虱，其作用刚好相反，在感染TYLCV的番茄植株上较健康的番茄植株上，B型烟粉虱的发育历期延长、寿命缩短、存活率和产卵量下降及雌成虫体长变短，TYLCV不利于B型烟粉虱的生长发育。
进一步将B型和Q型烟粉虱按各占50%比例分别接种到健康和感染TYLCV的番茄植株上，一定时间后随机采集寄主植物上一定数量的烟粉虱个体并鉴定其生物型，明确TYLCV对两种生物型烟粉虱种群间互作的影响。结果发现，未感染TYLCV的番茄上，B型烟粉虱经过5代替代了Q型烟粉虱，在感染TYLCV的番茄上，B型烟粉虱经过6代替代了Q型烟粉虱，TYLCV的感染不能逆转番茄植株上B型烟粉虱替代Q型烟粉虱的方向，但明显的减缓了B型烟粉虱替代Q型烟粉虱的速度，烟粉虱与TYLCV的互作一定意义上有助于田间Q型烟粉虱替代B型烟粉虱。
六、烟粉虱Q型替代B型促进中国TYLCV的暴发流行
对2009年采自我国18个省，55个烟粉虱种群（12个B型和43个Q型）的2750个烟粉虱个体进行TYLCV检测，结果发现，B型烟粉虱的带毒率为4.2%、Q型烟粉虱带毒率为24.4%，Q型烟粉虱的带毒率显著高于B型。
B型和Q型烟粉虱获取TYLCV的能力研究发现，相同时间内Q型烟粉虱获取病毒的量要显著的高于B型烟粉虱，B型烟粉虱需要48小时体内的病毒量才能达到饱和，而Q型烟粉虱仅需12小时体内的病毒量就能达到饱。
进一步的传毒特性研究发现，B型和Q型烟粉虱均不能垂直传播TYLCV，两者均仅能垂直传播TYLCV到卵和1龄若虫，而不能垂直传播到伪蛹和成虫。但两者均能水平传播TYLCV，Q型烟粉虱水平传播TYLCV的平均效率为52.2%，B型烟粉虱为10.5%，Q型烟粉虱水平传播TYLCV的能力显著高于B型烟粉虱。TYLCV在田间的暴发流行与Q型烟粉虱对B型烟粉虱的替代以及Q型烟粉虱水平传播TYLCV能力显著强于B型烟粉虱密切相关。
总之，本论文密切结合生产实际，针对重大外来入侵害虫—B型和Q型烟粉虱及其传播的TYLCV，通过系统监测，首次阐明了烟粉虱入侵我国十年来，田间危害优势生物型的变化动态；率先揭示了Q型烟粉虱替代B型烟粉虱的生态学机制和近年来TYLCV在我国暴发流行的机制。
Biological invasions endanger national security, ecological security, and economic security.Bemisia tabaci, which has been regarded as a species complex consisting of many biotypes, is aninvasive and destructive pest of numerous protected and field crops worldwide. Among the B.tabaci biotypes, the B biotype and Q biotype are two damaging and invasive crop pests. In themid-1990s, the B biotype was introduced into China and caused tremendous losses to agriculturalproduction and the national economy. After the Q biotype was first detected in Yunnan Provincein2003, it spread quickly into East China and North China. The invasion and spread of the Qbiotype was accompanied by outbreaks of a B. tabaci-transmitted virus called Tomato yellow leafcurl virus (TYLCV). In this dissertation, a systematic monitoring of B. tabaci on many crops inmore than20provinces in China was conducted. The results indicated that over the last10yearsthe Q biotype has been displacing the previously well-established B biotype as the predominantbiotype in China. Studies on the influence of biotic factors (host plant, plant virus, and symbioticbacteria) and an abiotic factor (insecticide application) on the interactions between B and Qbiotypes showed that insecticide application is the key ecological factor contributing to thedisplacement of the B biotype by the Q biotype. Controlled laboratory experiments wereconducted to compare the virus acquisition capability of the two biotypes and to investigate themode of viral transmission (horizontal and vertical). The results showed that the displacement ofthe B biotype by the Q biotype contributed to TYLCV epidemics in China. The research resultswill help guide the sustainable control of B. tabaci and TYLCV.
The main research contents and conclusions are as follows:
1. The distribution and population dynamics of B and Q biotypes in China
PCR primers specific for mtCO1of biotypes B and Q and gene sequencing were used in anextensive survey (covering19provinces in2009and27provinces in2011) to determine thebiotype status of B. tabaci in China. The results in2009showed that the Q biotype was dominantin44locations while B biotype was dominant in17locations and that the Q biotype was mainlydistributed in the Yangtze River and the eastern coastal areas. Among98populations examined in2011,57were pure Q biotype,12were pure B biotype, and29were mixtures of B and Q biotypes(with the Q biotype representing>50%of the individuals in20of the29); the Q biotype waspredominant across China except in southeast provinces. Overall, the results showed that over thelast10years, the main damaging biotype of B. tabaci in China has changed from the B biotype tothe Q biotype.
2. The impact of host plant on the competitive displacement between B and Q biotypes
To explore the impact of host plant on the competitive displacement between B and Qbiotypes, equal numbers of B and Q biotypes were added to the same poinsettia, pepper, tomato,and cabbage plants, and the biotypes of individuals in the developing populations wereperiodically determined. The results showed that the Q biotype could displace the B biotype on poinsettia and pepper after8and2successive generations, respectively. In contrast, the B biotypecould displace the Q biotype on tomato and cabbage after5and4successive generations,respectively. Different host plants had different effects on the interactions between B and Qbiotypes, resulting in the differences in the speed and direction of displacement of one biotype bythe other.
3. The impact of insecticide on the competitive displacement between B and Q biotypes
To explore the impact of insecticide on the competitive displacement between B and Qbiotypes, equal numbers of both B and Q biotypes were added to the same cotton and cucumberplants. The plants were either treated or not treated with an insecticide, and the biotypes ofindividuals in the developing populations were periodically determined. The results showed thatthe Q biotype displaced the B biotype on cotton after6successive generations without insecticidebut after only4successive generations with insecticide (Thiamethoxam). On cotton, the use ofinsecticide significantly increased the rate at which the Q biotype displaced the B biotype.
Although the B biotype displaced the Q biotype on cucumber after9successive generationswithout insecticide, the Q biotype replaced the B biotype when cucumber was treated withinsecticide. The Q biotype replaced the B biotype after5successive generations with theinsecticide Movento or Thiamethoxam, and after4successive generations with the insecticideBifenthrin. On cucumber, application of insecticide reversed the impact of host plant on theinteractions between B and Q biotypes, resulting in the displacement of the B biotype by the Qbiotype.
Overall, the use of insecticide enhanced displacement of the B biotype by the Q biotype inone of two ways depending on the host: if the untreated host plant was more suitable to the Qbiotype (as is cotton), an insecticide increased the rate at which the Q biotype displaced the Bbiotype; if the untreated host plant was more suitable to the B biotype (as is cucumber), aninsecticide reversed the direction of displacement. These results demonstrate that insecticideapplication plays a key role in the displacement of the B biotype by the Q biotype in the field.
4. Diversity of symbionts in Bemisia tabaci and its impact on the interactions between Band Q biotypes
Symbiotic bacteria can influence the growth, development, and reproduction of insect hosts.In this study, gene-specific PCR was used to screen for the presence of the primary symbiontPortiera and the six secondary symbionts (Hamiltonella, Arsenophonus, Cardinium, Wolbachia,Rickettsia, and Fritschea) in61field populations (17B and44Q biotypes) collected fromdifferent plant species and locations in China and in5laboratory populations that had similargenetic background but that were cultured on5different host plant species for5years. The resultsindicated that the diversity and infection frequency of secondary symbionts in B. tabaci wasinfluenced by B. tabaci biotype, B. tabaci sex, host plant, and geographical location. Nocorrelation was detected between the diversity and infection frequency of symbionts in B. tabaci and the displacement of the B biotype by the Q biotype.
An additional experiment was conducted to investigate the infection densities of symbionts in3laboratory populations that had similar genetic background but that were cultured on3differenthost plant species for6years of the B biotype and of a Thiamethoxam-resistant and-susceptible Bbiotype population. Overall, the results revealed that host plant and insecticide affected theinfection densities of symbionts in B. tabaci.
5. The impact of TYLCV on the competitive displacement between B and Q biotypes
The direct effect of TYLCV on the growth and development of B. tabaci was investigated bycomparing the performance of viruliferous and non-viruliferous B and Q biotypeson cotton (anon-host for TYLCV). The results showed that TYLCV significantly decreased the survival,fecundity, and adult body size of viruliferous individuals of the B biotype but not of the Q biotypefeeding on cotton.
The indirect effect of TYLCV on the growth and development of B. tabaci was investigatedby comparing the performance of non-viruliferous B and Q biotypes on healthy andTYLCV-infected tomato plants. The results showed that TYLCV-infected tomato plants improvedthe performance of the Q biotype but decreased the performance of the B biotype with respect tothe fecundity, survival rate, development time, and female body size.
The impact of TYLCV on the competitive displacement between B and Q biotypes wasfurther examined by adding equal numbers of both B and Q biotypes to the same healthy andTYLCV-infected tomato plants, respectively and periodically determining the biotypes ofindividuals in the developing populations. The results showed that the B biotype displaced the Qbiotype after5successive generations on healthy tomato plants but after6successive generationson TYLCV-infected tomato plants. Although TYLCV infection did not prevent the displacementof the Q biotype by the B biotype, it significantly delayed the rate of displacement. The overallresults indicate that the mutualistic relationship between TYLCV and the Q biotype contributes tothe displacement of the B biotype by the Q biotype in China.
6. The displacement of B biotype by Q biotype contribute to the epidemics of TYLCV inChina
A nation-wide field survey of18provinces that included55B. tabaci populations (12B and43Q biotypes) and2750individuals was conducted in2009to investigate the infection frequencyof TYLCV. The results showed that the percentage of TYLCV-infected individuals wassignificantly lower among B biotype (4.2%) than Q biotype (24.4%) individuals.
Virus acquisition experiments indicated that the relative viral gene expression was greater inthe Q biotype than in the B biotype after each of five acquisition access periods (AAPs) and thatthe B and Q biotypes required AAPs of48and12h, respectively, to reach their maximum viralloads.
Transmission experiments demonstrated that TYLCV was not transmitted vertically, i.e., was not transmitted to the next generation by the B and Q biotypes. TYLCV was detected in eggs andcrawlers but was not detected in the pupa or adult progeny from viruliferous B or Q whiteflies.TYLCV could be transmitted horizontally by both B and Q biotypes, and the overall percentage ofviruliferous individuals that transmitted TYLCV was52.2%for the Q biotype but only10.5%forthe B biotype, indicating that horizontal transmission of TYLCV was greater with the Q biotype.The epidemics of TYLCV are related to the displacement the B biotype by the Q biotype and thegreater horizontal transmission of TYLCV by the Q biotype than the B biotype.
In summary, in this dissertation, we found that the predominant biotype of B. tabaci haschanged during the past decade, the ecological mechanisms of displacement of B biotype by Qbiotype and the outbreaks of TYLCV in China were fist revealed.
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