Resistance of soybean genotypes to Bemisia tabaci (Genn.) Biotype B (Hemiptera: Aleyrodidae)

The silverleaf whitefly Bemisia tabaci (Genn.) biotype B has become a serious problem for soybean cultivation because it can significantly reduce soybean productivity. The use of soybean cultivars resistant to whitefly attack is an important strategy in an integrated pest management (IPM) program. This study evaluated the preference for oviposition and colonization by B. tabaci biotype B on different soybean genotypes. In the free-choice test, the genotypes studied were 'IAC 17' and 'IAC 19' as the standards for resistance and 'IAC Holambra Stwart' as the standard for susceptibility, as well as BABR01-0492, BABR01-0173, BABR01-1259, BABR01-1576, BABR99-4021HC, BABR99-4021HP, 'Barreiras', 'Conquista', 'Corisco', 'BRS Gralha', PI274454, PI227687, and PI171451. In the no-choice test, the four best genotypes selected in the free-choice test, in addition to the susceptible and resistant standards were evaluated. Our data indicated 'Barreiras' as the most resistant genotype against B. tabaci biotype B. 'BRS Gralha', which was the least attractive to whitefly adults in the free-choice test, did not show resistance to insect attack when they were confined in cages in the no-choice test. Despite the high number of eggs observed, BABR01-1576 and BABR99-4021HC showed a reduced number of nymphs, indicating antibiosis. The genotypes with a high level of resistance can be used as a tool against B. tabaci in IPM or as a source of resistance in plant-breeding programs.     

Factors affecting oviposition of Bemisia tabaci (Genn.) biotype B in sweet pepper

Bemisia tabaci (Gen.) biotype B is considered a pest of economical importance for several vegetables. The oviposition behaviour of the whilefly was evaluated in sweet pepper plants. The trials were carried out under greenhouse condition and in the Laboratory of Entomology of DEFERS/UNESP, Campus of Ilha Solteira-SP, with the sweet pepper Magali-R genotype. The effect of plant age on the whitefly oviposition was evaluated in free-choice tests, in plants, 25, 30, 35, 40 and 45-day-old, as egg distribution in the plant and on the leaf blade was evaluated in 35-days-old plants. In a no-choice tests, 35-day-old plants were used to evaluate the effect of the densities of 50, 100, 150, 200 and 250 adults per plant on the number of eggs laid by insects. The silverleaf whitefly preferred to oviposit on the third to sixth leaflets, of the medium and superior part of plants of sweet pepper; the leaf blade areas, located in the lobes right and left close the base of the leaf were the preferential site for whitefly oviposition. Older plants, 40- and 45-day-old, were preferentially used for oviposition, and 200 and 250 adults per plant were both enough to lay a number of eggs that allowed to differentiate among sweet pepper genotypes with different whitefly resistance levels.     

Biological aspects of Bemisia tabaci (Genn.) B biotype (Hemiptera: Aleyrodidae) on six bean genotypes

The silverleaf whitefly is one of the most harmful pests that attack bean crops, mainly for extracting large quantities of phloem sap and transmitting the bean golden mosaic virus. Resistant germoplasm plants can be an important method for controlling this pest. The biological aspects of Bemisia tabaci B biotype on bean (Phaseolus vulgaris) genotypes were evaluated. The tests were conducted under laboratory conditions, with the following genotypes: Arc 1, Arc 3s, Arc 5s, G13028, G11056 and Porrillo 70. The bean plants in a stage IV-1 were infested during one day with silverleaf whiteflies. Afterwards the eggs and nymphs were observed until adult emergence. Longevity and fecundity of emerged insects were also evaluated. The longest development time occurred for nymphs fed on Arc 3s genotype (26.5 days), following by G11056 (25.9 days) and G13028 (25.3 days). The development period was 5.5 days longer in Arc 3s when compared with Porrillo 70. Also, the wild genotypes Arc 3s and G11056 showed higher mortality rates (94.7% and 83.1%, respectively), which may suggest antibiosis and/or feeding nonpreference resistance type. For this reason, although longevity and fecundity were not influenced when the whitefly fed on resistant genotypes (Arc 3s, G11056, G13028 and Arc 5s), those genotypes can be used for bean breeding program towards B. tabaci B biotype.     

Establishment of a silverleaf whitefly, Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae) population in western Sydney

A field survey in western Sydney found populations of Bemisia tabaci biotype B (silverleaf whitefly) that have the potential to damage greenhouse vegetable crops prevalent in the region. Adult whiteflies were collected from five farms at 2-week intervals and the proportion of silverleaf whitefly in the overall whitefly population was determined using polyacrylamide gel electrophoresis. High levels of silverleaf whitefly were detected at most sampling dates on two of the farms surveyed and a third farm exhibited lower but relatively continuous silverleaf whitefly abundance. The presence of permanent silverleaf whitefly populations must be considered by the greenhouse industry when formulating whitefly management plans for western Sydney.     

豇豆轻斑驳病毒CPMMV感染菜豆对B型烟粉虱寄主选择和生物学特性的影响

豇豆轻斑驳病毒（Cowpea mild mottle virus,CPMMV）的流行与其媒介昆虫烟粉虱Bemisia tabaci密切相关,但CPMMV感染寄主植物对B型烟粉虱寄主选择行为和生物学特性的影响尚不清楚。本研究探究了1）CPMMV感染菜豆Phaseolus vulgaris不同时间对B型烟粉虱寄主选择和产卵偏好性的影响;2）CPMMV感染菜豆对B型烟粉虱生物学特性的影响。结果发现：1）相较于健康菜豆,摩擦接种CPMMV后1周,B型烟粉虱却偏好在CPMMV感染的植株上停留和产卵;而摩擦接种后4周,B型烟粉虱却偏好在健康的植株上停留和产卵;2）无论是在摩擦接种后1周和4周的CPMMV感染植株上,B型烟粉虱在健康和CPMMV感染植株上的产卵量和存活率差异不明显。可见,CPMMV感染菜豆对于B型烟粉虱的寄主选择偏好性具有时间效应,而CPMMV感染菜豆对于B型烟粉虱的生物学特性没有显著影响。     

Differential indirect effects of two plant viruses on an invasive and an indigenous whitefly vector: implications for competitive displacement

The role of vector–begomovirus–plant interactions in the widespread invasion by some members of the whitefly species complex Bemisia tabaci is poorly understood. The invasive B biotype of B. tabaci entered China in the late 1990s and had become the predominant or only biotype of the whitefly in many regions of the country by 2005–2006. Meanwhile epidemics of begomoviruses have been observed in many crops including tomato for which Tomato yellow leaf curl China virus (TYLCCNV) and Tomato yellow leaf curl virus (TYLCV) have been identified as two major disease-causing agents. Here, we conducted laboratory experiments to compare the performance of the invasive B and indigenous ZHJ1 whitefly biotypes on uninfected, TYLCCNV-infected and TYLCV-infected plants of tomato cv. Hezuo903, a cultivar that has been widely cultivated in many regions of China. The infection of tomato plants by either of the viruses had no or only marginal effects on the development, survival and fecundity of the B biotype. In contrast, survival and fecundity of the ZHJ1 biotype were significantly reduced on virus-infected plants compared to those on uninfected plants. Populations of the B biotype on uninfected and TYLCCNV-infected plants increased at similar rates, whereas population increase of the ZHJ1 biotype on TYLCCNV-infected plants was affected adversely. These asymmetric responses to virus infection of tomato plants between the B and ZHJ1 biotypes are likely to offer advantages to the B biotype in its invasion and displacement of the indigenous biotype.     

Development of silverleaf assay, protein and nucleic acid-based diagnostic techniques for the quick and reliable detection and monitoring of biotype B of the whitefly, Bemisia tabaci (Gennadius)

The aim of this study was to develop and optimize silverleaf bioassay, esterase analysis and PCR-based techniques to distinguish quickly and reliably biotype B of the whitefly, Bemisia tabaci (Gennadius), from Indian indigenous biotypes. Zucchini and squash readily develop silverleaf symptoms upon feeding by the B biotype, but they are not readily available in Indian markets. A local pumpkin variety 'Big' was, therefore, used in silverleaf assay, which developed symptoms similar to those on zucchini and squash and can be used reliably to detect B biotype. Analysis of non-specific esterases of B and the indigenous biotypes indicated both quantitative and qualitative differences in esterase patterns. Two high molecular weight bands were unique to B biotype and they occurred in abundance. These esterases were used to develop quick and field-based novel detection methods for differentiating B from the indigenous biotypes. Development of these simple and cost-effective protocols has wider application as they can be potentially used to identify other agricultural pests. Mitochondrial cytochrome oxidase I gene sequences and randomly amplified polymorphic DNA (RAPD) polymorphisms, generated using the primer OpB11, were also found useful for detecting B. tabaci biotypes. A B biotype-specific RAPD band of 800 bp was sequenced, which was used to a develop sequence characterized amplified region (SCAR) marker. The SCAR marker involved the development of B biotype-specific primers that amplified 550 bp PCR products only from B biotype genomic DNA. Silverleaf assay, esterases, RAPDs or a SCAR marker were used in combination to analyse whitefly samples collected from selected locations in India, and it was found that any of these techniques can be used singly or in combination to detect B biotype reliably. The B biotype was found in southern parts of India but not in the north in 2004-06.     

喷施吲哚-3-乙酸 (IAA) 对番茄及其上B型烟粉虱的影响

【目的】不同植物生长调节物质对植食性昆虫产卵、发育、取食等常有促进或抑制作用,进而影响植物与昆虫的互作关系。本研究旨在阐明常用的植物生长调节剂吲哚-3-乙酸对B型烟粉虱Bemisia tabaci (Gennadius)的影响,进而探索防治烟粉虱环境友好的方法,同时期望获得更加优质的烟粉虱用作替代寄主饲养寄生蜂。【方法】对番茄 Solanum lycopersicum L.植株外源喷施吲哚-3-乙酸(IAA),检测其生理生化的响应;记录B型烟粉虱存活率、产卵量并测量解毒酶等指标。【结果】喷施IAA后番茄叶片含水量略有上升,叶面积增大,叶绿素含量增加,过氧化物酶(POD)活性降低,多酚氧化酶(PPO)活性升高,超氧化物歧化酶(SOD)活性变化不显著(P＞0.05)。喷施IAA后番茄上B型烟粉虱成虫存活率高于对照组,其中50 mg/L IAA番茄上B型烟粉虱成虫存活率和总产卵量最高,250 mg/L IAA存活率最低;喷施IAA 120 h后烟粉虱总产卵量升高;碱性磷酸酶(AKP)、超氧化物歧化酶(SOD)活性较对照升高。【结论】喷施不同浓度的IAA溶液后,番茄植株的生理生化水平发生不同程度的改变,烟粉虱成虫的存活状况、产卵量以及体内解毒活性也会产生不同程度的改变。适宜剂量的IAA处理能够使烟粉虱的生存表现更加优异,从而为饲养寄生蜂提供优质的替代寄主。     

不同作物田烟粉虱发生的时空动态

2008～2009年连续2年系统调查了番茄、茄子、棉花、大豆、玉米等寄主植物上烟粉虱种群发生的时间与空间动态。结果表明：不同寄主植物上的烟粉虱成虫及其伪蛹数量有显著性差异,其密度大小依次为：茄子>棉花>番茄>大豆>玉米。其中,在玉米上除了发现极少量的成虫逗留外,没有发现烟粉虱的卵及若虫。在发生的时间序列上,烟粉虱成虫及伪蛹的数量呈现为先逐渐上升后又下降的变化过程,发生高峰期集中在8月5日到8月31日,9月初以后烟粉虱数量慢慢减少。在空间分布上,表现为烟粉虱成虫喜食寄主的上部叶片。统计分析显示,寄主对烟粉虱成虫和伪蛹的数量的影响极显著,而年份对其数量的影响没有显著差异。由此得出的烟粉虱发生和达到高峰的时间,可为烟粉虱预测预报和区域性综合治理提供重要理论依据。     

华东、华南和西南地区烟粉虱的种群及寄主分化

烟粉虱Bemisia tabaci是一个复合种,它具有的生物型分化、较强的传播病毒的能力和抗药性、较快的繁殖速率等特征使其成为我国农业生产中重要害虫之一。本研究利用细胞色素线粒体氧化酶Ⅰ基因,对采集自江苏、广东和云南三省的烟粉虱样本进行了生物型鉴定,并对烟粉虱生物型与寄主植物之间的关联性开展了调查。结果表明,在广东和云南省,都存在未鉴定的土著种群与入侵的B型、Q型共存的现象;同时,在本研究中广东省尚未采集到Q型烟粉虱,而在江苏采集到的粉虱样本全部为入侵型。研究结果还表明,相对于入侵种而言,土著种群显示出更强的寄主植物趋同性;丰富的寄主植物以及本身具有的多食性特性有助于B型、Q型等生物型在世界各地的广泛入侵。     

Influence of trichomes on attractiveness and ovipositional preference of Bemisia tabaci (Genn.) B biotype (Hemiptera: Aleyrodidae) on tomato genotypes

Brazil is one of the world's largest tomato producer, but considerable part of the production is lost due to the attack of Bemisia tabaci (Genn.) B biotype. Resistant germoplasm plants can be an important method for B. tabaci control in integrated pest management approaches. The attractiveness and ovipositional preference of B. tabaci for 17 tomato genotypes were evaluated in a free-choice test. Trials were set up in a randomized block design with ten replicates. Each replicate (one tomato plant per pot) was placed in a cage (80 x 50 x 50 cm) and infested with 1,000 adults during four days. Linear correlation tests were applied between the number of insects and eggs and number of trichomes in each tomato genotype. LA716, LA444-1 and PI134418 genotypes were the least attractive, while the 'Santa Clara' was the most attractive; PI134417 trapped the largest number of adults. LA716 genotype (4.1 eggs/leaflet and 2.1 cm2/eggs per leaflet) was the least preferred for whitefly oviposition; NAV1062, 'Fanny', LA1335, 'Santa Clara' and IAC294 were the most preferred genotypes. The glandular trichomes density was negatively correlated with whitefly's attractiveness and oviposition per leaflet and per leaf, and positively with the number of trapped insects. The non-glandular trichomes density was negatively correlated with the number of trapped insects and positively with whitefly's oviposition per cm2/leaflet andper cm2/leaf. LA716 had high antixenosis level (ovipositional nonpreference) toward B. tabaci B biotype related with type IV glandular trichome.     

烟粉虱和温室粉虱在甘蓝上的刺探取食行为比较

利用刺吸电波图技术研究B型、ZHJ_1型烟粉虱Bemisia tabaci(Gennadius)和温室粉虱Trialeurodes vaporariorum(Westwood)在甘蓝上的取食行为,将3种粉虱的电波图进行比较,其中,B型和ZHJ_1型烟粉虱记录到np,C,pd,E1,E2,F和G波7种波形,温室粉虱只记录到刺探波形,少有取食波形。B型的20个记录中只有1个没有持续吸食波形;ZHJ_1型的25个记录中有10个记录没有持续吸食波形;温室粉虱没有持续吸食记录。甘蓝叶片韧皮部的结构或汁液的化学成分与温室粉虱的抗性密切相关。     

Extraordinary resistance to insecticides reveals exotic Q biotype of Bemisia tabaci in the New World

A strain of the whitefly Bemisia tabaci (Gennadius) possessing unusually high levels of resistance to a wide range of insecticides was discovered in 2004 in the course of routine resistance monitoring in Arizona. The multiply resistant insects, collected from poinsettia (Euphorbia pulcherrima Willd. ex Klotzsch) plants purchased at a retail store in Tucson, were subjected to biotype analysis in three laboratories. Polyacrylamide gel electrophoresis of naphthyl esterases and sequencing of the mitochondrial cytochrome oxidase I gene (780 bp) confirmed the first detection of the Q biotype of B. tabaci in the New World. This U.S. Q biotype strain, referred to as Poinsettia'04, was highly resistant to two selective insect growth regulators, pyriproxyfen and buprofezin, and to mixtures of fenpropathrin and acephate. It was also unusually low in susceptibility to the neonicotinoid insecticides imidacloprid, acetamiprid, and thiamethoxam, relative to B biotype whiteflies. In 100 collections of whiteflies made in Arizona cotton (Gossypium spp.), vegetable, and melon (Cucumis melo L.) fields from 2001 to 2005, no Q biotypes were detected. Regions of the United States that were severely impacted by the introduction of the B biotype of B. tabaci in the 1980s would be well advised to promote measures that limit movement of the Q biotype from controlled environments into field systems and to formulate alternatives for managing this multiply-resistant biotype, in the event that it becomes more widely distributed.     

Biological aspects of Bemisia tabaci biotype B and the chemical causes of resistance in soybean genotypes

In Brazil, the silverleaf whitefly (SLW), Bemisia tabaci biotype B, is a serious soybean pest. SLW management is difficult and new control strategies, such as host–plant resistance, are required. Herein, we aimed to evaluate the biology of SLW, from eggs to adults, on seven soybean cultivars. The emergence of adult insects was monitored daily. Defense-related compounds were identified and quantified from the V3 to the V8 stages in SLW-infested and non-infested plants. The rates of emergence of SLW adults were lower in cultivars ‘IAC 17,’ ‘IAC 19’, and ‘IAC 24’ compared with the susceptible cultivar ‘IAC Holambra Stwart’. Rutin, genistin, genistein, and salicylic acid were identified and quantified in plant extracts. The rutin, genistin, and genistein levels decreased after SLW infestation. Rutin concentrations increased in infested plants of ‘IAC 17’ (V6) and ‘IAC 19’ (from V5 to V8). ‘Barreiras’ showed the highest genistein content in non-infested plants; from V5 growth stage, it was not detected in cultivars Doko (infested), Vencedora, ‘IAC 17’, and ‘IAC 24’ (non-infested). High levels of salicylic acid were observed in ‘IAC ‘19’-infested plants (at V3 and V5). The results suggest that rutin can be related to SLW adults’ emergence only when the feeding source was ‘IAC 19’. Consequently, further studies are needed to access the associated gene expressions and the effect of other secondary metabolites, mainly volatile compounds from SA pathway, including its consequences on feeding preference and mostly in relation to IAC cultivars.     

抗烟粉虱大豆种质资源筛选和抗性机制初探

烟粉虱(Bemisia tabaci Gennadius)近年来发生日益猖獗、危害日趋严重,防治比较困难,却未被列为主要经济害虫.筛选抗性种质和选育抗性品种是防治烟粉虱的有效措施.本研究对223份大豆种质资源进行了抗烟粉虱鉴定,筛选出滑皮豆等6份抗性较强的种质资源.调查了这223份种质资源的茸毛性状,测定了部分抗感烟粉虱品种的籽粒蛋白质和脂肪含量.结果表明,大豆品种对烟粉虱的抗性与叶片的茸毛性状有密切关系,无茸毛型抗性最强,茸毛紧贴型次之,茸毛直立型较差,茸毛斜立型抗性最差.大豆受烟粉虱危害程度与籽粒蛋白质和脂肪含量有密切关系,蛋白质含量越高受害越严重,脂肪含量越高受害越轻.根据鉴定结果,本研究提出了单叶平均感染烟粉虱0头为免疫,0.1～3.0头为高抗,3.1～10.0头为中抗,10.1～20.0头为中感,20头以上为高感的抗性鉴定标准.     

Performance of <Emphasis Type="Italic">Bemisia tabaci</Emphasis> Biotype B on Soybean Genotypes

Bemisia tabaci (Genn.) (Hemiptera: Aleyrodidae) has been recognized as an important pest of many agricultural systems including soybean [Glycine max (L.) Merrill] crops. As an alternative to chemical control, the use of resistant genotypes represents an important tool for integrated pest management (IPM). This study aimed to evaluate the biological development of Bemisia tabaci biotype B confined on 13 soybean genotypes under greenhouse conditions. Initially, the nymphal period, complete development period (egg–adult), and the viability of the silverleaf whitefly nymphs were evaluated in all genotypes. Then, four genotypes promising for resistance (‘Jackson,’ UX-2569-159, ‘P98Y11,’ and ‘TMG132 RR’) and a susceptible genotype (PI-227687) were selected for further assays, where two insect populations were compared: a first population from the initial rearing (cabbage plants) and another corresponding to insects previously reared out on the selected genotypes. In addition to the parameters evaluated in preliminary tests, we also determined the viability and incubation period of eggs. Moderate levels of resistance (antibiosis/antixenosis) to B. tabaci biotype B were found in three genotypes. ‘P98Y11’ and ‘TMG132 RR’ were less suitable for insect development, extending the development cycle, and UX-2569-159 caused high nymphal mortality. We did not observe a significant increase in the level of plant resistance by the use of previously stressed insects. This suggests that the evaluation of a single whitefly generation may be sufficient to make correct decisions on promising soybean genotypes.     

Biotype-dependent secondary symbiont communities in sympatric populations of Bemisia tabaci

The sweet potato whitefly, Bemisia tabaci, harbors Portiera aleyrodidarum, an obligatory symbiotic bacterium, as well as several secondary symbionts including Rickettsia, Hamiltonella, Wolbachia, Arsenophonus, Cardinium and Fritschea, the function of which is unknown. Bemisia tabaci is a species complex composed of numerous biotypes, which may differ from each other both genetically and biologically. Only the B and Q biotypes have been reported from Israel. Secondary symbiont infection frequencies of Israeli laboratory and field populations of B. tabaci from various host plants were determined by PCR, in order to test for correlation between bacterial composition to biotype and host plant. Hamiltonella was detected only in populations of the B biotype, while Wolbachia and Arsenophonus were found only in the Q biotype (33% and 87% infection, respectively). Rickettsia was abundant in both biotypes. Cardinium and Fritschea were not found in any of the populations. No differences in secondary symbionts were found among host plants within the B biotype; but within the Q biotype, all whiteflies collected from sage harboured both Rickettsia and Arsenophonus, an infection frequency which was significantly higher than those found in association with all other host plants. The association found between whitefly biotypes and secondary symbionts suggests a possible contribution of these bacteria to host characteristics such as insecticide resistance, host range, virus transmission and speciation.     

Comparison of greenhouse screening materials for excluding whitefly (Homoptera: Aleyrodidae) and thrips (Thysanoptera: Thripidae)

Twenty-eight greenhouse screening materials, with predetermined airflow resistance values, were evaluated for exclusion of the silverleaf whitefly Bemisia argentifolii Perring & Bellows and thrips from a mixed-species population. Screens differed in exclusion efficacy, expressed as a percentage of the fiberglass window screen control and at an approach velocity of 92 m/min, from -35 to 94% for silverleaf whitefly and from -13 to 95% for thrips. Seventeen screens excluded more silverleaf whitefly than did the window screen control, whereas only seven excluded more thrips. One material differentially excluded whitefly over thrips; many more differentially excluded thrips over whitefly. Airflow resistance, indicative of mesh hole size, did not necessarily correspond with degree of exclusion. Not all materials characterized as highly resistant to airflow provided significant exclusion. Exclusion of both types of pests was attained with several moderate- and one low-resistance screen. Another low-resistance screen excluded silverleaf whitefly only.     

Discovering new insect viruses: whitefly iridovirus (Homoptera: Aleyrodidae: Bemisia tabaci)

Adult whiteflies, Bemisia tabaci (Gennadius), collected from the field were screened for viral pathogens using a cell line from the silverleaf whitefly, B. tabaci, B biotype (syn. B. argentifolii). Homogenates from the field-collected whiteflies were applied to cell cultures and checked for cytopathic effects (CPE). Cells were observed to develop cytoplasmic inclusions and to have a change in morphology. Cells displaying CPE were observed using a transmission electron microscope and found to be infected with a virus. The virus particles had an icosahedral shape and an approximate size of 120-130 nm. The virus was observed in defined areas of the cytoplasm adjacent to the cell nucleus. Analysis using polymerase chain reaction, Southern blot hybridization, and DNA sequencing confirmed that the virus discovered infecting the whitefly cell cultures was an iridovirus. Sequence analysis showed that the amplimer (893 bp) had a 95% homology to the invertebrate iridescent virus type 6 major capsid protein gene. Discovery of new viruses of whiteflies may provide renewed interest in using pathogens in the development of innovative management strategies. This is the first report of an iridescent virus isolated from whiteflies, B. tabaci, collected from the field.     

世界性重要害虫B型烟粉虱的入侵机制

B型烟粉虱是近二十年来入侵世界各国并暴发成灾的一种世界性重要害虫。近年来, B型烟粉虱的入侵机制研究表明,其成功定殖、扩散、暴发与其对入侵地区的土著烟粉虱种群和其它昆虫种群的竞争取代有关;竞争取代包括生态位竞争,生殖干涉,其危害寄主对其它昆虫的影响等方面。B型烟粉虱抗药性的形成也是这种生物成功入侵的重要影响因素。同时,生态环境（包括寄主植物、气候因子、越冬场所等）的影响也不容忽视。最后本文还探讨了B型烟粉虱入侵机制研究动态及其意义。