Variation of pre-ovipositional period in the brown planthopper, Nilaparvata lugens, collected in tropical, subtropical and temperate Asia

Abstract:  We investigated the pre-ovipositional period of Nilaparvata lugens originating from tropical, subtropical and temperate regions of East and Southeast Asia using laboratory experiments to compare the migration capability of the populations. Macropter females collected in 1992 from Japan and subtropical North Vietnam had a longer immature period than those from tropical Indochina Peninsula, i.e. the day of the first oviposition after eclosion was about 3 days later in the former populations. Populations collected in 1993 and 1994 from Japan, Central and South China, and North Vietnam also had a longer immature period than that of the tropical Malaysian population. Thus, a general trend was demonstrated that macropters originating in temperate and subtropical East Asia had a longer pre-ovipositional period, which is presumably beneficial for pre-reproductive long-distance migration. This finding supports a current hypothesis for the migration system of N. lugens existing in East Asia, which has maintained the capability of long-distance migration throughout years in spite of the disadvantage of prolonged oviposition for multiplication.     

Responses by the brown planthopper,Nilaparvata lugens,to conspecific density on resistant and susceptible rice varieties

This study examines the nature of intraspecific interactions among Nilaparvata lugens (Stål) (Hemiptera: Delphacidae) planthoppers feeding on resistant and susceptible rice varieties. Planthopper nymphs produced less honeydew and gained less weight when feeding on rice variety IR62 (resistant) compared to susceptible rice varieties. A series of bioassays was conducted that varied N. lugens nymph densities on IR62 and IR22 (susceptible). Increasing nymph density facilitated feeding by conspecifics; however, intraspecific competition increased mortality of nymphs on IR62 (but rarely on IR22). Furthermore, nymph weights declined with increasing conspecific density on IR22, and the effects were weak on IR62. More female nymphs than males survived on IR62 but this was not affected by density. Nitrogenous fertilizer increased competition among N. lugens on young plants of IR22, but not on IR62. Results indicate that nymphs have a low efficiency in accessing resources when feeding on IR62, even where the plants have received fertilizer. Female‐biased survival and biomass compensation for mortality may promote population recovery after development on the resistant plant and accelerate adaptation to the resistant variety.     

Meiosis in males of the brown planthopper, Nilaparvata lugens

Meiosis in spermatocytes was studied among the three biotypes of the brown planthopper, Nilaparvata lugens (Stål). The chromosome number, morphology, and behavior of the biotypes were similar. The autosomes divided normally, reductional for the first division and equational for the second. The X chromosome had an unstable segregation behavior; it may segregate equationally or reductionally during the first division depending on its chromatid's orientation. Metaphase I plates contained 14 autosomal bivalents and an X chromosome (14_II + X). The co-oriented sister chromatids of the X chromosome moved together to either one of the poles during anaphase I, while auto-oriented sister chromatids segregated to opposite poles. Because of the anomalous segregation behaviour of the X chromosome, metaphase II plates may both contain 14_I + X chromatid, or one with only 14_I and the other with 14_I + X chromosome. Male N. lugens has a diploid chromosome number of 29.

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Zusammenfassung Die Meiose in Spermatozyten wurde in drei Biotypen von Nilaparvata lugens (Stål) studiert. Die Anzahl der Chromosomen, die Morphologie und das Verhalten der Biotypen waren ähnlich. Die Autosomen teilten in einer normalen Weise, reduzierend für die erste und äquatorial für die zweite Teilung. Das X-Chromosom hatte ein unstabiles Teilungsverhalten; es kann sich äquatorial oder aber auch reduktional trennen, je nachdem wie sich sein Chromatid orientiert hat. Die Ebenen der Metaphase I enthalten 14 autosome Bivalente und ein X-Chromosom (14_II + X). Die co-orientierten Schwester-chromatiden des X-Chromosoms bewegen sich während der Anaphase I zusammen zu einem der beiden Pole, während auto-orientierte Schwester-chromatiden sich nach entgegengesetzten Polen trennen. Wegen des ungewöhnlichen Teilungsverhaltens des X-Chromosoms, können beide Metaphase II Ebenen 14_I und X Chromatid enthalten, oder aber eine Ebene enthält 14_I und die andere 14_I + X.

     

Inheritance of body color in the brown planthopper, Nilaparvata lugens

The segregation frequencies of color phenotypes of F₁, F₂, and backcross progenies revealed that the body coloration of the brown planthopper, Nilaparavata lugens(Stål), exhibited multiple allelism. Monogene for color morphism exists in three allelic forms >-b ⁺ (brown or wild type), b ^o (orange) and b ^b (black). The dominance relationship was b ^b>b ⁺>b ^o. The relationship between phenotype and genotype in N. lugens was not fixed. The six possible genotypes produced eight phenotypes.

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Résumé Les fréquences de ségrégation des différentes colorations de F₁, F₂ et des croisements en retour ont montré que la coloration du corps de Nilaparvata lugens est due à un multi-allèlisme. Le monogène pour la couleur existe sous trois formes allèliques: b⁺ (brun ou sauvage), b^o (orange) et b^b (noir). Les relations de dominance étaient: b^b>b⁺>b^o. On n'a pas établi les relatios entre phénotypes et génotypes chez N. lugens. Les 6 génotypes possibles produisent 8 phénotypes.

     

褐飞虱抗药性研究现状

褐飞虱Nilaparvata lugens(Stal)对杀虫剂产生抗药性是其近年来暴发频繁的重要原因。文章综述国内外关于褐飞虱抗药性的研究成果,包括褐飞虱抗性测定方法、抗药性的发展、交互抗性、抗性遗传、抗性机理及抗性治理等。田间褐飞虱种群对新烟碱类药剂产生不同程度的抗药性,其中对吡虫啉产生高水平到极高水平抗性,对氯噻啉和噻虫嗪分别产生中等水平和低水平的抗药性,对呋虫胺和烯啶虫胺仍然处于敏感性阶段。此外,褐飞虱种群对噻嗪酮(昆虫生长调节剂)产生低水平到中等水平抗性。长期大面积使用化学药剂是褐飞虱产生抗药性的重要原因。因此,必须加强褐飞虱的抗性治理,以延缓其抗药性进一步发展。     

Dynamics of imidacloprid resistance and cross-resistance in the brown planthopper, Nilaparvata lugens

Extensive use of imidacloprid for suppressing the brown planthopper, Nilaparvata lugens (Stål) (Homoptera: Delphacidae), has placed heavy selection pressure on the target insect. A systematic study was carried out to determine imidacloprid resistance dynamics and cross-resistance. Data collected from a 3-year study (2005–2007) showed that in 2005, the resistance levels in Nanning (Guangxi), Haiyan (Zhejiang), and Nanjing and Tongzhou (Jiangsu) populations ranged from 200- to 799-fold compared with the susceptible strain. However, the resistance levels decreased to 135- to 233-fold in 2007, after reduced application of the chemical. A laboratory population was challenged with imidacloprid in successive generations. After 23 generations, the resistance ratio had increased from 200- to 1 298-fold. Continuous selection with imidacloprid could increase the resistance level even more than has already been developed in the population. Stopping selection with imidacloprid led to a rapid decrease of resistance from 759- to 114-fold after 17 generations. Resistance levels then became stable without decreasing any further. A similar result was also obtained from a study involving a field population (resistance ratio = 625-fold) collected from Tongzhou. At first, the population showed a rapid decrease in resistance right after imidacloprid selection was stopped, and then the resistance stabilized at a level of 105–129-fold. More interestingly, resistance increased again when selection was resumed. In addition, the resistant strain selected with imidacloprid showed substantial cross-resistance to imidaclothiz, thiacloprid, and acetamiprid, and slight levels of cross-resistance to dinotefuran and thiamethoxam, but no obvious cross-resistance to nitenpyram, buprofezin, and fipronil. The information from this study is valuable for formulating resistance-management strategies against N. lugens .     

Starvation tolerance of macropter brown planthopper, Nilaparvata lugens, from temperate, subtropical, and tropical populations in East and South-East Asia

We compared the starvation tolerance of macropter brown planthopper, Nilaparvata lugens Stål (Homoptera: Delphacidae), a characteristic favoring long-distance migration, among populations collected from subtropical and temperate East Asia, tropical Indochina, and the Malay Peninsula. Starvation tolerance of planthoppers was significantly affected by climatic zone in which the planthoppers had been collected and by feeding duration after adult eclosion. After 24 h feeding on rice, newly emerged macropters originating from East Asia lived longer without feeding (starvation tolerance) than macropters from the tropical populations. The difference in longevity between the two groups of populations became more conspicuous when macropters fed on rice for 48 or 72 h, indicating that post-eclosion feeding markedly increased starvation tolerance in East Asian populations relative to the tropical populations. These facts provide evidence that N. lugens populations that are adapted for long-distance migration are distributed in subtropical and temperate East Asia. From the data on post-eclosion feeding together with starvation tolerance, we discuss the timing of planthopper takeoff from paddies and the difference in resource allocation (vitellogenesis or stored resources) between East Asian and tropical populations.     

Genetic mapping of the rice resistance-breaking gene of the brown planthopper Nilaparvata lugens

Host plant resistance has been widely used for controlling the major rice pest brown planthopper (BPH, Nilaparvata lugens). However, adaptation of the wild BPH population to resistance limits the effective use of resistant rice varieties. Quantitative trait locus (QTL) analysis was conducted to identify resistance-breaking genes against the anti-feeding mechanism mediated by the rice resistance gene Bph1. QTL analysis in iso-female BPH lines with single-nucleotide polymorphism (SNP) markers detected a single region on the 10th linkage group responsible for the virulence. The QTL explained from 57 to 84% of the total phenotypic variation. Bulked segregant analysis with next-generation sequencing in F₂ progenies identified five SNPs genetically linked to the virulence. These analyses showed that virulence to Bph1 was controlled by a single recessive gene. In contrast to previous studies, the gene-for-gene relationship between the major resistance gene Bph1 and virulence gene of BPH was confirmed. Identified markers are available for map-based cloning of the major gene controlling BPH virulence to rice resistance.     

Electrical penetration graphs indicate that tricin is a key secondary metabolite of rice,inhibiting phloem feeding of brown planthopper,Nilaparvata lugens

Tricin (5,7,4′‐trihydroxy‐3′,5′‐dimethoxyflavone) is a valuable secondary metabolite which is widely present in gramineous plants, including cultivated rice (Oryza sativa L.) (Poaceae). It can defend the rice plant against damage by the brown planthopper (BPH), Nilaparvata lugens (Stål) (Hemiptera: Delphacidae), one of the most important pests of rice. This study was conducted to elucidate the mechanisms of action of tricin on BPH feeding behavior. BPH feeding behavior in resistant (Rathu Heenati, RHT) and susceptible (Taichuang native 1, TN1) rice varieties and artificial diets was monitored using the electrical penetration graph (EPG) technique. Tricin concentrations in leaves of varieties RHT and TN1 were quantitatively analyzed by liquid chromatography, coupled to tandem mass spectrometric techniques. Six (NP and N1‐5) and four (NP, N1, N2, and N4) types of waveforms occurred during feeding on rice plants and artificial diets, respectively. The tricin concentration of rice varieties was correlated with total and average durations of N4. Moreover, EPG data indicated that tricin significantly increased the duration of non‐probing and pathway periods and strongly inhibited phloem ingestion (N4). The inhibition was strongly dose dependent, resulting in complete suppression of activity in the phloem region when the tricin concentration was increased to 1 g l^?1. This study revealed that tricin disturbed the feeding behavior of BPH mainly by increasing the non‐probe period and inhibiting phloem ingestion. We confirmed the hypothesis that tricin is a ‘stylet probing stimulant’ of rice planthoppers as proposed in previous studies. The information on the ecological effect of tricin from this study may be useful to clarify the resistance mechanism against BPH of RHT and other tricin‐containing rice varieties.     

Effects of triflumezopyrim on the reproduction of brown planthopper,Nilaparvata lugens

The brown planthopper, Nilaparvata lugens Stål (Hemiptera: Delphacidae), is a notorious pest on rice in Asia and has evolved serious resistance to traditional chemical insecticides in recent years. Triflumezopyrim, a newly commercialized mesoionic insecticide, exhibited an excellent control effect on sucking insects, including N. lugens. However, the effects of triflumezopyrim on the reproduction of N. lugens have not yet been fully elucidated. In this study, we investigated the impacts of different concentrations of triflumezopyrim (LC₁₀, LC₅₀ and LC₉₀) on the reproduction and the reproduction related genes of N. lugens exposure to the insecticide via systemic and contact route. The results indicated that the preoviposition periods of N. lugens females in the LC₅₀ and LC₉₀ triflumezopyrim-treated groups (♀ t × ♂ t) were prolonged in contact toxicity treatment, compared with control (acetone treatment). The fecundities for three pair combinations (♀ c ×♂ t, ♀ t ×♂ c and ♀ t ×♂ t) treated with LC₅₀ and LC₉₀ concentrations of triflumezopyrim showed a significant decrease compared to water/acetone-treated control via contact and systemic route. No significant effects of LC₁₀ triflumezopyrim on preoviposition period and fecundity were found in contact and systemic toxicity treatments. However, the triflumezopyrim did not influence the egg hatchability. By microscope observation, the LC₅₀ and LC₉₀ triflumezopyrim impended the ovarian development in females. qRT-PCR analysis showed that the vitellogenin (NlVg) relative expression level of female adults exposed to LC₅₀ and LC₉₀ triflumezopyrim decreased by 30.3 – 39.5% in two toxicity treatments. The titers of another reproduction related hormones juvenile hormone III (JH III) in LC₅₀ and LC₉₀ triflumezopyrim-treatments were 27.4% and 25.8% lower than control in contact toxicity treatment and 27.08% and 37.18% lower than control in systemic toxicity treatment. In summary, these results suggested that the median lethal and high lethal concentrations of triflumezopyrim inhibited the reproduction and reduced the expression of NlVg and JH III in N. lugens. This study provided a reference for the efficient and scientific utilization of triflumezopyrim in the field.     

褐飞虱对吡虫啉抗性监测方法的研究

从结果的稳定性、准确性和操作方便等角度出发 ,建立了适合于监测褐飞虱 (NilaparvatalugensSt l)对吡虫啉抗性的稻苗浸根法。与其他方法相比 ,该方法除具有简单易行的特点外 ,还表现出很好的监测准确性。通过建立不同虫态和不同若虫龄期敏感基线以及对桂林、安庆、东台 3个地区的抗性监测 ,发现 3地褐飞虱对吡虫啉都处于敏感性下降阶段 ,安庆和东台地区的敏感性弱低于桂林     

有机稻田褐稻虱种群动态及天敌作用分析

为组建有机稻田褐稻虱Nilaparvata lugens(Stl)种群生态控制系统,系统调查实施有机耕作1年后褐稻虱种群动态,并应用生命表技术评价自然天敌的作用。结果表明,有机晚稻田中褐稻虱种群发生2个世代。高峰期分别为9月底~10月初、10月下旬,平均百丛达502.3,164.5头,种群增长率为常规稻田的31.76%。水稻移栽后天敌数量迅速回升,有机稻田中蜘蛛、捕食性天敌、寄生性天敌分别比常规稻田增长66.10%,166.67%,44.24%。将各类天敌作用相加,以"捕食及其它"、"寄生"对褐稻虱种群的控制作用最强,其EIPC值均显著高于常规稻田相应天敌因子EIPC。     

Wing polymorphism in Japanese and foreign strains of the brown planthopper, Nilaparvata lugens

Great variations in nymphal density and wing-form relationships occur in field populations of the brown planthopper, Nilaparvata lugens (Stål) (Homoptera: Delphacidae). When tested under uniform laboratory conditions, half of the female populations collected from 20 locations in Japan exhibited higher ratios of macropters with increasing nymphal density, but one quarter were macropterous over broad ranges of density, both types being classified as group III. Three predominantly brachypterous populations (group I) were found, even under crowded conditions and one intermediate (group II).The populations from South Asian countries, i.e. Philippines (one), Indonesia (two) and Malaysia (one) belonged to group I, but the three populations from the coastal areas of China belonged to group II. Those from Taiwan differed in responses to density; one was of group II type, and two of group III type.From these comparisons, immigration routes of N. lugens to Japan were estimated, and survival strategy of this migratory species is discussed.

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Zusammenfassung In Feldpopulationen von Nilaparvata lugens (Stål) (Homoptera: Delphacidae) gibt es grosse Variationen in der Larvendichte und in den Flügelform-Beziehungen. Unter einheitlichen Laboratoriumsbedingungen wurden diese Verhältnisse geprüft. Von 20 japanischen Herkünften zeigte die Hälfte der Weibchenpopulationen mit zunehmender Larvendichte einen höheren Macropterenanteil; ein Viertel was über einen grossen Dichtebereich macropter; beide Typen werden als Gruppe III bezeichnet. Drei Populationen wurden gefunden, die auch bei hoher Dichte vorwiegend brachypter waren (Gruppe I). Einer Zwischengruppe (II) gehörte eine einzige Population an.Die Populationen aus südasiatischen Ländern (1 aus den Philippinen, 2 aus Indonesien un 1 aus Malaysia) gehörten zu Gruppe I, die 3 Populationen aus den Küstenregionen von China jedoch zu Gruppe II. Die Populationen aus Taiwan unterschieden sich in der Reaktion auf die Dichte: 1 gehörte zu Gruppe II and 2 zu Gruppe III.Von diesen Daten werden Einwanderungswege von N. lugens nach Japan abgeleitet und die Überlebensstrategie dieser Wanderart diskutiert.

     

Gene knockdown by intro-thoracic injection of double-stranded RNA in the brown planthopper,Nilaparvata lugens

RNA interference (RNAi) is a powerful strategy for gene function study in insects. Here, we described the development of a RNAi technique by microinjection of double-stranded RNA (dsRNA) in the brown planthopper Nilaparvata lugens. Based on the mortality and RNAi efficiency criteria, the conjunctive between prothorax and mesothorax was selected as the injection site and 50 nl as injection volume. Three genes with different expression patterns were selected to evaluate the RNAi efficiency. A comparable 40% decrease of gene expression was observed at the 4th day after injection for the ubiquitously expressed calreticulin and the gut specific cathepsin-B genes, but only 25% decrease at the 5th day for the central nervous system specific Nlβ2 gene. Double injection could increase the RNAi efficiency, such as from 25% to 53% for Nlβ2 gene. The gene knockdown technique developed in this study will be an essential post-genomic tool for further investigations in N. lugens.     

水稻品种对褐飞虱的田间抗性及其生态学机制

评价了7个水稻品种(浙粳22、宜香845、中组14、秀水123、二优倍九、感虫对照品种TN1和抗虫对照品种IR36)在田间成株期对褐飞虱Nilaparvata lugens(Stl)的抗性表现,并初步分析了它们的抗性机制。结果表明,在田间网室人工诱发条件下,浙粳22、宜香845、秀水123和中组14等4个品种的抗性水平与抗虫品种IR36相同,均小于3级,表现为抗虫,而二优培九则与感虫品种TN1相同,表现为感虫。在自然发生条件下的水稻分蘖期至乳熟期,宜香845和中组14上褐飞虱数量仅为TN1上的7.84%～24.44%,秀水123和浙粳22则为TN1上的30.19%～60.27%。在4个抗性品种上褐飞虱的产卵量和卵孵化率明显减少,而若虫存活率仅为TN1上的20%～30%。与IR36相同,4个抗性品种上褐飞虱的若虫存活率和种群增长倍数均显著低于TN1上,说明他们对褐飞虱的抗性主要表现为抗生性。