Electrical penetration graphs from Nilaparvata lugens on resistant and susceptible rice varieties

The feeding behaviour of Nilaparvata lugens (Stal.) was electronically monitored on three rice varieties showing different levels of resistance in the Philippines, over an 8 h period, using a DC amplifier and a video-assisted observation method to simultaneously record honeydew production.Six electrical penetration graph (EPG) patterns are described by their frequency and amplitude characteristics. Patterns 4 and 5 were associated with the production of two different types of honeydew; basic honeydew containing amino acids was produced during Pattern 4 and acidic honeydew with no amino acids was produced during Pattern 5. These patterns are thought to indicate phloem and xylem ingestion by N. lugens respectively.Insects produced shorter periods of Pattern 4 on moderately resistant IR46 and resistant IR62 rice varieties than on susceptible IR22, although the time taken to produce the first period of Pattern 4 from the start of EPG recording did not differ between varieties. This suggests that N. lugens has the ability to initiate phloem ingestion on all three varieties, but that resistance mechanisms in IR46 and IR62 prevent sustained phloem ingestion, at least within the first 8 h of contact.

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Résumé Le comportement alimentaire de N. lugens Stahl a été enregistré életroniquement pendant 8 heures aux Philippines sur 3 variétés de riz présentant différents degrés de résistance, en utilisant un amplificateur DC associé à un système vidéo pour enregistrer simultanément la production de miellat.Six types d'enregistrements de pénétration (EPG) sont décrits avec leurs caractéristiques de fréquence et d'amplitude. Les types 4 et 5 sont associés à la production de 2 types différents de miellat; le miellat de base contenant des acides aminés est produit pendant le type 4, et le miellat acide sans acides aminés est produit pendant le type 5. On considére que ces types indiquent respectivement l'ingestion de phloème et xylème par N. lugens.Les périodes de type 4 ont été plus courtes sur les variétés modérément résistante IR46 et résistante IR62 que sur la variété sensible IR22, bien que le temps passé à produire la première période de type 4 depuis le début des enregistrements des EPG ne diffère pas suivant les variétés. Ceci laisse penser que N. lugens a la possibilité de commencer la consommation de phloème sur les trois variétés, mais que les mécanismes de résistance chez IR46 et IR62 empêchent une ingestion soutenue de phloème, au moins pendant les 8 premières heures de contact.

     

水稻品种中抗褐飞虱抗原次生物质的分析

借助高效液相色谱(HPLC)技术,研究了130份对褐飞虱生物型Ⅱ具有不同抗性水平的水稻样品(26个品种)中13个次生物质含量(峰面积)的差异;运用主成分分析和多元回归分析,建立了水稻品种抗性级别的预测模型：Y=3．4593-0．02491X1+0．08475X2-0．04227X8+0．1174X12结果表明,水稻的抗性水平与峰面积值之间极显著相关(r^2=0．84,P<0．01),峰1、峰2、峰8、峰12对应的次生化合物是影响水稻对褐飞虱生物型Ⅱ抗性水平的主要抗原次生化合物;水稻品种中起抗虫作用的抗原次生物质不止一种,而是几种的组合,而且它们对水稻抗虫性的贡献权重是不完全相同的．     

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.

     

Role of rice volatiles in the foraging behaviour of the predator Cyrtorhinus lividipennis for the rice brown planthopper Nilaparvata lugens

Experiments were conducted inlaboratory to study the impacts of ricegenotypes and rice plants treated withdifferent levels of nitrogen on the predationrates of the predator, Cyrtorhinuslividipennis Reuter, for eggs of the rice brownplanthopper (BPH), Nilaparvata lugens(Stål), and their relation to the ricevolatiles. In two-choice test, the predatorfemales showed different predation rates forBPH eggs on different rice genotypes, but notfor BPH eggs on rice plants treated withdifferent levels of nitrogen. The olfactoryresponse test revealed that more femalepredators oriented to the odours from healthyrice plants compared with the plain air(control); the predator preferred plant-BPHnymph complex over healthy plants, andplant-BPH gravid female complex over plant-BPHnymph complex. Additional experimentsdemostrated that there were obvious differencesin attractiveness of the healthy plants andplant-BPH gravid female complex of various ricegenotypes, while no significant differenceswere found between rice plants treated withdifferent levels of nitrogen. The results ofpredation rates of the predator on variousvarieties and plants treated with differentlevels of nitrogen were consistent with that ofits olfactory response in 5 of the 6 casesevaluated, which showed that rice volatilesplayed an important role in the foragingbehaviour of C. lividipennis.Implications for augmenting the effectivenessof natural enemies by adjusting rice attributesand cultural practices are discussed.     

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.

     

Determination and distribution of a female-specific protein in the brown planthopper, Nilaparvata lugens Stal (Homoptera: Delphacidae)

Electrophoretic analyses of hemolymph and body or ovary homogenates from reproducing females, males, and 5th instar nymphs of the brown planthopper, Nilaparvata lugens Stal, revealed a protein band of 175 kDa in females. An immunoblot test using antibody against this protein showed a positive reaction with a 175 kDa protein from female body or ovary homogenates. It is likely that this protein in hemolymph is vitellogenin (Vg). Distribution of Vg was determined by immunofluorescence and immunogold labeling techniques. The results showed that the positive immunofluorescence reactions were present in yolk particles, the intercellular space of follicle cells, hemolymph, and the epithelial plug of ovarioles. In addition, the yeast-like symbiotes (YLS) in mycetocytes of adults and various nymphal instars as well as those free in hemolymph or entering oocytes also exhibited a positive reaction. Electron micrographs showed that immunogold particles were found most in yolk mass and YLS over other tissues. Especially the YLS in various developmental stages all contained immunogold particles, implying that the symbiote is somewhat related with production of the female-specific protein.     

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.

     

抗性水稻品种对褐飞虱和白背飞虱种群参数的影响

在半自然状态下 ,用生命表方法初步比较了褐飞虱Nilaparvatalugens和白背飞虱Sogatellafurcif era在 4个抗褐飞虱水稻品种ASD7,IR3 6,JX89和Mudgo上的种群参数。初步的结果表明 ,白背飞虱在 4个抗褐飞虱水稻品种上具有较高的繁殖率 ,较短的发育时间和较大的种群增长。对褐飞虱各种群参数影响最大的是JX89,对白背飞虱影响最大的是Mudgo。与感性品种TN1比较 ,这些抗褐飞虱的水稻品种对 2种稻飞虱均具有不同程度的抗性。这些结果说明 ,抗褐飞虱水稻品种对白背飞虱同样具有一定的抗性 ,但白背飞虱对这种抗性具有较强的适应性 ,如果在一个地区长期推广抗褐飞虱的水稻品种 ,势必会导致白背飞虱的大发生     

褐飞虱对抗性水稻品种Mudgo个体致害性指标

在28±1℃下,观测了褐飞虱Nilaparvata lugens羽化24?h内短翅型雌成虫在敏感水稻品种TN1及抗性品种Mudgo上的寿命及体重增量。结果表明,取食抗性品种时,试虫的平均增重与寿命之间表现出明显的正相关。寿命在7天内的个体,存活期间平均体重增量为负值,而存活7天以上的个体则平均增重明显。进一步研究了初羽化的褐飞虱长翅型雌成虫在TN1及Mudgo上取食第1～4天的体重增量、寿命及产卵量,结果发现试虫在Mudgo上的这3个生物学参数之间存在明显正相关。其中,寿命及体重增量较好地反映出害虫个体对抗性品种的致害能力,可作为个体致害性指标。根据研究结果,提出将羽化后在Mudgo上存活7天以上或最初4天内体重增量大于0.1mg的雌成虫定为能够致害该抗性品种的个体。     

Detection and analysis of QTLs for resistance to the brown planthopper, Nilaparvata lugens, in a doubled-haploid rice population

 We used a mapping population of 131 doubled-haploid lines, produced from a cross between an improved indica rice variety (IR64) and a traditional japonica variety (Azucena), to detect quantitative trait loci (QTLs) for resistance to the brown planthopper (BPH), Nilaparvata lugens. We evaluated the parents and mapping population with six tests that measure varying combinations of the three basic mechanisms of insect host plant resistance, i.e., antixenosis, antibiosis, and tolerance. To factor-out the effect of the major resistance gene Bph1 from IR64, the screening was done with two BPH populations from Luzon Island, The Philippines, that are almost completely adapted to this gene. A total of seven QTLs associated with resistance were identified, located on 6 of the 12 rice chromosomes. Individual QTLs accounted for between 5.1 and 16.6% of the phenotypic variance. Two QTLs were predominantly associated with a single resistance mechanism: one with antixenosis and one with tolerance. Most of the QTLs were derived from IR64, which has been shown to have a relatively durable level of moderate resistance under field conditions. The results of this study should be useful in transferring this resistance to additional rice varieties. Received: 10 May 1998 / Accepted: 4 June 1998     

Allozyme variation among biotypes of the brown planthopperNilaparvata lugens in the Philippines

Allozyme variation was studied in threeNilaparvata lugens biotypes infesting specific rice varieties and a biotype infesting a weed grass,Leersia hexandra. Of the 20 enzymes inN. lugens for which activity was noted, 9 were polymorphic. Eleven enzyme loci were monomorphic for the same allele in all biotype populations; the rest were polymorphic for two or more alleles. The mean number of alleles per polymorphic locus was 2.3, while the mean number of alleles per locus was 1.5; heterozygosity ranged from 0.02 to 0.06 (biotype 1 > biotype 3 >Leersia-infesting biotype > biotype 2). Allelic frequency differences were observed in five loci among the four biotypes. However, the coefficient of genetic identity (I) of 0.99+ showed that the four biotype populations were genetically close relatives or merely populations ofN. lugens undergoing genetic differentiation. This work was partly supported by a financial grant received from the Directorate for Technical Cooperation and Humanitarian Aid, Switzerland.     

Differential gene expression in response to brown planthopper feeding in rice

Plant responses to herbivores are complex. 108 cDNA clones representing genes relating to plant responses to chewing insect-feeding, pathogen infection, wounding and other stresses were collected. Northern blot and cDNA array analysis were employed to investigate gene expression regulated by piercing-sucking insect, brown planthopper (BPH), Nilaparvata lugens (Homoptera: Dephacidae) on both the resistant and susceptible rice genotypes. After BPH feeding in rice for 72 h, the expression of most tested genes was affected. 14 genes in resistant rice variety B5 and 44 genes in susceptible MH63 were significantly up- or down-regulated. Most of the well-regulated genes were grouped in the categories of signaling pathways, oxidative stress/apoptosis, wound-response, drought-inducible and pathogen-related proteins. Those related to the flavonoid pathway, aromatic metabolidsm and the octadecanoid pathway were mostly kept unchanged or down-regulated. Our results indicate that BPH feeding induces plant responses which would take part in a jasmonic acid-independent pathway and crosstalk with those related to abiotic stress, pathogen invasion and phytohormone signaling pathways.     

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.     

A Simple Sequence Repeat- and Single-Nucleotide Polymorphism-Based Genetic Linkage Map of the Brown Planthopper,Nilaparvata lugens

In this study, we developed the first genetic linkage map for the major rice insect pest, the brown planthopper (BPH, Nilaparvata lugens). The linkage map was constructed by integrating linkage data from two backcross populations derived from three inbred BPH strains. The consensus map consists of 474 simple sequence repeats, 43 single-nucleotide polymorphisms, and 1 sequence-tagged site, for a total of 518 markers at 472 unique positions in 17 linkage groups. The linkage groups cover 1093.9 cM, with an average distance of 2.3 cM between loci. The average number of marker loci per linkage group was 27.8. The sex-linkage group was identified by exploiting X-linked and Y-specific markers. Our linkage map and the newly developed markers used to create it constitute an essential resource and a useful framework for future genetic analyses in BPH.     

利用抑制差减杂交技术分离受水稻抗性调控的褐飞虱基因

为分离受水稻抗性调控的褐飞虱Nilaparvata lugens基因, 以取食感虫水稻台中1号和高抗水稻B5的2叶1芯秧苗24 h的褐飞虱4龄若虫为起始材料, 采用抑制差减杂交技术构建了两个群体间的正反向差减cDNA文库。通过斑点杂交从差减文库中筛选代表受水稻抗性调控的基因的cDNA克隆, 进行测序和功能分析, 挑选具功能的基因进行Northern杂交验证。结果表明, 通过斑点杂交筛选到的98个阳性克隆代表92个互不重复的单基因, 其中25个与动物的已知蛋白基因存在较高的同源性。Northern杂交表明, 这25个基因有11个表达上调, 8个表达下调, 提示它们可能在褐飞虱适应抗性水稻过程中发挥了重要作用。本研究结果为克隆上述新基因的全长cDNA序列及进一步研究其在褐飞虱与水稻互作中的功能奠定了基础。     

A migration analysis of the rice planthopper Nilaparvata lugens from the Philippines to East Asia with three-dimensional computer simulations

Migrations of the rice planthopper Nilaparvata lugens (Stål) (Delphacidae) from the Philippines to Taiwan, southern China, and southern Japan were analyzed using three-dimensional migration simulations. The results strongly suggested that the Southeast Asian population of N. lugens mixes with the East Asian population. This highlighted the possibility that planthoppers from the Southeast Asian population, which have properties different from those in the East Asian population such as feeding of resistant rice varieties and wing polymorphism, could migrate to Japan via southern China and Taiwan. This study, therefore, emphasizes the special care that should be taken to monitor the properties of immigrants to Japan.     

十六个水稻品种(系)对褐飞虱的抗虫性评价

本文通过苗期抗性鉴定、田间抗虫性鉴定、稻田节肢动物功能团优势度比较及产量测定的方法来评价16个水稻品种(系)对褐飞虱Nilaparvata lugens(Stl)的抗性。结果表明,有12个品种(系)对褐飞虱表现为抗级,2个品系表现为中抗。这14个抗虫-中抗的品种(系)在成株期也表现一定的抗虫性,且与对照TN1之间差异显著。本试验条件下,广占63-3S/华恢15、广两优106、Y58S/华15、广两优476、广占63-4S/华恢15这5个品种(系)不仅抗虫性好,而且产量也较高,分别比TN1和主栽品种扬两优6号高63.49%～66.57%和3.47%～11.57%,具有广泛的开发应用前景。     

褐飞虱在香稻上的发生与产卵特征

研究褐飞虱NilaparvatalugensStl在香稻上的危害与产卵特征,结果表明香稻田中褐飞虱的自然种群密度基数远高于非香型稻。从7月22日至8月31日香稻田中褐飞虱的自然种群密度一直增长,在武香988和鄂香1号田间的若虫分别从7月22日的7头丛和3头丛上升至8月31日的126头丛和73头丛。香型稻田间的褐飞虱的自然种群密度增长慢于非香型稻田间。香稻与非香稻田间褐飞虱成虫出现的高峰日相差不大。香稻田中褐飞虱产卵株率高于非香型稻田,其中最高的为鄂香1号,达到74.27%。香稻田间株平均产卵块数几乎为非香型稻上的2倍,但是香稻上的卵死亡率(鄂香1号为39.24%)显著高于非香型稻上的(鄂中5号为27.58%,两优培九为27.21%),且产卵部位有一定差异。鄂香1号第Ⅲ叶鞘上卵块最多,占总数的60.98%,武香988的第Ⅱ和第Ⅳ叶鞘上产卵较多,第Ⅰ叶鞘上没有稻飞虱产卵。非香型稻鄂中5号上产卵最多的是第Ⅳ叶鞘,第Ⅰ和第Ⅱ叶鞘上没有褐飞虱产卵。两优培九除第Ⅰ和第Ⅴ叶鞘上没有褐飞虱产卵外,第Ⅱ、第Ⅲ和第Ⅳ叶鞘上均有褐飞虱产卵,而且卵量差异不大。     

Quantitative genetic analysis of biotypes of the brown planthopper Nilaparvata lugens: heritability of virulence to resistant rice varieties

The virulence of Nilaparvata lugens Stål (Homoptera: Delphacidae) to a resistant variety of rice, Oryza sativa L., is suggested to be under polygenic control. To determine whether the virulence of N. lugens is a quantitative character that has continuous distribution or a threshold character that has a few discontinuous phenotypic forms and is determined by some underlying continuous variable, I examined the frequency distribution of honeydew excretion that has been used as a measure of ability of attacking the rice plants, and the relationship between honeydew excretion and the oviposition rate of N. lugens females using a japonica rice line Saikai 190 which has a resistance gene Bph 1. The frequency distribution in honeydew excretion significantly differed from the normal distribution, but two distributions below and above 10 mg honeydew excretion for two days did not significantly deviate from normality, suggesting a bimodal distribution. There were significant differences in the proportion of females ovipositing and the number of eggs deposited between the female groups that excreted 0–10 mg honeydew and more than 20 mg honeydew. Within these female groups, however, the reproductive performances were not different. Thus, virulence of N. lugens can be analyzed as a threshold character that has two distinct phenotypes, virulent and avirulent. I estimated the heritabilities of the virulence of N. lugens by parent-offspring regression using the percentage of virulent females in a full-sib family. The estimates of heritability were 0.41 on Saikai 190 and 0.55 on ASD7, carrying the bph 2 gene. These results show that the N. lugens population has substantial genetic variation in virulence. The regression coefficients on female and male parents were similar, suggesting similar genetic contributions by both parents. When the parental families were examined on Saikai 190 and the progeny on ASD7, the regression coefficient approached zero. There may be a low genetic correlation between virulences to the two varieties.     

Effects on the feeding behaviour of Nilaparvata lugens (Stål) of sublethal concentrations of the foliarly applied nitromethylene heterocycle 2-nitromethylene-1, 3-thiazinan-3-yl-carbamaldehyde

Abstract. An electrical recording technique, combined with simultaneous videofilming, was used to investigate the detailed feeding behaviour of Nilaparvata lugens (Stål) (Homoptera: Delphacidae) on rice. Phases of brief probing, of non-feeding activities, such as walking and of sustained feeding were observed. Monitoring of electrical signals recorded during long, uninterrupted periods of contact between the insect and its host plant, revealed three pattern classes. Two clearly distinguishable wave-like patterns represent the hopper sucking from xylem and phloem tissues. The third represents a 'complex' of signals believed to be associated with stylet penetration, salivation and sucking activities in non-vascular tissues of the host plant. This recording method and simple counts of honeydew excretion activity were used to assess the effect on the feeding behaviour of N. lugens of a sublethal concentration of the nitromethylene heterocycle insecticide 2-nitromethylene-l, 3-thiazinan-3-yl-carbamaldehyde applied to rice. The treatment caused a general reduction in feeding activity. Analysis of the electrical trace revealed that phloem feeding was strongly inhibited. Since these effects were largely reversed on transferring hoppers from treated to untreated hosts, the test compound was considered to have an antifeedant rather than a conventional poisoning effect on the hoppers when applied at sublethal doses.