13 resultados para Ephestia Kuhniella


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The innate immune system of insects consists of humoral and cellular components involved in the recognition of and responses to intruding foreign micro- or macroorganisms. Several molecules have been identified so far that recognize molecular patterns present on microorganisms, such as lipopolysaccharides, peptidoglycans and lipoteichonic acid. These molecules, acting as opsonins, trigger immune responses such as phagocytosis, nodule formation, melanization and encapsulation. Here, we investigated the role of calreticulin (CRT) present on the surface of Pieris rapae hemocytes in phagocytosis. Comparative phagocytosis assays using yeast cells showed that hemocytes from different insects exhibit significant variation in their phagocytosing potential and relative CRT involvement. (C) 2003 Elsevier Science Ltd. All rights reserved.

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Fatores de risco que afetam a produção de sementes em Ipomoea pes-caprae (L.) R. Br. (Convolvulaceae) foram avaliados em 10 praias da Ilha de Santa Catarina, SC. Ipomoea pes-caprae é auto-incompatível, possui longos estolões, com difícil individualização dos genetas em campo. Assim, manchas de I. pes-caprae foram monitoradas, sendo definidas como áreas ocupadas pela espécie, descontínuas entre si. Nestas, 9,4% a 76,0% das inflorescências estavam abortadas. Soterramentos causaram perda de botões e frutos, mas a ineficiência de polinização parece explicar a grande taxa de abortos após a floração. A predação de sementes pelas larvas dos bruquídeos Megacerus baeri e M. reticulatus e por lagartas da mariposa Ephestia kuhniella foi expressiva. Os bruquídeos ovipõem preferencialmente em frutos em amadurecimento, danificando até 65,7% das sementes produzidas nas manchas. Ephestia kuhniella danificou até 57,4% das sementes em 1996 e até 5,4% em 1997, ano com menor abundância de frutos. Frutos com mariposa não abrem para dispersão, permanecendo três meses e meio presos à planta. As sementes intactas destes frutos são viáveis e com sobrevivência de plântulas similar às provindas de frutos sem infestação. Os danos registrados podem reduzir as sementes viáveis a 9% do total produzido, estimando-se de 0,5 a 30,6 sementes viáveis m-2 nas manchas estudadas. As maiores densidades de sementes viáveis ocorreram nas manchas com mais frutos, reforçando como vantajosa, uma alta produção de frutos. A baixa densidade de sementes viáveis encontradas em algumas localidades talvez possa comprometer a regeneração local da espécie.

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The use of Bacillus thuringiensis (Bt) endotoxins to control insect vectors of human diseases and agricultural pests is threatened by the possible evolution of resistance in major pest species. In addition to high levels of resistance produced by receptor insensitivity (5, 16, 17), several cases of tolerance to low to medium levels of toxin have been reported in laboratory colonies of lepidopteran species (3, 18). Because the molecular basis of some of these cases of tolerance to the toxin are not known, we explored alternative mechanisms. Here, we present evidence that tolerance to a Bt formulation in a laboratory colony of the flour moth Ephestia kuehniella can be induced by preexposure to a low concentration of the Bt formulation and that the tolerance correlates with an elevated immune response. The data also indicate that both immune induction and Bt tolerance can be transmitted to offspring by a maternal effect and that their magnitudes are determined by more than one gene.

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A nested polymerase chain reaction protocol yielded positive detection of the maternally inherited cytoplasmic proteobacterium Wolbachia in total genomic DNA from coffee berry borers collected in Benin, Brazil, Colombia, Ecuador, El Salvador, Honduras, India, Kenya, Mexico, Nicaragua, and Uganda. Wolbachia was not detected in specimens from Cameroon, the Dominican Republic, Indonesia, Jamaica, and Peru. Amplified bands from India and Brazil were cloned and sequenced. The 438-bp sequence clearly conformed to Wolbachia group B and was nearly identical to that of Ephestia kuehniella. The possible implications of Wolbachia infection in the coffee berry borer are discussed.

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Dissertação de Mestrado, Biotecnologia em Controlo Biológico, 27 de Junho de 2013, Universidade dos Açores.

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The aim of this contract was to finalise in vitro rearing on artificial diets of entomophagous insects useful to control insect pests mainly in greenhouses through an analytical and rational approach. The work focuses on the development and optimisation of artificial diets for one coccinellid (Harmonia axyridis), two heteropteran predators (Orius laevigatus, Dicyphus tamaninii), and a braconid parasitoid of aphids (Aphidius ervi). The definition of the artificial diets was based on biochemical analyses of their natural food (aphids) or substitution food for the predators (Ephestia kuehniella eggs). Reliable quality control parameters of the entomophages produced were used in order to adjust dietary composition and formulation of the different diets tested.

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BACKGROUND: The use of Trichogramma species is a potential key strategy in integrated pest management. However, its effectiveness depends on the use of chemicals that do not interfere with parasitism and parasite population growth. Here, a study was made of the effects of synthetic insecticides on Trichogramma pretiosum and Trichogramma exiguum in different hosts (Ephestia kuehniella, Plutella xylostella and Spodoptera frugiperda) and the influence of International Organisation for Biological Control (IOBC/WPRS) methodology in selectivity studies using different Trichogramma species. The insecticides used were commercial formulations (triflumuron at a concentration of 0.2 mL L-1 water, etofenprox at a concentration of 0.47 mL L-1 water and endosulfan at a concentration of 7.5 mL L-1 water); the control treatment consisted of distilled water. Eggs attached to cardboard cards were offered to parasitoids inside glass cages. Parasitised eggs, parasitism and adult emergence rates and parasitism reduction were evaluated.RESULTS: Endosulfan and etofenprox, classified as class-4 toxic products, were extremely toxic to the parasitoids. Triflumuron, classified as a non-toxic product, was selective to the parasitoids in eggs of all hosts.CONCLUSIONS: The methodology recommended by IOBC/WPRS influenced results regarding the use of different species of parasitoids, and the use of a single parasitoid species in their experiment is questionable. (C) 2011 Society of Chemical Industry

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The objective of this study was to determine the influence of Metarhizium anisopliae on the predatory capacity and functional response of Chrysoperla externa fed Bemisia tabaci B biotype rearing cotton. The stock rearings of B. tabaci B biotype and C. externa were maintained in cages according to preexisting methodologies. Upon hatching, predator larvae were fed on eggs of Ephestia küehniella until their 3rd instar and then fed on whitefly nymphs in density exceeding their intake capacity (150 nymphs/individual). The nymphs were offered on cotton leaf disks (5 cm diameter), 24 hours after the treatments application following: T1: Control (water); T2: M. anisopliae at the concentration of 105 conidia cm-3; T3: 10 8 conidia cm-3 origined from Metharril® formulation of the 109 viable conidia cm-3 M. anisopliae active principle. The predatory capacity was obtained by the difference between the numbers of nymphs supplied and nymphs left. Five different nymphal densities (130, 160, 190, 220, and 250) were selected for the functional response study. The number of nymphs preyed was recorded after 24, 48, 72, and 96 hours. The M. anisopliae entomopathogenic fungi did not influence predatory capacity of C. externa which ranged from 70.6 individuals at the highest concentration to 87.1 in the control. Conclued that the C. externa predatory capacity was not affected by fungi M. anisopliae concentrations. The predator showed a type-II functional response after 24 hours, and a type-I response after 48, 72s, and 96 hours independent from fungi concentrations.

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Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

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El 1 de enero de 2014 entró en vigor la Directiva Europea 2009/128/CE sobre uso sostenible de plaguicidas y el Real Decreto 1311/2012 por el cual se traspone dicha normativa comunitaria al ámbito nacional. Estos reglamentos establecen el marco legal por el que las explotaciones agrícolas deben cumplir los principios generales de la Gestión Integrada de Plagas (GIP). Los principios de la GIP dan preferencia a aquellos métodos de control que sean sostenibles y respetuosos con el medio ambiente, dando prioridad al control biológico, al físico y a otros de carácter no químico. Sin embargo, el uso de insecticidas selectivos con los enemigos naturales es necesario en ocasiones para el adecuado manejo de las plagas en cultivos hortícolas. Por ello, el objetivo general de esta Tesis ha sido aportar conocimientos para la mejora del control de plagas en cultivos hortícolas, mediante la integración de estrategias de lucha biológica, física y química. La primera de las líneas de investigación de esta Tesis se centró en el estudio del efecto de la presencia dos depredadores, larvas Chrysoperla carnea y adultos de Adalia bipunctata, en la dispersión del virus de transmisión no persistente Cucumber mosaic virus (CMV) y del virus de transmisión persistente Cucurbit aphid-borne yellows virus (CABYV), transmitidos por el pulgón Aphis gosypii en cultivo de pepino. La tasa de transmisión de CMV fue baja para los dos tiempos de evaluación ensayados (1 y 5 días), debido al limitado movimiento de su vector A. gossypii. Las plantas que resultaron infectadas se localizaron próximas a la fuente de inóculo central y la presencia de ambos enemigos naturales no incrementó significativamente el porcentaje de plantas ocupadas por pulgones ni la tasa de transmisión de CMV. Los patrones de distribución de A. gossypii y de CMV tan solo fueron coincidentes en las proximidades de la planta central infectada en la que se liberaron los insectos. En los ensayos con CABYV, la presencia de C. carnea y de A. bipunctata respectivamente provocó un incremento significativo de la dispersión de A. gossypii tras 14 días, pero no tras 7 días desde la liberación de los insectos. La reducción en el número inicial de pulgones en la planta central infectada con CABYV fue siempre mayor tras la liberación de C. carnea en comparación con A. bipunctata. Sin embargo, la tasa de transmisión de CABYV y su distribución espacial no se vieron significativamente modificadas por la presencia de ninguno de los depredadores, ni tras 7 días ni tras 14 días desde el inicio de los ensayos. Al igual que se estudió el efecto de la presencia de enemigos naturales en el comportamiento de las plagas y en la epidemiología de las virosis que transmiten, en una segunda línea de investigación se evaluó el posible efecto del consumo de pulgones portadores de virus por parte de los enemigos naturales. Este trabajo se llevó a cabo en el Laboratorio de Ecotoxicología del Departamento de Entomología de la Universidade Federal de Lavras (UFLA) (Brasil). En él se evaluó la influencia en los parámetros biológicos del enemigo natural Chrysoperla externa al alimentarse de Myzus persicae contaminados con el virus de transmisión persistente Potato leafroll virus (PLRV). El consumo de M. persicae contaminados con PLRV incrementó significativamente la duración de la fase larvaria, reduciendo también la supervivencia en comparación a otras dos dietas a base de M. persicae no contaminados con el virus y huevos del lepidóptero Ephestia kuehniella. La duración de la fase de pupa de C. externa no difirió significativamente entre las dietas a base de pulgones contaminados con PLRV y pulgones no contaminados, pero ambas fueron menores que con la dieta con huevos de E. kuehniella. Sin embargo, ni la supervivencia en la fase de pupa ni los parámetros reproductivos de los adultos emergidos mostraron diferencias significativas entre las dietas evaluadas. Por el contrario, la supervivencia de los adultos durante los 30 primeros días desde su emergencia sí se vio significativamente afectada por la dieta, siendo al término de este periodo del 54% para aquellos adultos de C. externa que durante su fase larvaria consumieron pulgones con PLRV. Dentro de la GIP, una de las estrategias de carácter físico que se emplean para el control de plagas y enfermedades en cultivos hortícolas protegidos es el uso de plásticos con propiedades fotoselectivas de absorción de la radiación ultravioleta (UV). Por ello, la tercera línea de investigación de la Tesis se centró en el estudio de los efectos directos e indirectos (mediados por la planta) de condiciones especiales de baja radiación UV sobre el crecimiento poblacional del pulgón A. gossypii y los parámetros biológicos del enemigo natural C. carnea, así como sobre las plantas de pepino en las que se liberaron los insectos. Los ensayos se realizaron en jaulones dentro de invernadero, utilizándose en el primero de ellos plantas de pepino sanas, mientras que en el segundo las plantas de pepino fueron previamente infectadas con CABYV para estudiar de qué manera afectaba la incidencia del virus en las mismas condiciones. Las condiciones de baja radiación UV (bajo plástico Térmico Antivirus®) ejercieron un efecto directo en las fases iniciales del cultivo de pepino, promoviendo su crecimiento, mientras que en fases más avanzadas del cultivo indujeron un aumento en el contenido en nitrógeno de las plantas. Las plantas de pepino que fueron sometidas a mayor intensidad de radiación UV (bajo plástico Térmico Blanco®) al inicio del cultivo mostraron un engrosamiento significativo de las paredes de las células epidérmicas del haz de las hojas, así como de la cutícula. El uso del plástico Térmico Antivirus®, utilizado como barrera fotoselectiva para crear condiciones de baja radiación UV, no alteró con respecto al plástico Térmico Blanco® (utilizado como control) el desarrollo poblacional del pulgón A. gossypii ni los parámetros biológicos evaluados en el depredador C. carnea. En el segundo experimento, realizado con plantas infectadas con CABYV, la incidencia de la virosis enmascaró las diferencias encontradas en experimento con plantas sanas, reduciendo aparentemente la influencia de las distintas condiciones de radiación UV. Por último, para el desarrollo de las estrategias de GIP es importante estudiar los posibles efectos secundarios que los plaguicidas pueden tener en los enemigos naturales de las plagas. Es por ello que en la Tesis se evaluaron la toxicidad y los efectos subletales (fecundidad y fertilidad) de flonicamida, flubendiamida, metaflumizona, spirotetramat, sulfoxaflor y deltametrina en los enemigos naturales C. carnea y A. bipunctata. Los efectos secundarios fueron evaluados por contacto residual tanto para larvas como para adultos de ambos enemigos naturales en condiciones de laboratorio. Flonicamida, flubendiamida, metaflumizona y spirotetramat fueron inocuos para larvas de último estadio y adultos de C. carnea y A. bipunctata. Por este motivo, estos insecticidas se presentan como buenos candidatos para ser incorporados dentro de programas de GIP en combinación con estos enemigos naturales para el control de plagas de cultivos hortícolas. Sulfoxaflor fue ligeramente tóxico para adultos de C. carnea y altamente tóxico para larvas de último estadio de A. bipunctata. Para A. bipunctata, sulfoxaflor y deltametrina fueron los compuestos más dañinos. Deltametrina fue también el compuesto más tóxico para larvas y adultos de C. carnea. Por tanto, el uso de deltametrina y sulfoxaflor en programas de GIP debería tomarse en consideración cuando se liberasen cualquiera de estos dos enemigos naturales debido al comportamiento tóxico que mostraron en condiciones de laboratorio. ABSTRACT On 1 January 2014 came into effect the Directive 2009/128/EC of the European Parliament about sustainable use of pesticides and the Royal Decree 1311/2012 that transposes the regulation to the Spanish level. These regulations establish the legal framework that agricultural holdings must adhere to in order to accomplish the general principles of Integrated Pest Management (IPM). The guidelines of IPM give priority to sustainable and eco-friendly pest control techniques, such as biological and physical measures. Nevertheless, the use of pesticides that are selective to natural enemies is sometimes a necessary strategy to implement accurate pest management programs in horticultural protected crops. Therefore, the general objective of this Thesis was to contribute to the improvement of pest management strategies in horticultural crops, by means of the integration of biological, physical and chemical techniques. The first research line of this Thesis was focused on the evaluation of the effects of two aphidophagous predators, Chrysoperla carnea larvae and Adalia bipunctata adults, on the spread of the non-persistently transmitted Cucumber mosaic virus (CMV, Cucumovirus) and the persistently transmitted Cucurbit aphid-borne yellows virus (CABYV, Polerovirus), by the aphid vector Aphis gossypii in a cucumber crop under greenhouse conditions. The CMV transmission rate was generally low, both after 1 and 5 days, due to the limited movement of its aphid vector A. gossypii. Infected plants were mainly located around the central virusinfected source plant, and the percentage of aphid occupation and CMV-infected plants did not differ significantly in absence and presence of natural enemies. The distribution patterns of A. gossypii and CMV were only coincident close to the central plant where insects were released. In the CABYV experiments, the presence of C. carnea larvae and A. bipunctata adults induced significant A. gossypii dispersal after 14 days but not after 7 days. The reduction in the initial aphid population established in the central plant was always higher for C. carnea than for A. bipunctata. Nevertheless, CABYV spread was not significantly modified by the presence of each predator either in the short term (7 days) or in the long term (14 days). Furthermore, the percentage of CABYV-infected plants did not significantly differ when each natural enemy was present in any evaluation period. It is important to evaluate the influence that natural enemies have on pest dynamics and on the spread of viral diseases, but it should be also taken into account the possible effect on the performance of natural enemies when they feed on preys that act as vectors of viruses. Thus, in a second research line developed in the Laboratory of Ecotoxicology, Department of Entomology, of the Universidade Federal de Lavras (UFLA) (Brazil), it was evaluated the performance of Chrysoperla externa under the condition of consuming Myzus persicae acting as vector of Potato leafroll virus (PLRV). The diet composed of PLRV-infected M. persicae significantly increased the length and reduced the survival rate, of the larval period in regard to the other two diets, composed of non-infected M. persicae and Ephestia kuehniella eggs. The lengths of the pupal stage were not significantly different between the aphid diets, but both were significantly shorter than that of E. kuehniella eggs. Neither pupal survival nor reproductive parameters revealed significant differences among the diets. Nevertheless, the adult survival curves during the first 30 days after emergence showed significant differences, reaching at the end of this interval a value of 54% for those C. externa adults fed on PLRVinfected aphids during their larval period. According to the IPM guidelines, one of the physical strategies for the control of pests and diseases in horticultural protected crops is the use of plastic films with photoselective properties that act as ultraviolet (UV) radiation blocking barriers. In this sense, the third research line of the Thesis dealt with the study of the direct and plant-mediated influence of low UV radiation conditions on the performance of the aphid A. gossypii and on the biological parameters of the natural enemy C. carnea, as well as on the cucumber plants where insects were released. The experiments were conducted inside cages under greenhouse conditions, using for the first one healthy cucumber plants, while for the second experiment the cucumber plants were previously infected with CABYV in order to assess the influence of the virus in the same conditions. The low UV radiation conditions (under Térmico Antivirus® plastic film) seemed to exert a direct effect in the early stages of cucumber plants, enhancing their growth, and in an increasing nitrogen content at further developmental stages. The higher UV radiation exposure (under Térmico Blanco® plastic film) in the early stages of the cucumber crop induced the thickening of the adaxial epidermal cell walls and the cuticle of leaves. The use of Térmico Antivirus® plastic film as a photoselective barrier to induce low UV radiation conditions did not modify, in regard to Térmico Blanco® plastic film (used as control), neither the population development of A. gossypii nor the studied biological parameters of the predator C. carnea. In the second experiment, done with CABYV-infected cucumber plants, the incidence of the virus seemed to mask the direct and plant-mediated influence of the different UV radiation conditions. In last term, for the development of IPM strategies it is important to study the potential side effects that pesticides might have on natural enemies. For this reason, in the Thesis were tested the toxicity and sublethal effects (fecundity and fertility) of flonicamid, flubendiamide, metaflumizone, spirotetramat, sulfoxaflor and deltamethrin on the natural enemies C. carnea and A. bipunctata. The side effects of the active ingredients of the insecticides were evaluated with residual contact tests for the larvae and adults of these predators under laboratory conditions. Flonicamid, flubendiamide, metaflumizone and spirotetramat were innocuous to last instar larvae and adults of C. carnea and A. bipunctata. Therefore, these pesticides are promising candidates for being incorporated into IPM programs in combination with these natural enemies for the control of particular greenhouse pests. In contrast, sulfoxaflor was slightly toxic to adults of C. carnea and was highly toxic to last instar larvae of A. bipunctata. For A. bipunctata, sulfoxaflor and deltamethrin were the most damaging compounds. Deltamethrin was also the most toxic compound to larvae and adults of C. carnea. In accordance with this fact, the use of sulfoxaflor and deltamethrin in IPM strategies should be taken into consideration when releasing either of these biological control agents, due to the toxic behavior observed under laboratory conditions.

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A necessidade por recursos alimentares move os insetos predadores na busca por presas, para garantir a sobrevivência. Existem vários fatores que podem influenciar o comportamento de busca e escolha por presas e, sob determinadas circunstâncias, o inseto pode escolher se alimentar de presas não alvo, como, por exemplo, inimigos naturais e assim prejudicar a população de organismos benéficos. Garantir a estabilidade do agroecossistema é uma forma de manter as populações de pragas em níveis compatíveis com a ação de inimigos naturais e, consequentemente obter o sucesso no controle biológico. Programas bem sucedidos de manejo de pragas requerem previamente a realização de estudos sobre comportamento de insetos. Com o intuito de compreender os fatores que governam o comportamento predatório e as interações ecológicas entre Harmonia axyridis e Chrysoperla externa, considerando a presença de Diaphorina citri como presa, objetivou-se estudar o efeito de interações intraguilda sobre a predação de D. citri, experimentalmente e utilizando modelos estatísticos para compreender a dinâmica de interações tróficas no contexto de potenciais guildas presentes em citros. Três experimentos distintos foram realizados para investigar o comportamento dos insetos predadores. Foram realizados testes com escolha entre larvas dos predadores C. externa e H. axyridis e testes com e sem escolha comparando machos e fêmeas do predador H. axyridis, a fim de investigar padrões de escolha de presas. O segundo experimento foi realizado para investigar o comportamento predatório entre larvas de segundo instar dos predadores sob diferentes densidades de D. citri como presa. As combinações foram: sem escolha de presas intraguilda, combinações com escolha de presas intraguilda e combinações com a presença de dois predadores da mesma espécie para possibilitar o canibalismo. As densidades de D. citri utilizadas foram 5, 10, 15, 30, 60, 80. No último experimento avaliou-se a interação entre larvas de C. externa e H. axyridis, ao longo do desenvolvimento, simulando a existência de apenas uma presa, como fonte de alimento. O desenvolvimento dos predadores foi avaliado e comparado utilizando-se duas presas diferentes, ninfas de D. citri e ovos de Ephestia kuehniella para cada larva de primeiro instar dos predadores C. externa e H. axyridis. As combinações de predadores foram as mesmas citadas no experimento anterior. Observou-se a duração do desenvolvimento larval nas diferentes combinações de predadores e presa, bem como o percentual de predações intraguilda e canibalismos ocorridos. Nem a densidade de presas, nem o tipo de presa ofertada influenciou o comportamento das larvas dos predadores para a realização de predação intraguilda ou canibalismo. O principal fator foi a diferença de idade que reflete no tamanho dos predadores, podendo direcionar a predação intraguilda ou o canibalismo e até mesmo acelerar o ciclo de desenvolvimento do predador intraguilda envolvido. Predadores em estágio inicial de vida buscaram por qualquer tipo de presa para garantir sua sobrevivência, não demonstrando qualquer padrão em suas escolhas. No caso dos adultos de H. axyridis, há diferença de comportamento entre os sexos, dependendo da densidade de presas disponíveis, fêmeas poderão ser mais ágeis na busca e consumo de presas.

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Parasitoid wasps use a variety of mechanisms to alter their host's physiology to the benefit of the developing endoparasite inside the host larva. Association of certain wasps with viruses and virus-like particles (VLPs) that contribute to their success in parasitism is one of the fascinating evolutionary adaptations conferring active or passive protection for the endoparasite from the host immune system. Venturia canescens has been shown to produce VLPs that provide protection for the developing parasitoid egg inside the host, Ephestia kuehniella. Here, we report on the presence of a novel small RNA-containing virus from V. canescens, designated as VcSRV, occurring in the ovaries of the wasp. The virus particles are found together with VcVLPs in the lumen of the calyx region of the ovaries and are injected together with the egg and VcVLPs into E kuehniella larvae where they enter hemocytes. Alignment of the RNA-dependent RNA polymerase gene of VcSRV indicates that the virus most likely belongs to the recently described genus Iflavirus. (c) 2004 Elsevier Ltd. All rights reserved.

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Insect host-parasitoid interactions provide fascinating examples of evolutionary adaptations in which the parasitoid employs a variety of measures and countermeasures to overcome the immune responses of its host. Maternal factors introduced by the female wasps during egg deposition play an important role in interfering with cellular and humoral components of the host's immune defence. Some of these components actively suppress host immune components and some are believed to confer protection for the developing endoparasitoid by rather passive means. The Venturio conescens/Ephestia kuehniella parrositoid-host system is unique among other systems in that the cellular defence capacity of the host remains virtually intact after parasitization. This system raises some important questions that are discussed in this mini-review: If immune protection of the egg and the emerging larva is achieved by surface properties comprising glycoproteins and virus-like particles (VLPs) produced by the female wasp, why is the prophenoloxidose activating cascade blocked in parasitized caterpillars? Another question is the evolutionary origin of these particles, given that the functional role and structural features of V. canescens VLP proteins are more related to cellular proteins than to viruses.