Biological characteristics and thermal requirements of a Brazilian strain of the parasitoid Trichogramma pretiosum reared on eggs of Pseudoplusia includens and Anticarsia gemmatalis

A new strain of the parasitoid Trichogramma pretiosum, was collected in Rio Verde County, State of Goiás, Central Brazil, and designated as T. pretiosum RV. This strain was then found to be the most effective one among several different strains of T. pretiosum tested in a parasitoid selection assay. Therefore, its biological characteristics and thermal requirements were studied, aiming at allowing its multiplication under controlled environmental conditions in the laboratory. The parasitoid was reared on eggs of Pseudoplusia includens and Anticarsia gemmatalis at different constant temperatures within an 18–32 °C temperature range. The number of annual generations of the parasitoid was also estimated at those temperatures. Results have shown that T. pretiosum RV developmental time, from egg to adult, was influenced by all temperatures tested within the range, varying from 6.8 to 20.3 days and 6.0 to 17.0 days on eggs of P. includens and A. gemmatalis, respectively. The emergence of T. pretiosum RV from eggs of A. gemmatalis was higher than 94% at all temperatures tested. When this variable was evaluated on eggs of P. includens, however, the figures were higher than that within the 18–30 °C range (more than 98%), and were also statistically higher than the emergence observed at 32 °C (90.2%). The sex ratio of the parasitoids emerged from eggs of A. gemmatalis decreased from 0.55 to 0.29 at 18–32 °C, respectively. However, for those emerged from eggs of P. includens, the sex ratio was similar (0.73, 0.72 and 0.71) at 20, 28 and 32 °C, respectively. The lower temperature threshold (Tb) and thermal constant (K) were 10.65 °C and 151.25 degree-days when the parasitoid was reared on eggs of P. includens; and 11.64 °C and 127.60 degree-days when reared on eggs of A. gemmatalis. The number of generations per month increased from 1.45 to 4.23 and from 1.49 to 4.79 when the parasitoid was reared on eggs of P. includens and A. gemmatalis, respectively, following the increases in the temperature.     

Host Preferences of Trichogramma pretiosum and the Influence of Prior Ovipositional Experience on the Parasitism of Plutella xylostella and Pseudoplusia includens Eggs

Trichogramma wasps are generalist egg parasitoids used in biological control efforts. In a multi host situation they may preferentially parasitize a non-target host species to the detriment of the control program. Plutella xylostella Linnaeus (Lepidoptera: Plutellidae) is a very serious pest of cabbage, but is only one in a number of species in the ‘cabbageworm’ complex. We investigated the host preferences of Trichogramma pretiosum Riley (Hymenoptera: Chalcidoidea) when offered the eggs of Plutella xylostella and the eggs of Pseudoplusia includens Hübner (Lepidoptera: Noctuidae), another species in the ‘cabbageworm’ complex. Trichogramma pretiosum reared on the eggs of the factitious host Ephestia kuehniella Zeller (Lepidoptera: Pyralidae) parasitized both Plutella xylostella and Pseudoplusia includens eggs under laboratory conditions. For both choice and no-choice experiments, T. pretiosum parasitized significantly more P. xylostella eggs than P. includens eggs. Prior ovipositional experience with one or other of the two host species had no effect on the subsequent parasitism levels of the two host species. The preference for P. xylostella eggs was also not affected by this prior ovipositional experience.     

Acceptability and suitability of Tuta absoluta eggs from irradiated parents to parasitism by Trichogramma nerudai and Trichogramma pretiosum (Hymenoptera: Trichogrammatidae)

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Cytopathology of the soybean looper,Pseudoplusia includens,infected with the Pseudoplusia includens icosahedral virus

Ultrastructural responses of soybean looper cells of various tissues infected with Pseudoplusia includens icosahedral virus (PIIV), a newly characterized RNA virus [Y. C. Chao, H. A. Scott, and S. Y. Young (1983)J. Gen. Virol.64, 1835–1838], were studied in situ. Most cells of fat body and epidermis consistently contained virus particles and associated cytopathic structures. Virus particles, corresponding to those of purified PIIV in morphology and size, always occurred in the cytoplasm either in membrane-bound virogenic stroma and/or freely in the ground cytoplasm. Virogenic stroma, which appeared to be distinct from those induced by other insect viruses, consisted of electron-dense matrix material, in which virus particles were embedded, and membranous vesicles, 70 or 80 nm in diameter, containing nucleic acid-like fibrils. Virus particles in virogenic stroma occurred as hexagonally arranged crystalline arrays made up primarily of homogeneously dense particles, while those in the ground cytoplasm were dispersed randomly and had an electron-lucent central core. Extremely large numbers of virus particles were also located in noncellular cuticle layers of the integument.     

Nocturnal parasitism of moth eggs by Trichogramma wasps

Parasitoid wasps of the genus Trichogramma are used worldwide as biological control agents against lepidopteran pests. Trichogramma wasps develop inside eggs of a wide range of host species, most of them moths. They are generally considered as diurnal insects. Here, we investigated whether Trichogramma wasps can also successfully parasitise host eggs at night under controlled laboratory conditions. Eggs of the moth Ephestia kuehniella were offered under dark conditions (scotophase) to females of Trichogramma brassicae and Trichogramma evanescens either from 9:00 PM to 9:00 AM or from 11:00 AM to 5:00 PM at four different temperatures (5°C, 10°C, 15°C and 20°C). Both species are known to parasitise E. kuehniella eggs in the photophase during daytime. The results show that T. brassicae did not parasitise eggs in the scotophase at night and only very few in the artificially induced scotophase during daytime from 10°C to 20°C. In contrast, T. evanescens parasitised more eggs in the dark both at night and artificially induced scotophase during daytime. Parasitism in the scotophase already started at 5°C, with more eggs being parasitised and more offspring being produced at higher temperatures. T. evanescens displayed higher parasitism activity in the induced scotophase during daytime than in the scotophase at night. The present study suggests that Trichogramma are capable of successfully parasitising host eggs at night, even at low temperatures, but that nocturnal activity with respect to parasitism varies between wasp species.     

Fertility life table of Trichogramma pretiosum (Hym., Trichogrammatidae) in eggs of Tuta absoluta and Phthorimaea operculella (Lep., Gelechiidae) at different temperatures

Abstract: The aim of this work was to study the reproductive potential of Trichogramma pretiosum reared on Tuta absoluta and Phthorimaea operculella eggs through fertility life tables at different temperatures. The development cycle and the parasitization capacity of this parasitoid was determined in order to calculate the net reproductive rate ( R _o), the infinitesimal increase ratio ( r _m), the finite increase rate ( λ ) and the mean duration of the generation ( T ). The mean duration of one generation of T. pretiosum kept on both eggs was observed to show an inverse relation with the increase of temperature. The net reproduction rate varied according to the temperature variation for both species. The maximum increase in capacity of T. pretiosum on the first host ( T. absoluta ) was reached at 22°C and on the second host ( P. operculella ) between 22 and 25°C. The infinitesimal increase rate and the finite increase rate for both moths had a relationship with the increase of temperature ranging from 18 to 30°C. The highest value of λ for both moths occurred at 30 and 32°C according to the lesser duration of a generation.     

Hematopoiesis in larval Pseudoplusia includens and Spodoptera frugiperda

Maintenance of circulating hemocytes in larval Lepidoptera has been attributed to both mitosis of hemocytes already in circulation and the release of hemocytes from hematopoietic organs. In this study, we compared hematopoiesis in the noctuids Pseudoplusia includens and Spodoptera frugiperda. For both species, hemocyte densities per microl of blood increased with instar. Differential hemocyte counts indicated that plasmatocytes were the most abundant hemocyte type during early instars but granular cells were the most abundant hemocyte type in the last instar. Hematopoietic organs were located in the meso- and metathorax of S. Frugiperda and P. Includens. These organs contained large numbers of hemocytes in S. Frugiperda, but contained few hemocytes in P. Includens. The majority of the hemocytes recovered from hematopoietic organs were identified as plasmatocytes. Using hemocyte type-specific markers and bromodeoxyuridine (BrdU) incorporation experiments, we determined that all hemocyte types with the exception of oenocytoids synthesize DNA. BrdU labeling indices for both species also fluctuated with the molting cycle. Ligation experiments suggested that hematopoietic organs are an important source of circulating plasmatocytes in S. Frugiperda but not in P. Includens. Injection of heat killed bacteria into larvae induced higher levels of BrdU labeling than injection of sterile saline, suggesting that infection and wounding induce different levels of hemocyte proliferation. Arch.     

Pseudoplusia includens Densovirus Genome Organization and Expression Strategy

The genome of a densovirus of a major phytophagous pest, Pseudoplusia includens, was analyzed. It contained 5,990 nucleotides (nt) and included inverted terminal repeats of 540 nt with terminal Y-shaped hairpins of 120 nt. Its DNA sequence and ambisense organization with 4 typical open reading frames demonstrated that it belonged to the genus Densovirus in the subfamily Densovirinae of the family Parvoviridae.     

The genetics and evolution of obligate reproductive parasitism in Trichogramma pretiosum infected with parthenogenesis-inducing Wolbachia

Parthenogenesis-inducing (PI) Wolbachia belong to a class of intracellular symbionts that distort the offspring sex ratio of their hosts toward a female bias. In many PI Wolbachia-infected species sex ratio distortion has reached its ultimate expression-fixation of infection and all-female populations. This is only possible with thelytokous PI symbionts as they provide an alternative form of reproduction and remove the requirement for males and sexual reproduction. Many populations fixed for PI Wolbachia infection have lost the ability to reproduce sexually, even when cured of the infection. We examine one such population in the species Trichogramma pretiosum. Through a series of backcrossing experiments with an uninfected Trichogramma pretiosum population we were able to show that the genetic basis for the loss of female sexual function could be explained by a dominant nuclear effect. Male sexual function had not been completely lost, though some deterioration of male sexual function was also evident when males from the infected population (created through antibiotic curing of infected females) were mated to uninfected females. We discuss the dynamics of sex ratio selection in PI Wolbachia-infected populations and the evolution of non-fertilizing mutations.     

Regulation of melanization by glutathione in the moth Pseudoplusia includens

The phenoloxidase (PO) cascade regulates the melanization of hemolymph, which serves as a conserved humoral immune response in insects and other arthropods. The reductant glutathione (GSH) has long been used to inhibit melanization of hemolymph from insects but whether GSH levels in hemolymph are sufficient to play a physiological role in regulating melanization is unknown. Here, we characterized the abundance and effects of GSH on the melanization of plasma from larval stage Pseudoplusia includens (Lepidoptera: Noctuidae). GSH concentration in newly collected plasma from day two fifth instars ranged from 50 to 115 μM, while the titer of tyrosine, a substrate for the PO cascade, was 141 μM. GSH titers rapidly declined in plasma after collection from larvae, but no melanin formation occurred until GSH levels fell below 20 μM. Added GSH dose-dependently blocked melanization while PO substrates overrode GSH inhibition. Experiments conducted in the absence of oxygen and presence of PO cascade inhibitors further suggested that depletion of GSH from plasma was primarily due to formation of reactive intermediates produced by activated PO. Additional studies identified hemocytes as a potential source of plasma GSH. Hemocyte lysates recycled oxidized glutathione (GSSG) into GSH using NADPH, while intact hemocytes released GSH into the medium. These results suggest that in addition to protease cascade-releated mechanisms that regulate phenoloxidase, GSH exerts another level of control on melanization of insect hemolymph.     

拟澳洲赤眼蜂和短管赤眼蜂在米蛾多卵上的种间竞争

1引言 小菜蛾(Plutella xylostella)是十字花科蔬菜的一种重要害虫.20世纪70年代以来,化学杀虫剂的大量使用和抗性的产生.使小菜蛾成为南亚、东南亚和中国南方蔬菜生产中最为严重的害虫[8,22].因此,寻求安全有效的生物防治措施是防治小菜蛾的必要途径[23].赤眼蜂(Trichogramma)是一类多食性的卵寄生蜂,近百年来已被成功地用于防治为害多种作物及森林的害虫[21],但研究和利用赤眼蜂防治小菜蛾只有10来年的历史[12,23].     

Ultrastructure of the natural and factitious host eggs of Trichogramma galloi Zucchi and Trichogramma pretiosum Riley (Hymenoptera: Trichogrammatidae)

The surface and structure of the chorion of eggs of Diatraea saccharalis (F.) (Lepidoptera: Crambidae), Anticarsia gemmatalis (Huebner), Heliothis virescens F., Spodoptera frugiperda (Smith) (Lepidoptera: Noctuidae), Sitotroga cerealella (Oliver) (Lepidoptera: Gelechiidae), Ephestia kuehniella Zeller and Corcyra cephalonica Stainton (Lepidoptera: Pyralidae), that are hosts of Trichogramma galloi Zucchi and Trichogramma pretiosum Riley (Hymenoptera: Trichogrammatidae) were studied on SEM and TEM. Other characteristics of these eggs, such as changes in their color during embryonic development, size and volume were also recorded. Sculpturing and texture of the surface of the chorion greatly varied among the species studied, as well as the number of layers of the chorion and their thickness. Eggs of the factitious hosts were among the smallest and their volume was very close to each other. All these characteristics would provide basic information for a better understanding of the host selection behavior and are useful for the development of a suitable artificial host egg for the in vitro rearing of these egg parasitoids.     

拟澳洲赤眼蜂和短管赤眼蜂在米蛾卵上的种间竞争

By means of single and mixed inoculation, this paper studied the interspecific competition between T. confusum and T. pretiosum on the factitious eggs of Corcyra cephaloica under different parasitoid densities, host densities and inoculated spaces. The results showed that for both singly and mixed inoculated groups, the parasitism increased with parasitoid density but decreased with host density, whereas the percentage of female progeny dropped with parasitoid density but enhanced with host density. No significant effect was observed on adult emergence for all parasitoid and host density treatments. In mixed inoculated group, the proportion of T. pretiosum in the progeny decreased with parasitoid density but increased with host density, and was more than 50% in all treatments, indicating that T. pretiosum had a stronger competitive ability than T. confusum. In the spaces ranging from 4cm^3 to 102cm^3, the parasitism decreased gradually in both singly and mixed inoculated groups. The percentage of female progeny and adult emergence had no significant difference among different inoculated spaces. In mixed inoculated group, the proportion of T. pretiosum in the progeny was more than 50% in all treatments except space 102 cm3, but decreased with space, which suggested that T. confusum could improve their competitive ability through increasing their search areas and looking for more hosts.     

Parasitism of cotton leafworm Alabama argillacea eggs by Trichogramma pretiosum in commercial cotton fields

Release of natural enemies in commercial fields is challenging and has been inconsistent in the results achieved. This work discusses the augmentative releases of Trichogramma pretiosum to control the cotton leafworm (CLW) Alabama argillacea and also examines the parasitoid–host interaction under grower field conditions. The treatments consisted of fields with and without releases of T. pretiosum set up in Primavera do Leste and Campo Verde Counties, MT, Brazil, during three different seasons (2003 and 2004 dry and 2004 regular summer seasons). Trichogramma wasps were weekly released in the treated fields throughout the entire sampling period (14–15 week period) at a rate of 100 000 wasps per hectare. One‐way repeated measures analysis of variance was performed for the number of parasitized eggs, followed by a meta‐analysis procedure to determine the contribution of T. pretiosum release on overall parasitism. In addition, regression analysis was conducted with each season’s data sets to study the relationship of the host density and parasitism response by T. pretiosum. The overall results of Trichogramma augmentative releases did not result in significant increase of CLW egg parasitism beyond the natural parasitism in the areas studied. However, based on Cohen’s d effect sizes from the meta‐analysis, the parasitism rate was greater in fields under T. pretiosum releases during four out of 15 weeks surveyed. The parasitism of CLW eggs by T. pretiosum exhibited host density‐dependence only in one out of three seasons surveyed. These findings are encouraging as they are evidence that T. pretiosum is able to maintain a considerable level of parasitism under commercial field conditions, highlighting their potential value in large‐scale commercial areas of cotton as previously found at the small and diverse farming scale. Future studies should address the potential of early‐season, low density releases of the parasitoid.     

Parasitism capacity of Trichogramma pretiosum Riley (Hymenoptera: Trichogrammatidae) reared under different temperatures on Bonagota salubricola (Meyrick) (Lepidoptera: Tortricidae) eggs

The parasitism capacity of Trichogramma pretiosum Riley strain bonagota on Bonagota salubricola (Meyrick) eggs was studied under the temperatures of 18, 20, 22, 25, 28, 30 and 32 degrees C. The number of days with parasitism, accumulated parasitism, total number of eggs parasitized per female and parasitoid longevity was evaluated. In the first 24h, parasitism ranged from 1.6 (32 degrees C) to 8.8 (22 degrees C) eggs of B. salubricola. Accumulated egg parasitism of B. salubricola reached 80% in 1st to 4th day at 20 degrees C to 32 degrees C, respectively, and in the 7th day at 18 degrees C. Temperatures from 18 degrees C to 22 degrees C were the best suited for the total eggs parasitized for female, resulting in 35.4 and 24.6 eggs/male respectively. T. pretiosum female longevity ranged from 7.8 to 2.5 days, at 18 degrees C and 32 degrees C, respectively. The results showed that T. pretiosum strain bonagota is better adapted to temperatures from 18 degrees C to 22 degrees C.     

Methods for Detection of Anticarsia gemmatalis Nucleopolyhedrovirus DNA in Soil

Two methods, phenol-ether and magnetic capture-hybridization (MCH), were developed and compared with regard to their sensitivities and abilities to extract the DNA of the insect baculovirus Anticarsia gemmatalis nucleopolyhedrovirus (AgMNPV) from soil and to produce DNA amplifiable by PCR. Laboratory experiments were performed with 0.25 g of autoclaved soil inoculated with different viral concentrations to optimize both methods of baculovirus DNA extraction and to determine their sensitivities. Both procedures produced amplifiable DNA; however, the MCH method was 100-fold more sensitive than the phenol-ether procedure. The removal of PCR inhibitors from the soil appeared to be complete when MCH was used as the viral DNA isolation method, because undiluted aliquots of the DNA preparations could be amplified by PCR. The phenol-ether procedure probably did not completely remove PCR inhibitors from the soil, since PCR products were observed only when the AgMNPV DNA preparations were diluted 10- or 100-fold. AgMNPV DNA was detected in field-collected soil samples from 15 to 180 days after virus application when the MCH procedure to isolate DNA was coupled with PCR amplification of the polyhedrin region.     

Trichogramma pretiosum parasitism and dispersal capacity: a basis for developing biological control programs for soybean caterpillars

In order to succeed in biological control programs, not only is it crucial to understand the number of natural enemies to be released but also on how many sites per area this releasing must be performed. These variables might differ deeply among egg parasitoid species and crops worked. Therefore, these trials were carried out to evaluate the parasitism (%) in eggs of Anticarsia gemmatalis and Pseudoplusia includens after the release of different densities of the egg parasitoid Trichogramma pretiosum. Field dispersal was also studied, in order to determine appropriate recommendations for the release of this parasitoid in soybean fields. The regression analysis between parasitism (%) and densities of the parasitoid indicated a quadratic effect for both A. gemmatalis and P. includens. The maximum parasitism within 24 h after the release was reached with densities of 25.6 and 51.2 parasitoids per host egg, respectively, for the two pests. Parasitism of T. pretiosum in eggs of P. includens decreased linearly as the distance of the pest eggs from the parasitoid release sites increased. For P. includens, the mean radius of T. pretiosum action and the area of parasitoid dispersal in the soybean crop were 8.01 m and 85.18 m2, respectively. We conclude that for a successful biological control program of lepidopteran pests using T. pretiosum in soybean fields, a density of 25.6 parasitoids per host egg, divided into 117 sites per hectare, should be used.     

Separation and behavior in vitro of hemocytes from the moth,Pseudoplusia includens

Hemocytes collected from larvae of Pseudoplusia includens (Lepidoptera: Noctuidae) were separated by centrifugation on Percoll cushions. The procedure resulted in 95% purity of plasmatocytes and greater than 99% purity of granular and spherule cells. Medium supplemented with chicken serum enhanced cell viability and promoted spreading of plasmatocytes. Cell-free plasma and medium preconditioned by plasmatocytes or granular cells stabilized cells in vitro and also accelerated spreading of plasmatocytes relative to medium supplemented with chicken serum. Oenocytoids were the only morphotype that exhibited endogenous phenoloxidase activity, while granular cells and plasmatocytes were the only cells that endocytosed fluorescent beads in vitro. Granular cells and plasmatocytes ingested fluorescently labelled beads, both in mixed populations of hemocytes and after separation. Plasmatocytes were the only morphotype that encapsulated large foreign targets in vitro following separation. Separated granular cells attached and spread on the surface of foreign targets but never formed a multilayered capsule.     

不同温度下稻螟赤眼蜂在黏虫卵上的种群生命表

稻螟赤眼蜂Trichogramma japonicum Ashmead是水稻二化螟Chilo suppressalis(Walker)的优势卵寄生蜂。本文构建了以黏虫卵为中间寄主,稻螟赤眼蜂在20℃、25℃、30℃、33℃四个温度条件下的生命表。研究结果表明利用黏虫卵繁育稻螟赤眼蜂时,不同温度下其生长发育和繁殖均存在显著差异,雌蜂平均寿命随温度的升高呈下降趋势,在20℃条件下,雌蜂平均寿命为1.44 d,显著长于其它温度,当温度为30~33℃时,雌蜂寿命最短,平均为0.59 d。在温度为20℃和25℃时,稻螟赤眼蜂的羽化率显著高于温度较高的30℃和33℃,平均为94.73%。在25℃条件下,黏虫卵繁育的稻螟赤眼蜂的净生殖力(R 0)最大,达到31.9176,单雌寄生卵数最多,为32.88个,表现出最强的繁殖能力,而在30℃条件下,瞬时增长率r m和周限增长率λ达到最大值。综合生命表参数、单雌寄生黏虫卵数、寿命、羽化率等指标,利用黏虫卵工厂化繁殖稻螟赤眼蜂的最佳温度为25℃。本研究为利用黏虫卵大量繁殖稻螟赤眼蜂提供了有价值的参考。     

Effect of induced tolerance to Bt toxin on the egg size of Helicoverpa armigera and parasitism by Trichogramma pretiosum

Larvae of Helicoverpa armigera (Hübner) can develop a form of Bt tolerance after exposure to sub‐lethal doses of Bt‐toxin subclass Cry1Ac. Increasing levels of tolerance are produced over generations of larval exposure, which are not related to DNA sequence changes, and are largely maternally transmitted. The characteristic of maternal transmission, combined with the importance of egg parasitoids to cotton pest management, raises questions about the effects of Bt tolerance/exposure on the eggs of H. armigera and on some key metrics of egg parasitism. In the present study, the effect of inducible tolerance on eggs of H. armigera and parasitism by Trichogramma pretiosum (Riley) is investigated. First, the volumes of eggs laid by susceptible and tolerant H. armigera females are compared. In addition, the effect of inducible tolerance on egg parasitism is determined by comparing parasitism success, the number of adult wasps emerged per host egg, and the proportion of male and female offspring emerged per host egg. The results obtained suggest that Cry1Ac‐tolerance is associated with increased egg volume, even after just one generation of sub‐lethal exposure. When tolerant H. armigera are freed from ongoing sub‐lethal exposure, a corresponding decrease in egg volume is not detected. Although there is no difference in the percentage of eggs parasitized, there is an increase in the number of emergent parasitoids, especially males, from eggs laid by tolerant H. armigera. These results confirm that maternally‐transmitted Bt tolerance is reflected in the phenotype of the eggs of tolerant offspring, which affects egg parasitism.