Venom from the ectoparasitoid wasp Eulophus pennicornis disrupts host ecdysteroid production by regulating host prothoracic gland activity

Abstract. Attack by the ectoparasitoid Eulophus pennicornis Nees (Hymenoptera: Eulophidae) prevents larvae of Lacanobia oleracea L. (Lepidoptera: Noctuidae) from moulting. Prothoracic glands (PGs) excised from parasitized or artificially envenomated hosts show a reduced basal level of ecdysteroid release at a time when non-parasitized caterpillars produce an ecdysteroid surge (48 h post moult to 5th stadium = penultimate stadium in non-venomated hosts). By contrast, PGs from similarly parasitized or envenomated caterpillars release comparatively high levels of ecdysteroid at 120 h post-moult. Temporary inactivation of PGs cannot be attributed solely to a parasitoid-induced reduction in cell viability, and incubation in E. pennicornis venom in vitro does not exert any direct effect on either PG cell viability or ecdysteroid release. However, inactivated PGs are not stimulated by forskolin, which may indicate that the absence of the required pre-moult ecdysteroid surge in developmentally arrested L. oleracea is due to insensitivity to a prothoracicotropic hormone. Even though parasitized caterpillars never moult, reversed-phase HPLC separations and radioimmunoassay confirm that they produce active moulting hormone (20-hydroxyecdysone) at 120 h post-moult. These results suggest that E. pennicornis arrests host development through the indirect effects on their hosts' PGs. This effect is not achieved through the destruction of gland cells, but more likely reflects the interruption of an innate cycle in PG activity, such that they lose their ability to respond to a normal cue to produce an essential hormone peak at a crucial point in development.     

Parasitism of Lacanobia oleracea (Lepidoptera) by the ectoparasitoid, Eulophus pennicornis, is associated with a reduction in host haemolymph phenoloxidase activity

When haemolymph from fifth instar Lacanobia oleracea was incubated in vitro, rapid melanization occurred. Similar levels of melanization occurred in haemolymph from larvae that had been experimentally injected with venom from the ectoparasitic wasp, Eulophus pennicornis. In contrast, haemolymph from larvae parasitized by this wasp melanized more slowly and less extensively. Phenoloxidase assays indicated that enzyme activity was present in haemocyte lysate supernatants, serum and plasma from L. oleracea and that on day 5 post-parasitization, fractions prepared from parasitized larvae had significantly less phenoloxidase activity than similar fractions from untreated or experimentally envenomated larvae. In addition, no PO activity was detectable in wasp venom, and the venom had no effect on L. oleracea plasma phenoloxidase activity in vitro. These results indicate that parasitism of L. oleracea by E. pennicornis suppresses host haemolymph phenoloxidase activity and that this suppression is not induced by adult wasp venom. The results are discussed with reference to the survival advantages of suppressing the activity of this host enzyme, and to the possible source(s) of putative suppressive factors.     

Ability to host regulate determines host choice and reproductive success in the gregarious ectoparasitoid Eulophus pennicornis (Hymenoptera: Eulophidae)

Abstract.  The age of Lacanobia oleracea (L.) in the final (sixth) larval stadium influences host choice and developmental success significantly in the gregarious ectoparasitoid Eulophus pennicornis (Nees). In choice tests, parasitoids with prior oviposition experience parasitize hosts in the second day of the sixth stadium most frequently. Parasitoid brood survival on normally-reared (i.e. fed) hosts declines monotonically with age such that mean progeny survival (egg–adult) is less than 20% for wasps developing on hosts parasitized on day 5 of the sixth stadium, as opposed to almost 50% when developing on those parasitized on day 1. Neck ligation of hosts increases the survival of wasp larvae developing on older hosts (days 4 and 5), whereas starved hosts produce progeny in similar numbers to fed hosts on most days during the final larval stadium. Hosts parasitized early in the stadium (days 1–3), although continuing to grow, do not exhibit the characteristic physical changes that non-parasitized larvae exhibit prior to pupation. However, hosts parasitized on days 4 and 5 form prepupae in appreciable numbers, particularly on day 5 where, regardless of treatment, over 80% of hosts attain this stage. Envenomated hosts behave similarly, an observation that suggests that it is the wasp's inability to arrest completely development in older hosts that is the significant factor in reducing the developmental success of the wasp. The findings are discussed in the light of the known endocrinological events in the host, and in relation to previously reported host manipulations induced by this wasp.     

Inhibition of juvenile hormone esterase activity in Lymantria dispar (Lepidoptera, Lymantriidae) larvae parasitized by Glyptapanteles liparidis (Hymenoptera, Braconidae)

Glyptapanteles liparidis is a gregarious, polydnavirus (PDV)-carrying braconid wasp that parasitizes larval stages of Lymantria dispar. In previous studies we showed that parasitized hosts dramatically increase juvenile hormone (JH) titers, whereas JH degradation is significantly inhibited in the hemolymph. Here we (i) quantified the effects of parasitism on JH esterase (JHE) activity in hemolymph and fat body of penultimate and final instars of L. dispar hosts and (ii) assessed the relative contribution of individual and combined wasp factors (PDV/venom, teratocytes, and wasp larvae) to the inhibition of host JHE activity. The effects of PDV/venom was investigated through the use of gamma-irradiated wasps, which lay non-viable eggs (leading to pseudoparasitization), while the effects of teratocytes and wasp larvae were examined by injection or insertion of these two components in either control or pseudoparasitized L. dispar larvae. Parasitism strongly suppressed host JHE activity in both hemolymph and fat body irrespective of whether the host was parasitized early (premolt-third instar) or late (mid-fourth instar). Down-regulation of JHE activity is primarily due to the injection of PDV/venom at the time of oviposition, with only very small additive effects of teratocytes and wasp larvae under certain experimental conditions. We compare the results with those reported earlier for L. dispar larvae parasitized by G. liparidis and discuss the possible role of JH alterations in host development disruption.     

Juvenile hormone synthesis, metabolism, and resulting haemolymph titre in Heliothis virescens larvae parasitized by Toxoneuron nigriceps

Last instar larvae of the tobacco budworm, Heliothis virescens F., fail to pupate and have little 20-hydroxyecdysone when parasitized by Toxoneuron nigriceps (Viereck). In this paper, we extend these observations to juvenile hormone (JH) to determine if parasitism by this wasp affects other endocrine systems. To this end, we compared the production of JH by corpora cardiaca-corpora allata complexes (CC-CA), the metabolism of JH by haemolymph enzymes, and the haemolymph titre of JH in parasitized and non-parasitized control larvae of H. virescens during the last larval instar. CC-CA from parasitized and control larvae had similar peaks of JH synthesis on day 1 of the fifth instar, with JH II accounting for more than 90% of total JH in both groups. On subsequent days, JH synthesis dropped to undetectable levels more quickly in non-parasitized controls than in parasitized larvae. JH metabolism by haemolymph of parasitized and control animals increased from low levels on day 1 of the fifth instar to high levels on days 2 and 3 of the instar. JH metabolism was significantly higher in control larvae than in parasitized larvae. After day 3, JH metabolism decreased in both groups, but was significantly higher in parasitized larvae. The major metabolite of JH in both groups was JH acid, though traces of JH diol and JH acid diol were also detected. The haemolymph titre of JH in both groups peaked on day 1 of the fifth instar and, similar to the synthesis of JH by CC-CA, decreased more rapidly in control larvae. As a result, non-parasitized animals had significantly lower JH titres on day 2. The higher JH titres observed in parasitized larvae during the early fifth instar may contribute to their developmental arrest. The possible role of these JH alterations in the host developmental and metabolic redirection is discussed and a more comprehensive physiological model accounting for host-parasitoid interactions is proposed.     

Developmental interaction between suboptimal instars of Spodoptera littoralis (Lepidoptera:Noctuidae) and its parasitoid Microplitis rufiventris (Hymenoptera:Braconidae)

Relative effects of parasitism by Microplitis rufiventris on the development of the third instar Spodoptera littoralis (preferable, optimal host) with the development of penultimate (5th) and last (6th) instars (suboptimal hosts) were investigated. Newly molted 6th instar hosts were more acceptable for parasitization by the wasp female than older hosts. In singly parasitized 3rd instar hosts, 82.0 +/- 3.9% of the parasitoid eggs developed to full-grown instar wasp larvae. However, parasitoid eggs deposited singly in 73.9 +/- 3.3% of 5th and 100% of 6th instar hosts failed to develop. Superparasitization in the 3rd instar hosts reduced the production of pseudoparasitized larvae and, conversely, all parasitized hosts yielded viable parasitoid offspring. In suboptimal hosts, the development interaction between the parasitoid and its host larvae was highly influenced by the age of hosts at parasitism, load of deposited eggs, and other parasitoid factors. The latter factors, e.g., mainly calyx fluid particles, might be involved in establishing parasitoid eggs in the suboptimal hosts. In the last two host instars, superparasitization significantly increased the number of parasitoid larvae successfully reaching their final instar. Variation in host quality, e.g., physiological status, might be attributed, in part, to the partial breakdown of the solitary habit observed in the earlier instars. More parasitoid eggs developed to mature parasitoid larvae in hosts superparasitized as 6th instar than parasitoid eggs laid in 5th instar hosts. Superparasitization significantly lengthened the developmental period of 5th and 6th host instars and inhibited their development to the pupal stage. Studying parasitoid development in suboptimal instars of its habitual host provided physiological insight, as shown here. The results may have implication for biological control and in vitro mass rearing programs with solitary parasitoids.     

Does superparasitism improve host suitability for parasitoid development? A case study in the Microplitis rufiventris-Spodoptera littoralis system

Superparasitism refers to the oviposition behavior of parasitoid females who lay their eggs in an already parasitized host. Recent studies have shown that allocation of additional eggs to an already parasitized host may be beneficial under certain conditions. In the present work, mortality of Microplitis rufiventris wasps was significantly influenced by both host instar of Spodoptera littoralis larvae at parasitism and level of parasitism. In single parasitization, all host instars (first through sixth) were not equally suitable. Percentage of emergence success of wasp larvae was very high in parasitized first through third (highly suitable hosts), fell to 60% in the fourth instar (moderate suitable) and sharply decreased in the penultimate (5th) instars (marginally suitable). Singly parasitized sixth (last) instar hosts produced no wasp larvae (entirely unsuitable), pupated and eclosed to apparently normal adult moths. The scenario was different under superparasitism, whereas supernumerary individuals in the highly suitable hosts were almost always killed as first instars, superparasitization in unsuitable hosts (4th through 6th) had significant increase in number of emergence success of wasp larvae. Also, significantly greater number of parasitoid larvae successfully developed in unsuitable hosts containing three wasp eggs than counterparts containing two wasp eggs. Moreover, the development of surplus wasp larvae was siblicidal in earlier instars and nonsiblicidal gregarious one in the penultimate and last “sixth” instars. It is suggested that the optimal way for M. rufiventris to deal with high quality hosts (early instars) is to lay a single egg, while the optimal way to deal with low quality hosts (late instars) might be to superparasitize these hosts.     

The effect of snowdrop lectin (GNA) delivered via artificial diet and transgenic plants on Eulophus pennicornis (Hymenoptera: Eulophidae), a parasitoid of the tomato moth Lacanobia oleracea (Lepidoptera: Noctuidae)

Snowdrop lectin (Galanthus nivalis agglutinin, GNA) has previously been shown to confer significant levels of protection against the lepidopteran pest Lacanobia oleracea when expressed in transgenic potato. The effect of GNA on the parasitism of L. oleracea by the gregarious ectoparasitoid Eulophus pennicornis was investigated. Maize-based, and potato leaf-based diets containing GNA, and excised transgenic potato leaves expressing GNA, were fed to L. oleracea larvae from the beginning of either the third or fourth larval instar. Lacanobia oleracea larvae were individually exposed to single mated adult female E. pennicornis parasitoids from the fifth instar onwards.The success of the wasp was not reduced by the presence of GNA in any of the diets, or by the length of feeding of the host prior to parasitism. However, the mean number of wasps that developed on L. oleracea reared from the third instar on the GNA-containing maize diet was significantly higher than on the controls (20.6 and 9.3 adults/host respectively). In all other cases differences were not significant. Eulophus pennicornis progeny that developed on L. oleracea reared on GNA-containing diets showed little or no alteration in size, longevity, egg load and fecundity when compared with wasps that had developed on hosts fed the respective control diets.The results suggest that expression of GNA in transgenic crops to confer resistance to lepidopteran pests will not adversely affect the ability of the ectoparasitoid E. pennicornis to utilise the pest species as a host.     

Infection by the microsporidium Vairimorpha necatrix (Microspora: Microsporidia) elevates juvenile hormone titres in larvae of the tomato moth, Lacanobia oleracea (Lepidoptera: Noctuidae)

The effects of infection by a microsporidium, Vairimorpha necatrix (Kramer), on the endogenous levels of juvenile hormones in tomato moth (Lacanobia oleracea L.) larvae were investigated. Levels of juvenile hormone II (JH II) were 10-fold greater in the infected larvae on day two of the sixth stadium but no significant difference was observed on day seven. Juvenile hormone I (JH I) was also detected in day two and day seven sixth instar infected larvae but was not detected in non-infected larvae. The duration of the fifth and sixth stadia was significantly longer for infected larvae when compared with non-infected larvae. No evidence was found to suggest that supernumerary moults are a feature of infection by V. necatrix in L. oleracea larvae. Experiments were performed to determine whether the elevation in JH levels, which probably prevents pupation, is an adaptive mechanism of the microsporidium for extending the growth phase of the host, thereby allowing increased spore production. A proportion of infected larvae were collected on days 9 and 24 of the sixth stadium and spore extracts prepared from each larva. These days represent the average duration of the sixth stadium required for uninfected larvae to reach pupation, and the average number of days that V. necatrix-infected larvae survive in the sixth stadium before dying from infection. The mean spore yields from infected larvae 24 days into the sixth stadium were significantly higher than the spore yields obtained from day nine sixth instar larvae. The hypothesis that V. necatrix manipulates host endocrinology (i.e. prolong the host larval state to maximise spore yield) is discussed in context with the results obtained.     

Stage-specific effects of Campoletis sonorensis parasitism on Heliothis virescens development and prothoracic glands

Abstract The ichneumonid endoparasitoid Campoletis sonorensis Cameron (Hymenoptera: Ichneumonidae) injects a polydnavirus when it oviposits into a host. We compared the development of Heliothis virescens (F.) (Lepidoptera: Noctuidae) larvae parasitized in the penultimate (fourth) stadium with those parasitized in the last (fifth) stadium by C.sonorensis and show that hosts stung in the fifth stadium exhibited arrested or delayed development compared to the controls. Parasitoids developed normally to the point of emergence in larvae stung in the fifth stadium but most did not successfully emerge from the host. The prothoracic glands in all successfully parasitized fifth stadium hosts and most unsuccessfully parasitized fifth stadium hosts showed some degree of virally-induced degeneration. Larvae stung in the fourth stadium developed more slowly than controls and either did not moult or developed to a fifth and sometimes a supernumerary sixth stadium before parasitoid emergence. Unsuccessfully parasitized hosts were delayed in their development but eventually moulted to the fifth and, in some cases, a supernumerary sixth stadium before pupating. Hosts stung in the fourth stadium showed no signs of prothoracic gland degeneration whether successfully parasitized or not. In addition, calyx fluid injections into early fourth stadium hosts did not cause prothoracic gland degeneration even after these hosts moulted to the fifth stadium, suggesting that degeneration induced by polydnavirus is specific to the last stadium of the host.     

The juvenile hormone titer of Trichoplusia ni and its potential role in embryogenesis of the polyembryonic wasp CopidosomaFloridanum

The hemolymph juvenile hormone (JH) titer of third through fifth stadia Trichoplusia ni parasitized by the polyembryonic parasitoid, Copidosoma floridanum, was measured by radioimmunoassay and compared to the titers of unparasitized larvae. The JH titer of parasitized larvae fluctuated from 28 pg/μl to undetectable levels. Maximum levels of hormone were present at ecdysis to the fourth and fifth stadium, and at the prepupal stage. Qualitatively, similar fluctuations were observed in unparasitized larvae. However, the titers in unparasitized larvae were much lower than those of parasitized larvae in the third and early fourth stadia, and the titer fell to undetectable levels in the fifth stadium 24 h earlier (48 h) than in parasitized larvae (72 h). Preventing the JH titer from falling during the fourth and fifth stadia by topical application of (RS)-methoprene or JH II had a juvenilizing effect on parasitized T. ni, and inhibited C. floridanum embryo morphogenesis. The effect of exogenous methoprene and JH on C. floridanum development depended on timing of application and dosage. Application of 100 pmol per day of methoprene beginning at 2 h of the host fourth stadium, prior to the large drop in the endogenous JH titer, inhibited morphogenesis in the majority of C. floridanum embryos. Application of methoprene at later times of host development did not inhibit morphogenesis although other developmental alterations were observed. The potential significance of host JH and ecdysteroid titers on polyembryonic development are discussed.     

Juvenile hormone inhibits the synthesis of major larval haemolymph proteins in rice moth, Corcyra cephalonica

Larval haemolymph proteins (LHP), LHP49 and LHP46 are produced in the penultimate and last larval instars. Starvation during the early and mid stage of last instar development prevents the production of both LHPs. Decapitation in early and mid last instar stimulates LHP synthesis and their concentration in haemolymph increases, while ligation in last instar larvae blocks the production of LHPs. Application of exogenous JH lowers the synthesis of LHP49 and LHP46 in Corcyra. These observations suggests that LHP49 and LHP46 synthesis is activated during the periods when JH titres are either low or undetectable.     

中红侧沟茧蜂寄生对寄主粘虫血淋巴糖类、脂类和蛋白质含量的影响

为揭示寄生蜂寄生对其寄主的生理调控机制, 室内对中红侧沟茧蜂Microplitis mediator寄生与未被寄生寄主粘虫Mythimna separata幼虫血淋巴中糖类、脂类和蛋白含量变化进行了测定。结果显示: 在滞育与非滞育条件下, 被寄生的粘虫血淋巴中糖原浓度均比未被寄生的粘虫高。滞育条件下寄生后12 d差异显著（P<0.05）, 被寄生粘虫糖原含量为7.93 μg/mL, 未被寄生粘虫糖原含量为4.70 μg/mL; 非滞育条件下寄生后6 d差异显著（P<0.05）, 被寄生粘虫糖原含量为14.35 μg/mL, 未被寄生粘虫糖原含量为5.47 μg/mL。海藻糖含量测定结果显示, 在滞育条件下寄生蜂对被寄生粘虫无明显影响, 而非滞育条件下影响效果差异显著（P<0.05）, 寄生后4 d被寄生粘虫海藻糖含量为46.82 μg/mL, 未被寄生粘虫含量为26.72 μg/mL。在滞育与非滞育两种条件下, 寄生与未被寄生寄主脂类和蛋白含量没有显著性差异。结果说明: 寄生蜂的存在使寄主血淋巴中的糖原含量增高; 非滞育条件是影响被寄生粘虫海藻糖含量变化主要因素; 粘虫对中红侧沟茧蜂的寄生表现相当强的适应性和忍受力。     

Granulovirus prevents pupation and retards development of Adoxophyes honmai larvae

Abstract Larvae of Adoxophyes honmai (Lepidoptera: Tortricidae) infected with granulovirus (AdhoGV) do not pupate; instead, they undergo prolonged larval development and die during the final stadium. Non-infected larvae, however, pupate after five larval stadia. Insect metamorphosis is regulated by fluctuations of ecdysteroid and Juvenile Hormone (JH). JH esterase activity and titres of ecdysteroid must be measured to understand fully the interaction of an insect virus and its host. JH esterase activity is consistently low in AdhoGV-infected larvae, which suggests that JH in AdhoGV-infected larvae is not degraded during the final stadium. The ecdysteroid titre in non-infected larvae showed a large peak in the final stadium before pupation, whereas that in AdhoGV-infected larvae increased from day 2 to day 5 in the final stadium, and then remained at a high level until death. Furthermore, an ecdysteroid UDP-glucosyltransferase (EGT) assay showed that this activity occurs in haemolymph from AdhoGV-infected larvae, but not in haemolymph of non-infected larvae. PCR and sequencing analysis revealed that the AdhoGV genome contains an egt gene, which encodes a protein of 445 amino acids, located approximately 1 kbp upstream from the granulin gene. These results suggest that AdhoGV-infected larvae are prevented from pupating because JHE activity is suppressed and EGT expression inactivates ecdysteroid in the haemolymph.     

Biology of Microplitis kewleyi (Hymenoptera: Braconidae): A parasite of Agrotis ipsilon (Lepidoptera: Noctuidae) larvae

Microplitis kewleyi Muesebeck is a gregarious internal parasite of larvae of the black cutworm Agrotis ipsilon (Hufnagel). Studies of the biology of the parasite revealed that there was an inverse relationship between host instar and parasite preference. Duration of development from egg to pupa ranged from 18 days at 27°C to 68.7 days at 16°C. Development from egg to pupa took 13.5–21.6 days when fourth and first instar host larvae, respectively, were parasitized. A larger number of parasites emerged from hosts parasitized in the fourth instar (22.4) than the first instar (11.5). Parasite pupation occurred when the host was in the fifth/sixth instar, depending on the instar parasitized. Thirty‐nine per cent of host larvae exposed as first instars to parasites died before parasite emergence. This decreased to 0% for host larvae exposed as fourth instars. The sex ratio was 1:1.2 (M:F). Thirty‐seven per cent of hosts exposed diurnally were stung, compared to 24% exposed nocturnally. Mean daily progeny was highest (12) on the first day, decreasing to zero after 20 days. Percent host parasitism was also highest on the first day (35%) decreasing to nearly 0% after 18 days. There appear to be three parasite larval instars. Host larvae often remained alive after parasite emergence.     

Performance of Microplitis tuberculifer (Hymenoptera: Braconidae) parasitizing Mythimna separata (Lepidoptera: Noctuidae) in different larval instars

The solitary parasitoid Microplitis tuberculifer (Wesmael) is an important biological control agent of various lepidopteran pests in Asia. We examined the preference of M. tuberculifer for different instars of its common host, Mythimna separata (Walker), host instar effects on parasitoid development, and the consequences of parasitism in different stages for growth and consumption of host larvae. The wasp successfully parasitized the first four larval instars of M. separata, but not the fifth, which appeared to be behaviorally resistant. First and second instars were parasitized at higher rates compared to thirds and fourths in no-choice situations, ostensibly due to longer handling times for the latter, but second instars were most preferred in a choice test that presented all stages simultaneously. Although later instar hosts yielded heavier cocoons, the fastest parasitoid development was obtained in second instars. Lower sex ratios were obtained from first instars as females appeared to lay a smaller proportion of fertilized eggs in small hosts. Both weight gain and food consumption of parasitized larvae were reduced significantly within 24 h of parasitism, regardless of the stage parasitized, and final body weights were less than 10% those of unparasitized larvae. Thus, M. tuberculifer has good potential as a biological control agent of M. separata, successfully parasitizing the first four larval instars and dramatically reducing plant consumption by the host in all cases.     

The biology of Meteorus gyrator (Hymenoptera: Braconidae), a solitary endoparasitoid of the tomato moth, Lacanobia oleracea (Lepidoptera: Noctuidae)

There is a need to identify potential biological control agents for use against lepidopterous pests in greenhouses. The solitary endoparasitoid Meteorus gyrator (Thunberg) attacks a range of macrolepidopterous larvae, including those of some important horticultural pest species. Laboratory trials designed to investigate the biology of M. gyrator on larvae of the tomato moth, Lacanobia oleracea Linnaeus, reveal that this parasitoid is capable of parasitizing all larval stages of its host, third instars being parasitized most frequently. Each female parasitoid lives for up to 40 days (at 25 degrees C), ovipositing into an average of 78 hosts. Preadult development is rapid ( approximately 2 weeks), and the sex ratio of offspring is 1:1. Parasitism by M. gyrator suppresses the growth of both early and late host instars, and there is a concomitant reduction in the amount of food consumed (overall feeding reduction over a 12 day period is 68%). Our results indicate that inoculative releases of M. gyrator could provide effective biological control of L. oleracea and other noctuid pests of greenhouses.     

Effect of entomopoxvirus infection of the smaller tea tortrix, Adoxophyes sp. on the development of the endoparasitoid, Ascogaster reticulatus

Infection of Adoxophyes sp. (Lepidoptera: Tortricidae) larvae by an entomopoxvirus (AsEPV) adversely affected the development of the endoparasitoid, Ascogaster reticulatus Watanabe (Hymenoptera: Braconidae). Parasitoid larvae developing in AsEPV-infected hosts grew more slowly and spent more time in their hosts than did parasitoid larvae developing in noninfected hosts. Percentages of emergence of larval parasitoids that developed in AsEPV-infected hosts were significantly lower than those of parasitoids that developed in noninfected hosts. Parasitoid larvae in AsEPV-infected host perished when their hosts died of AsEPV infection. Significant numbers of parasitized and infected larvae exhibited apolysis to the final instar, whereas noninfected-parasitized larvae died in the penultimate instars due to emergence of parasitoids.     

Changes in haemolymph proteins of the fall armyworm, Spodoptera frugiperda (J. E. Smith), associated with parasitism by the braconid parasitoid Cotesia marginiventris (Cresson)

Changes in haemolymph proteins of the fall armyworm, Spodoptera frugiperda, associated with parasitism by the parasitoid Cotesia (= Apanteles) marginiventris were monitored by sodium dodecyl sulphate polyacrylamide gel electrophoresis. As early as hour 4 after parasitization treatment, several electrophoretically slow-migrating, high-molecular-weight proteins were detected in the host's haemolymph. These proteins were detected earlier in haemolymph from parasitized larvae than in haemolymph from control larvae, and their concentrations were higher in heavily parasitized host larvae (≥ 3 eggs/host) than in lightly parasitized larvae (1 egg/host). Additionally, unique proteins that migrated electrophoretically with bovine serum albumin appeared in the haemolymph of parasitized larvae at hour 8 after parasitization treatment and were evident in haemolymph collected through to hour 64.     

Dose-dependent influence of Microplitis rufiventris factors on developmental rate and growth of last-instar Spodoptera littoralis larvae

The solitary endoparasitoid, Microplitis rufiventris, attacks and can develop in earlier instars of Spodoptera littoralis larvae with preference to third‐instar larvae. We used the last stadium (sixth instar), a stage which is not naturally parasitized. The newly moulted larvae (0–3 h old) of this stadium were more acceptable for parasitization by the wasp females than the older ones (24 h old). Parasitization by M. rufiventris wasp of last instar S. littoralis larvae leads to dose (no. of eggs + parasitoid factors)‐dependent effects which were more pronounced at 20°C than at 27°C. A single oviposition into a sixth instar host larva resulted in normal development of the host. However, superparasitization increased the proportions of developmentally arrested hosts and number of live wasp larvae. Development of supernumerary individuals of the parasitoid in the host larva leads to dose‐related adverse effects on host growth and development. The present study may provide interesting opportunities for studying the physiological bases of host–parasitoid interactions and parasitoid intra‐specific competition in the biological system considered.