Leptopilina heterotoma and L. boulardi: strategies to avoid cellular defense responses of Drosophila melanogaster

Eggs of three strains of the cynipid parasitoid Leptopilina heterotoma and a Tunisian strain (G317) of L. boulardi are not encapsulated by hemocytes of Drosophila melanogaster hosts, but the eggs of a Congolese strain (L104) of L. boulardi are encapsulated. To determine the reason for the difference in host response against the parasitoid eggs, lamellocytes (hemocytes that encapsulate foreign objects and form capsules around endogenous tissues in melanotic tumor mutants) were examined in host larvae parasitized by the five Leptopilina strains. Parasitization by the three L. heterotoma strains affected the morphology of host lamellocytes and suppressed endogenous melanotic capsule formation in melanotic tumor hosts. L104 did not alter the morphology of host lamellocytes nor block tumor formation in melanotic tumor mutant hosts. The morphology of some lamellocytes was affected by G317 parasitization but host lamellocytes were still capable of forming melanotic tumors and encapsulating dead supernumerary parasitoid larvae. Therefore, the eggs of strains affecting lamellocyte morphology are protected from encapsulation by the host's blood cells. L. heterotoma eggs float freely in the host hemocoel but L. boulardi eggs are attached to host tissue surfaces. Lamellocytes cannot infiltrate the attachment site so the capsule around the L104 egg remains incomplete. The wasp larva uses this gap in the capsule as an escape hatch for emergence.     

Effects of lamellolysin from a parasitoid wasp on Drosophila blood cells in vitro

Female parasitoid Leptopilina heterotoma inject a factor, lamellolysin, along with their eggs into the host hemocoel to destroy selectively host hemocytes that encapsulate foreign objects. In parasitized Drosophila melanogaster larvae, these hemocytes (lamellocytes) change from discoidal cells to bipolar cells that no longer adhere to each other to form capsules. To study the effects of lamellolysin on Drosophila lamellocytes in vitro, a giant strain of D. melanogaster was constructed to yield hemolymph with an abundance of lamellocytes. The effect of lamellolysin on the adhesivity of lamellocytes in vitro was demonstrated when the cells were gently rotated in the culture medium. Under these conditions, the bipolar shape of the affected lamellocytes resembled that of lamellocytes in parasitized hosts. When lamellocytes were exposed to lamellolysin in stationary culture medium, the elongation of the bipolar cells continued until they became threadlike. Lamellocytes fragmented in both stationary and rotating culture medium in the presence of lamellolysin, although loss of cellular material was more pronounced in the latter. This study demonstrates that lamellolysin acts directly and destructively on lamellocytes.     

Haemocyte changes in D. Melanogaster in response to long gland components of the parasitoid wasp Leptopilina boulardi: a Rho-GAP protein as an important factor

The hymenopteran wasp Leptopilina boulardi (Figitidae) is a larval solitary parasitoid of Drosophila larvae of the melanogaster sub-group. The factors used by parasitoid females to prevent encapsulation of their eggs by the host are localized in the female long gland and reservoir. We report here the physiological effects of these factors on host haemocytes using in vivo injection experiments. The total number of haemocytes, the number of plasmatocytes and the number of crystal cells were not modified by injection of long gland extracts. In contrast, long gland extracts either from virulent or avirulent strains had a significant effect on the lamellocyte number. Compared to the Ringer control, the avirulent long gland products induced an increase of the lamellocyte number while virulent extracts induced a drastic decrease together with an alteration of the morphology of these cells. Interestingly, changes in the lamellocyte morphology were also observed following injection of the P4 protein, a major component of L. boulardi female long glands that displays a strong immune suppressive effect on Drosophila larvae. The implication of the P4 protein in suppressing the host cellular immunity is discussed in correlation with its predicted molecular function as a Rho-GAP protein.     

A RhoGAP protein as a main immune suppressive factor in the Leptopilina boulardi (Hymenoptera, Figitidae)-Drosophila melanogaster interaction

To protect its eggs, the endoparasitoid wasp Leptopilina boulardi injects immune suppressive factors into Drosophila melanogaster host larvae. These factors are localized in the female long gland and reservoir. We analyzed the protein content of these tissues and found that it strongly differed between virulent and avirulent parasitoid strains. Four protein bands present in virulent long glands were eluted and their immune suppressive effect was assessed in vivo, allowing demonstrating a major effect of one of these proteins. The corresponding cDNA encodes a predicted 30 kDa subunit containing a Ras homologous GTPase Activating Protein (RhoGAP) domain, suggesting a possible involvement in the regulation of actin cytoskeleton changes. Using Western-blot experiments, we showed that this protein is abundant in virulent female long glands but is undetectable in virulent females deprived of long glands or in long glands from avirulent wasps. Its potential role in modifying the morphology and the adhesive properties of the host lamellocytes, involved in Drosophila cellular immune responses, is discussed.     

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.     

Protein tyrosine phosphatases of Toxoneuron nigriceps bracovirus as potential disrupters of host prothoracic gland function

The genomic sequence of the bracovirus associated with the wasp Toxoneuron nigriceps (Hymenoptera, Braconidae) (TnBV), an endophagous parasitoid of the tobacco budworm larvae, Heliothis virescens (Lepidoptera, Noctuidae), contains a large gene family coding for protein tyrosine phosphatases (PTPs). Here we report the characterization of cDNAs for two of the viral PTPs isolated by screening a cDNA library from haemocytes of parasitized host larvae. The two encoded proteins show 70% amino acid identity and are expressed in the fat body of parasitized hosts. In addition, one was expressed in inactivated prothoracic glands (PTGs), 24 h after parasitoid oviposition. The rapid block of ecdysteroidogenesis does not appear to be due to inhibition of general protein synthesis, as indirectly indicated by the unaltered S6 kinase activity in the cytosolic extracts of basal PTGs from parasitized host larvae. Rather, TnBV PTP over-expression in inactivated host PTGs suggests that gland function may be affected by the disruption of the phosphorylation balance of key proteins regulating points upstream from the ribosomal S6 phosphorylation in the PTTH signaling cascade.     

Effects of parasitization or injection of parasitoid-derived factors from the endoparasitic wasp Glyptapanteles porthetriae (Hym., Braconidae) on the development of the larval host, Lymantria dispar (Lep., Lymantriidae)

Second instar larvae of Lymantria dispar were parasitized or injected with parasitoid-derived factors such as venom, calyx fluid or parasitoid eggs from Glyptapanteles porthetriae . Growth and development of the host larvae were affected in all different groups compared to control larvae of the same age, injected with Ringer solution. The greatest impact on host growth and on the duration of the 3rd instar was caused by injecting parasitoid eggs. Treated larvae showed melanized capsules or nodules in the hemocoel. While the wasp age had no effect on parasitization efficiency or on the percentage of melanized particles in the hemocoel, significantly more encapsulations were found in larvae parasitized by old wasps as opposed to young wasps. Superparasitization (double or quadruple oviposition) increased the parasitization efficiency markedly. While none of the control larvae showed melanized particles, in the groups of single and superparasitized (2× and 4×) hosts a high percentage of melanized particles (capsules and nodules) occurred.     

拟寄生蜂搜索产卵过程中对寄主的竞争

综述拟寄生蜂搜索产卵过程中对寄主竞争的最新研究进展.这类竞争具有四种方式,即标记寄主、杀卵和杀幼、守护寄主和捕食寄主.（1）标记寄主常涉及寄主标记信息素,这是由雌蜂在产卵前、产卵时或产卵后分泌的化学物质.寄主标记信息素常介导拟寄生蜂对已寄生和健康寄主的辨别、减少过寄生和多寄生、减轻种内和种间竞争压力.（2）雌蜂遇到已寄生寄主时,很多种类杀死前一雌蜂遗留的卵和幼虫,再产下自己的卵.雌蜂使用三种方法杀卵和杀幼,即产卵器穿刺、取食和使用有毒物质.通过杀卵和杀幼,产卵雌蜂清除了前一雌蜂遗留的后代,主动改善了寄主品质,从而有利于自身后代的生存.（3）守护寄主在肿腿蜂科、缘腹细蜂科、金小蜂科、缨小蜂科和茧蜂科中均有报道,守护者驱逐入侵者以保护后代及健康寄主.（4）捕食寄主不仅减少了健康寄主数量,且直接导致已寄生寄主中拟寄生蜂卵和幼虫的死亡.雌蜂一般在体内成熟卵量较少时捕食寄主.讨论了研究拟寄生蜂搜索产卵过程中竞争寄主的理论意义和实际应用价值.     

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.     

Biogenesis, structure, and immune-suppressive effects of virus-like particles of a Drosophila parasitoid, Leptopilina victoriae

Drosophila melanogaster larvae are attacked by virulent strains of parasitoid wasps. Females of Leptopilina heterotoma produce virus-like particles (VLPs) that efficiently destroy lamellocytes, a major larval immune effector cell type. We report here that L. victoriae, a closely related wasp species, also produces VLPs that trigger immune suppression responses in fly hosts. We compare the ability of immune suppression of the two parasitoids using a mutant host strain hopscotch^{Tumorous-lethal} (hop^Tum-l). hop^Tum-l larvae have two defects of hematopoietic origin: overproliferation of hemocytes and constitutive encapsulation of self-tissue by lamellocytes. The encapsulation phenotype is suppressed weakly by L. victoriae and strongly by L. heterotoma. In vitro studies on hop^Tum-l lamellocytes show that VLP-containing fluid from either wasp species induces lamellocyte lysis, but with different kinetics.Previously undocumented precursors of L. victoriae VLPs are synthesized in the long gland and are first visible within canals connecting secretory cells to the long gland lumen. VLP assembly occurs in the lumen. VLPs show multiple electron-dense projections surrounding a central core. Maturing particles appear segmented, singly or in arrays, embedded in the reservoir matrix. In sections, mature particles are pentagonal or hexagonal; the polygon vertices extending into spikes. Our results suggest that L. victoriae is likely to promote immune suppression by an active mechanism that is mediated by VLPs, similar to that used by L. heterotoma.     

Parasitism-induced Effects on Host Growth and Metabolic Efficiency in Tobacco Hornworm Larvae Parasitized by Cotesia congregata

Parasitism by the braconid wasp Cotesia congregata affects the growth of Manduca sexta larvae in a parasitoid 'dose-dependent' fashion. Following parasitization of fourth-instar larvae, more heavily parasitized larvae grew larger compared to those containing fewer parasitoids due to an increase in host dry weight. The differences in host mass appeared to arise after oviposition. A 'dose-dependent' enhancement of host dry weight would appear nutritionally beneficial for the parasitoids developing in more 'crowded' hosts. The efficiencies of conversion of ingested and digested food to body mass and the approximate digestibility of the diet ingested by the host caterpillar did not vary significantly with clutch size although parasitoids took slightly longer to develop in the more heavily parasitized hosts. Larval parasitoids developing in the presence of many competitors weighed up to 50% less than those developing in hosts with fewer endoparasitoids, although the weight of adult female parasitoids did not vary significantly with wasp clutch size. The maximum number of emerging wasps was 200 parasitoids, possibly representing the host's 'carrying capacity' for larvae parasitized in the fourth-instar. The ratio of emerging to non-emerging parasitoids decreased as parasitoid clutch size increased, with few or none emerging from very heavily parasitized hosts containing more than 400 parasitoids. Copyright 1997 Elsevier Science Ltd. All right reserved     

Effects of the parasitization of a braconid, Apanteles, on the blood of its host, Pieris

Parasitization of a braconid wasp, Apanteles glomeratus, of larvae of a common cabbage butterfly, Pieris rapae crucivora, caused changes in differential haemocyte count (DHC), total haemocyte count (THC), and encapsulative capacity against dead eggs of Apanteles in the fourth and fifth instar host larvae.However, no correlation could be found between the number of Apanteles eggs deposited and THC of the middle fourth instar host larvae or between the number of parasitoid larvae and specific gravity of the haemolymph from the late fifth instar host larvae.From the changes in DHC and in THC of both non-parasitized and parasitized Pieris larvae, an increase in the number of plasmatocytes of non-parasitized Pieris larvae in the early fourth instar period was supposed to be due to transformation of prohaemocytes into plasmatocytes, and a low population of plasmatocytes of parasitized larvae in the comparable period was assumed to be due to a suppression of transformation of prohaemocytes by some factor released from the parasitoid eggs.Failure of the parasitized fourth instar Pieris larvae to encapsulate injected dead eggs of Apanteles indicated that the parasitoid embryos were, in some way, actively inhibiting the encapsulation reactions of the host.The increase in THC of the parasitized fifth instar larvae could not be ascribed to a decrease in the volume of host haemolymph. Rather it could be interpreted by a suppression of adhesive capacity of haemocytes in the host haemocoel to tissue surfaces.Reduced encapsulative capacity of the parasitized fifth instar larvae might be attributed either to a depression of the adhesive activity of plasmatocytes resulting from a depletion of energy source for haemocytes in the host haemolymph by parasitization, or from an active suppression of adhesiveness of the plasmatocytes by secretions from ‘giant cells’ (teratocytes) originated from the parasitoid.     

Egg dispersion is more important than competition type for herbivores attacked by a parasitoid

Herbivore fitness can be altered by a combination of interacting organisms, such as its food plant, conspecifics, and predators/parasitoids. Here, we tested relative effects of plant species, herbivore intraspecific competition type, and spatial distribution of the herbivore among plant units on herbivore survival and whether parasitoids modified these effects. We used an endophagous bruchine seed predator Callosobruchus maculatus for the herbivore, and a braconid wasp Heterospilus prosopidis for the parasitoid. The survival rate of C. maculatus was measured for each of 16 combinations of two plants (bean species, Vigna unguiculata and V. radiata), two competition types of C. maculatus larvae (contest and scramble), two spatial distributions of hosts [sparse (1 C. maculatus larva per seed over 20 seeds) and dense (2 C. maculatus larvae per seed over ten seeds)], and with/without a parasitoid pair. In the absence of the parasitoid, C. maculatus survival rate was lower with V. radiata and in the contest type. With the parasitoid, the proportion parasitized hosts was independent of total host density. Neither the proportion of parasitized hosts nor host survival rate was affected by plant species or host strain, but they were affected by host spatial distribution. When host distribution was dense, a higher proportion of hosts were parasitized, and C. maculatus survival rate was lower. Here we discuss parasitoid potential as a selective agent for the sparse within-pod distribution of its hosts in the field.     

Host refractoriness of the tobacco hornworm, Manduca sexta, to the braconid endoparasitoid Cotesia flavipes

Cotesia flavipes (Hymenoptera:Braconidae) is a gregarious endoparasitoid of several pyralid stemborer larvae of economic significance including the sugarcane borer, Diatraea saccharalis. In this study, the ability of this parasitoid to develop in a sphingid host, Manduca sexta, was tested. First, second, third, fourth, and even pharate fifth instar host tobacco hornworm larvae were readily parasitized by the female C. flavipes parasitoids but no wasp larvae hatched from the eggs in this refractory host. Instead, the parasitoid eggs were invariably encapsulated by the host's hemocytes and, ultimately, no parasitoids emerged from tobacco hornworm hosts. The first stages of encapsulation were evident at 2 h post-parasitization of the host M. sexta larvae, when the beginning stages of capsule formation were seen. The developmental fate of the host larvae with encapsulated parasitoids was variable. Most succumbed as abnormally small fifth instars or as post-wandering prepupal animals, while a few developed normally to the pupal stage. Dissection of all the larvae or pupae with encapsulated wasp eggs showed evidence of hemocytic encapsulation and melanization of the C. flavipes eggs. This report describes the association between C. flavipes and M. sexta, which appears to be an excellent model system for studying the physiological processes accompanying wasp egg encapsulation that result in death of the host as well as the parasitoid. Since the parasitoid egg never hatches, the system offers an excellent opportunity to identify and study the effects of parasitoid-injected polydnavirus and venom on host physiology.     

Interspecific competition between the braconid endoparasitoids Glyptapanteles porthetriae and Glyptapanteles liparidis in Lymantria dispar larvae

The effect of interspecific competition between the solitary endoparasitoid Glyptapanteles porthetriae Muesebeck (Hymenoptera: Braconidae) and the gregarious Glyptapanteles liparidis Bouché (Hymenoptera: Braconidae), was investigated in larvae of Lymantria dispar L. (Lepidoptera: Lymantriidae). Host larvae were parasitized by both wasp species simultaneously in premolt to the 2^nd or the 3^rd host instar or in an additional approach with a 4‐day delay in parasitization by the second wasp species. Host acceptance experiments revealed that both wasp species do not discriminate between unparasitized host larvae and larvae parasitized previously by the same or the other species. In more than 90% female wasps parasitized the larva they encountered first. During the period of endoparasitic development, larvae of the competing parasitoid species never attacked the egg stage of the other species. When host larvae were parasitized simultaneously by both wasp species, the rate of successful development of both species depended on the age of the host larva at the time of its parasitization; G. liparidis emerged successfully from 44% of host larvae parasitized during the premolt to 2^nd instar, G. porthetriae from 28%, and in 20% of the hosts both parasitoid species were able to develop in one gypsy moth larva. However, when host larvae were parasitized simultaneously during premolt to the 3^rd instar, G. liparidis was successful in 90% of the hosts, compared to 8% from which only G. porthetriae emerged. In the experiments with delayed oviposition, generally the species that oviposited first succeeded in completing its larval development. Larvae of the species ovipositing with four days delay were frequently attacked and killed by larvae of the first parasitizing species or suffered reduced growth. As the secondary parasitoid species, G. porthetriae‐larvae were never able to complete their development, whereas G. liparidis developed successfully in at least 12,5% of the multiparasitized host larvae. Thus, multiparasitism of gypsy moth larvae by both Glyptapanteles species corresponds to the contest type; however, G. porthetriae is only able to develop successfully as the primary parasitoid of young host larvae.     

高效液相色谱法检测菜蛾绒茧蜂幼虫体内多杀菌素残留

应用高效液相色谱法检测施用于寄主幼虫的多杀菌素可否传递到在其体内发育的寄生蜂幼虫。以对多杀菌素高抗的小菜蛾Plutella xylostella幼虫作为菜蛾绒茧蜂Cotesia plutellae的寄主,待绒茧蜂发育到1龄幼虫时,将浓度为50 mg/L的多杀菌素点滴到寄主幼虫背板上,随后让寄主幼虫取食经50 mg/L多杀菌素处理过的甘蓝叶片,寄主幼虫和其体内的蜂幼虫再发育3天后,将寄主幼虫解剖取出蜂幼虫,用高效液相色谱法对经多杀菌素处理的小菜蛾幼虫体液以及绒茧蜂幼虫匀浆液进行检测,结果多杀菌素的2个活性成分spinosyn A和spinosyn D均被检测到,两者的多杀菌素残留浓度分别是2.79 mg/L和0.94 mg/L。这表明,通过寄主幼虫体壁接触和取食进入其体内的多杀菌素,可通过寄生蜂幼虫体壁浸透、蜂幼虫对寄主血淋巴的取食,或这两种途经一起进入蜂幼虫的体内,对蜂幼虫产生作用。     

Interspecific competition between the braconid endoparasitoids Glyptapanteles porthetriae and Glyptapanteles liparidis in Lymantria dispar larvae

The effect of interspecific competition between the solitary endoparasitoid Glyptapanteles porthetriae Muesebeck (Hymenoptera: Braconidae) and the gregarious Glyptapanteles liparidis Bouché (Hymenoptera: Braconidae), was investigated in larvae of Lymantria dispar L. (Lepidoptera: Lymantriidae). Host larvae were parasitized by both wasp species simultaneously in premolt to the 2^nd or the 3^rd host instar or in an additional approach with a 4-day delay in parasitization by the second wasp species. Host acceptance experiments revealed that both wasp species do not discriminate between unparasitized host larvae and larvae parasitized previously by the same or the other species. In more than 90% female wasps parasitized the larva they encountered first. During the period of endoparasitic development, larvae of the competing parasitoid species never attacked the egg stage of the other species. When host larvae were parasitized simultaneously by both wasp species, the rate of successful development of both species depended on the age of the host larva at the time of its parasitization; G. liparidis emerged successfully from 44% of host larvae parasitized during the premolt to 2^nd instar, G. porthetriae from 28%, and in 20% of the hosts both parasitoid species were able to develop in one gypsy moth larva. However, when host larvae were parasitized simultaneously during premolt to the 3^rd instar, G. liparidis was successful in 90% of the hosts, compared to 8% from which only G. porthetriae emerged. In the experiments with delayed oviposition, generally the species that oviposited first succeeded in completing its larval development. Larvae of the species ovipositing with four days delay were frequently attacked and killed by larvae of the first parasitizing species or suffered reduced growth. As the secondary parasitoid species, G. porthetriae-larvae were never able to complete their development, whereas G. liparidis developed successfully in at least 12,5% of the multiparasitized host larvae. Thus, multiparasitism of gypsy moth larvae by both Glyptapanteles species corresponds to the contest type; however, G. porthetriae is only able to develop successfully as the primary parasitoid of young host larvae.     

Cellular immune competence of a Drosophila mutant with neoplastic hematopoietic organs

In the Tum^l mutant of Drosophila melanogaster, the larval hematopoietic organs undergo neoplastic changes and release into circulation large numbers of blood cells. The lamellocytes, and to a lesser extent the plasmatocytes from which they are derived, are the cells that encapsulate various endogenous tissues and form melanotic tumors. The mutation is temperature sensitive, with maximum gene expression manifested at 29°C. The ability of Tum^l larvae to encapsulate eggs of the wasp parasite Leptopilina heterotoma is dependent not only on temperature, with host larvae much more immune reactive at 29°C than at lower temperatures (15° or 21°C), but also on the interval of time following infection when temperature shift experiments are performed. When the shift of parasitized larvae from 21° to 29°C is delayed by 18 hr the hosts are not as immune reactive as those shifted immediately after infection. Since Tum^l larvae are potentially highly immune reactive at the time of infection (with sufficient numbers of lamellocytes in circulation to encapsulate parasites), the low degree of immune competence in hosts shifted to 29°C after 18 hr or maintained at lower temperatures suggests that the increased capacity of blood cells to react against foreign surfaces is dependent on the cells acquiring new or altered recognition and adherence properties at 29°C. The 18-hr delay may provide the parasite with an opportunity to interfere with the acquisition of these specific cellular alterations. Differential hemocyte counts from parasitized larvae show abnormally low lamellocyte counts in susceptible hosts, indicating that successfully developing parasites interfere with the differentiation of hemocytes.     

Immunogenetic aspects of the cellular immune response of Drosophila against parasitoids

Host-parasite relationships represent integrating adaptations of considerable complexity involving the host's immune capacity to both recognize and destroy the parasite, and the latter's ability to successfully invade the host and to circumvent its immune response. Compatibility in Drosophila-parasitic wasp (parasitoid) associations has been shown to have a genetic basis, and to be both species and strain specific. Studies using resistant and susceptible strains of Drosophila melanogaster infected with virulent and avirulent strains of the wasp Leptopilina boulardi demonstrate that the success of the host cellular immune response depends on the genetic status of both host and parasitoid. Immunological, physiological, biochemical, and genetic data form the bases of a two-component model proposed here to account for the observed specificity and complexity of two coevolved adaptations, host nonself recognition and parasitoid virulence.     

Gregarious development of the solitary endo-parasitoid, Microplitis rufiventris in its habitual host, Spodoptera littoralis

Abstract:  Earlier research has shown that the koinobiont parasitoid, Microplitis rufiventris , attacks and can develop on early instars of Spodoptera littoralis larvae with preference to third instars. However, the present study was carried out using the newly moulted sixth instar larvae at two different temperatures (20 ± 1 and 27 ± 1°C) to study the developmental interaction between the parasitoid and the last instar host larvae. Parasitoid eggs laid in singly parasitized host larvae invariably died. As the number of parasitoid eggs/host larvae increased, the proportion of eggs that hatched and number of viable parasitoid larvae successfully reached to their final instar increased. The effect of superparasitization seems to be dose (no. of eggs + parasitoid factors)-temperature-dependent. The results demonstrate a kind of 'Allee effect' suggesting that superparasitized last instar S. littoralis larvae provide a better host environment than singly parasitized hosts for the parasitoid, M. rufiventris . This may be due to host's hormone and/or low dose of factors injected with parasitoid eggs. The supernumerary individuals of wasp larvae developed normally to the point of emergence but most did not successfully emerge from the host. The improvement of both hatchability and post-embryonic development of parasitoid wasp was significantly (P < 0.01) greater at 20°C than at 27°C. The results of the present study are useful in understanding the evolution of life-history strategies and host range in parasitic hymenoptera.