Impact of a parasitoid on the bacterial symbiosis of its aphid host

Embryo production in aphids is absolutely dependent on the function of symbiotic bacteria, mainly Buchnera, and the growth and development of koinobiont parasitoids in aphids requires the diversion of nutrients from aphid embryo production to the parasitoid. The implication that the bacterial symbiosis may be promoted in parasitized aphids to support the growing parasitoid was explored by analysis of the number and biomass of mycetocytes, and the aphid cells bearing Buchnera, in the pea aphid Acyrthosiphon pisum Harris (Hemiptera: Aphididae) parasitized by the wasp Aphidius ervi Haliday (Hymenoptera: Braconidae). Aphids hosting a young larval parasitoid bore more mycetocytes of greater total biomass, and embryos of lower biomass than unparasitized aphids. Furthermore, one of the three aphid clones tested, which limited teratocyte growth (giant cells of parasitoid origin having a trophic role), bore smaller mycetocytes and larger embryos, than one or both of the two aphid clones with greater susceptibility to the parasitoid. These data suggest that susceptibility of the aphid‐Buchnera symbiosis to parasitoid‐mediated manipulation may, directly or indirectly, contribute to aphid susceptibility to parasitoid exploitation.     

Pea aphid clonal resistance to the endophagous parasitoid Aphidius ervi

The physiological mechanism of resistance to the endophagous braconid Aphidius ervi Haliday (Hymenoptera, Braconidae) by a pink clone (PC) of Acyrthosiphon pisum (Harris) (Homoptera, Aphididae) has been investigated. Comparative data on parasitoid development and associated host biochemical changes in the resistant PC aphids and in a susceptible green clone (GC) of A. pisum are reported. When the PC aphids were attacked as early 4th instars, the developing parasitoid larvae showed a strongly reduced increase in size, compared to those synchronously developing in GC aphids, and were unable to produce a regular mummy. In contrast, parasitism of 2nd instar PC aphids, allowed completion of parasitoid development, but adults had a prolonged developmental time, due to a longer duration of parasitoid’s final (3rd) instar. In all cases, teratocytes, cells deriving from the A. ervi serosal membrane, and the proteins abundantly synthesised by them, were never found in the haemolymph of parasitised PC aphids. Host castration, as demonstrated by total protein incorporation into reproductive tissues, was total in the majority of early (2nd instar) parasitised host aphids, while it was limited when later instars (4th) of PC aphids were parasitised. This is partly due to the absence of the cytolytic activity of teratocytes on host embryos, which, through their persistence, may compete for nutritional resources with the developing parasitoid larvae. In parasitised PC aphids, this competitive effect is further aggravated for the parasitoid by the absence of the regulated amino acid titre increase in the host haemolymph, which is regularly observed in GC aphids. Failure of teratocyte development in the PC clone of the pea aphid is, then, the major functional constraint accounting for the reduction/inhibition of A. ervi larval growth. The reported results allow to assess in vivo the role of teratocytes in the host physiological redirection and nutritional exploitation by the parasitoid, and to integrate and validate the proposed physiological model of host-parasitoid interactions in the system A. pisum-A.ervi.     

Does variation in host plant association and symbiont infection of pea aphid populations induce genetic and behaviour differentiation of its main parasitoid, Aphidius ervi?

The host-associated differentiation (HAD) hypothesis states that higher trophic levels in parasitic associations should exhibit similar divergence in case of host sympatric speciation. We tested HAD on populations of Aphidius ervi the main parasitoid of the pea aphid Acyrthosiphon pisum, emerging from host populations specialized on either alfalfa or red clover. Host and parasitoid populations were assessed for genetic variation and structure, while considering geography, host plant and host aphid protective symbionts Regiella insecticola and Hamiltonella defensa as potential covariables. Cluster and hierarchical analyses were used to assess the contribution of these variables to population structure, based on genotyping pea aphids and associated A. ervi with microsatellites, and host aphid facultative symbionts with 16S rDNA markers. Pea aphid genotypes were clearly distributed in two groups closely corresponding with their plant origins, confirming strong plant associated differentiation of this aphid in North America. Overall parasitism by A. ervi averaged 21.5 % across samples, and many parasitized aphids producing a wasp hosted defensive bacteria, indicating partial or ineffective protective efficacy of these symbionts in the field. The A. ervi population genetic data failed to support differentiation according to the host plant association of their pea aphid host. Potential for parasitoid specialization was also explored in experiments where wasps from alfalfa and clover aphids were reciprocally transplanted on alternate hosts, the hypothesis being that wasp behaviour and parasitic stages should be most adapted to their host of origin. Results revealed higher probability of oviposition on the alfalfa aphids, but higher adult emergence success on red clover aphids, with no interaction as expected under HAD. We conclude that our study provides no support for the HAD in this system. We discuss factors that might impair A. ervi specialization on its divergent aphid hosts on alfalfa and clover.     

Development of the aphid parasitoid,Aphidius ervi, in asexual and sexual females of the pea aphid,Acyrthosiphon pisum, and the blackberry-cereal aphid,Sitobion fragariae

The aphid parasitoid,Aphidius ervi Haliday, overwinters in larval diapause. The possibility that the parasitoid might prefer sexual (oviparae) rather than asexual females (virginoparae) as overwintering hosts (oviparae predominate in autumn when host numbers are generally declining) was tested by comparing these aphid morphs as potential hosts. Two host species were also examined, the pea aphid,Acyrthosiphon pisum (Harris), and the blackberry aphid,Sitobion fragariae (Walker). The parasitoids took longer to develop inS. fragariae than inA. pisum but the development of non-diapausingA. ervi was similar in sexual and asexual females. This observation, together with the greater variation in the duration of the different parasitoid stadia inS. fragariae, indicated that the parasitoid is specialized on the pea aphid. In photophases of 12 h and longer, the proportion ofA. ervi entering diapause inA. pisum oviparae was higher than in virginoparae. The critical daylength (where 50% of parasitoids entered diapause) was therefore longer in oviparae (12.6 h) than in virginoparae (11.7 h) with the inference that parasitoids developing in the oviparae would enter diapause earlier in the field. InS. fragariae, critical day-lengths were similar in both aphid morphs. The duration of diapause was unaffected by host morph and emergence in short days (10:14 L:D) occurred over a long period (c. 60 days).     

Avoidance of intraguild predation leads to a long-term positive trait-mediated indirect effect in an insect community

Intraguild predation among natural enemies is common in food webs with insect herbivores at their base. Though intraguild predation may be reciprocal, typically one species suffers more than the other and frequently exhibits behavioural strategies to lessen these effects. How such short-term behaviours influence population dynamics over several generations has been little studied. We worked with a model insect community consisting of two species of aphid feeding on different host plants (Acyrthosiphon pisum on Vicia and Sitobion avenae on Triticum), a parasitoid (Aphidius ervi) that attacks both species, and a dominant intraguild predator (Coccinella septempunctata) that also feeds on both aphids (whether parasitized or not). As reported previously, we found A. ervi avoided chemical traces of C. septempunctata. In population cages in the laboratory, application of C. septempunctata extracts to Vicia plants reduced parasitism on A. pisum. This did not increase parasitism on the other aphid species, our predicted short-term trait-mediated effect. However, a longer term multigenerational consequence of intraguild predator avoidance was observed. In cages where extracts were applied in the first generation of the study, parasitoid recruitment was reduced leading to higher population densities of both aphid species. S. avenae thus benefits from the presence of a dominant intraguild predator foraging on another species of aphid (A. pisum) on a different food plant, a long-term, trait-mediated example of apparent mutualism. The mechanism underlying this effect is hypothesized to be the reduced searching efficiency of a shared parasitoid in the presence of cues associated with the dominant predator.     

Host regulation by the aphid parasitoid Aphidius ervi: the role of teratocytes

The biochemical profile and metabolism of Acyrthosiphon pisum(Harris) (Homoptera, Aphididae) are markedly altered and redirected in response to parasitization by the endophagous braconid Aphidius erviHaliday (Hymenoptera, Braconidae). In the present study, the role played in the host regulation process by teratocytes, cells deriving from the dissociation of the embryonic membrane of the parasitoid, is taken into consideration. The protein synthesis activity of these cells of embryonic origin is analysed in vitroand an essential characterization of those proteins de novosynthesized and released in the incubation medium is provided. Teratocytes, obtained by dissecting parasitized host aphids, 3, 4 and 5 days after parasitoid oviposition, were incubated in vitroand, at the end of the incubation period, were separated from the medium for SDS-PAGE analysis of both cellular and secreted proteins. Various cellular proteins were more abundant as the time between parasitization and teratocyte collection increased. Furthermore, two proteins, showing an approximate molecular mass of 15 kD (p15) and 45 kD (p45) respectively, were abundantly secreted in the incubation medium by 5 day-old teratocytes. Incubations in presence of ³⁵S radiolabelled amino acids indicated that p15 and p45 are both synthesized by A. erviteratocytes. The amino acid composition of these two proteins was similar to that reported for other insect proteins with a demonstrated nutritional function. The p45 protein was found to be glycosylated. A tentative physiological model describing the host regulation role played by different parasitoid-derived factors is proposed.     

Waxy bloom in peas influences the performance and behavior of Aphidius ervi, a parasitoid of the pea aphid

Leaf surface waxy bloom can influence the predator–prey interactions that take place on peas. We tested whether the interaction between the pea aphid Acyrthosiphon pisum Harris (Homoptera: Aphidae) and a parasitoid, Aphidius ervi Haliday (Hymenoptera: Aphidiidae), is affected by reduced wax. We performed greenhouse experiments comparing aphid parasitism by individual A. ervi on two varieties of reduced wax peas to two normal wax sister varieties. We also observed the behavior of individual A. ervi in the greenhouse and measured field parasitism in small plots of reduced wax and normal wax peas. In the greenhouse, individual A. ervi parasitized more aphids on the reduced wax varieties than on their normal wax counterparts. Wasps spent more time actively foraging on reduced wax pea plants, which may contribute to the higher parasitism observed on those varieties. The greenhouse results suggested that the improved performance of individual A. ervi on reduced wax peas might contribute to a higher parasitism on reduced wax peas in the field. Field parasitism was significantly higher in reduced wax pea plots during 2000. Overall parasitism was higher in 2000 than in 2001 and 2002. In the latter years, parasitism was higher on reduced wax plants, but not significantly different from normal wax plants. Improved foraging by individual A. ervi resembles improved foraging by other carnivorous insects on reduced wax peas. The advantages of reduced wax for biological control of the pea aphid may hold when any of several different natural enemies is abundant.     

Plastic responses of some life history traits and cellular components of body size in Aphidius ervi as related to the age of its host Acyrthosiphon pisum

Phenotypic plasticity of wing size and shape has been evaluated in Aphidius ervi developing in its host, Acyrthosiphon pisum, parasitized at seven different ages. The parasitoid wing size was used as an estimator of both whole body size and its cellular composition. No size difference was observed in A. ervi adults emerged from aphids 1, 2 or 3 days old at parasitization. Body size then increased in A. ervi emerged from hosts older at parasitization. Body size values as related to host age at parasitization were achieved by adjusting developmental time, developmental rate or both. Parasitoids of similar size, but developed in hosts parasitized at different ages, had different wing cellular composition, while the increase of parasitoid body size was related to a general increase in both cell area and cell number. These results seem to suggest a trade‐off between adult size and developmental time, at least for parasitoids developed at the two extremes of host ages at parasitization, and that A. ervi can reach the same adult size via different trajectories, adapting its ontogenetic processes. Wing shape was typical for all the different parasitoid classes considered and differed strongly between males and females, independent of their size. Parasitoid males (haploids) and females (diploids) did not differ in either cell area or cell number, suggesting a possible sex‐determined dosage compensation in somatic tissue endoreplication. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 439–454.     

Survival to Parasitoids in an Insect Hosting Defensive Symbionts: A Multivariate Approach to Polymorphic Traits Affecting Host Use by Its Natural Enemy

Insect parasitoids and their insect hosts represent a wide range of parasitic trophic relations that can be used to understand the evolution of biotic diversity on earth. Testing theories of coevolution between hosts and parasites is based on factors directly involved in host susceptibility and parasitoid virulence. We used controlled encounters with potential hosts of the Aphidius ervi wasp to elucidate behavioral and other phenotypic traits of host Acyrthosiphon pisum that most contribute to success or failure of parasitism. The host aphid is at an advanced stage of specialization on different crop plants, and exhibits intra-population polymorphism for traits of parasitoid avoidance and resistance based on clonal variation of color morph and anti-parasitoid bacterial symbionts. Randomly selected aphid clones from alfalfa and clover were matched in 5 minute encounters with wasps of two parasitoid lineages deriving from hosts of each plant biotype in a replicated transplant experimental design. In addition to crop plant affiliation (alfalfa, clover), aphid clones were characterized for color morph (green, pink), Hamiltonella defensa and Regiella insecticola symbionts, and frequently used behaviors in encounters with A. ervi wasps. A total of 12 explanatory variables were examined using redundancy analysis (RDA) to predict host survival or failure to A. ervi parasitism. Aphid color was the best univariate predictor, but was poorly predictive in the RDA model. In contrast, aphid host plant and symbionts were not significant univariate predictors, but significant predictors in the multivariate model. Aphid susceptibility to wasp acceptance as reflected in host attacks and oviposition clearly differed from its suitability to parasitism and progeny development. Parasitoid progeny were three times more likely to survive on clover than alfalfa host aphids, which was compensated by behaviorally adjusting eggs invested per host. Strong variation of the predictive power of intrinsic (body color) and extrinsic traits (symbionts, host plant), indicate that host variables considered as key predictors of outcomes strongly interact and cannot be considered in isolation.     

Sources of variation in dietary requirements in an obligate nutritional symbiosis

The nutritional symbiosis between aphids and their obligate symbiont, Buchnera aphidicola, is often characterized as a highly functional partnership in which the symbiont provides the host with essential nutrients. Despite this, some aphid lineages exhibit dietary requirements for nutrients typically synthesized by Buchnera, suggesting that some aspect of the symbiosis is disrupted. To examine this phenomenon in the pea aphid, Acyrthosiphon pisum, populations were assayed using defined artificial diet to determine dietary requirements for essential amino acids (EAAs). Six clones exhibiting dependence on EAAs in their diet were investigated further. In one aphid clone, a mutation in a Buchnera amino acid biosynthesis gene could account for the clone''s requirement for dietary arginine. Analysis of aphid F₁ hybrids allowed separation of effects of the host and symbiont genomes, and revealed that both affect the requirement for dietary EAAs in the clones tested. Amino acid requirements were minimally affected by secondary symbiont infection. Our results indicate that variation among pea aphids in dependence on dietary amino acids can result from Buchnera mutation as well as variation in the host genotype.     

EVOLUTION OF AN APHID-PARASITOID INTERACTION: VARIATION IN RESISTANCE TO PARASITISM AMONG APHID POPULATIONS SPECIALIZED ON DIFFERENT PLANTS

The evolution of associations between herbivorous insects and their parasitoids is likely to be influenced by the relationship between the herbivore and its host plants. If populations of specialized herbivorous insects are structured by their host plants such that populations on different hosts are genetically differentiated, then the traits affecting insect-parasitoid interactions may exhibit an associated structure. The pea aphid (Acyrthosiphon pisum) is a herbivorous insect species comprised of genetically distinct groups that are specialized on different host plants (Via 1991a, 1994). Here, we examine how the genetic differentiation of pea aphid populations on different host plants affects their interaction with a parasitoid wasp, Aphidius ervi. We performed four experiments. (1) By exposing pea aphids from both alfalfa and clover to parasitoids from both crops, we demonstrate that pea aphid populations that are specialized on alfalfa are successfully parasitized less often than are populations specialized on clover. This difference in parasitism rate does not depend upon whether the wasps were collected from alfalfa or clover fields. (2) When we controlled for potential differences in aphid and parasitoid behavior between the two host plants and ensured that aphids were attacked, we found that pea aphids from alfalfa were still parasitized less often than pea aphids from clover. Thus, the difference in parasitism rates is not due to behavior of either aphids or wasps, but appears to be a physiologically based difference in resistance to parasitism. (3) Replicates of pea aphid clones reared on their own host plant and on a common host plant, fava bean, exhibited the same pattern of resistance as above. Thus, there do not appear to be nutritional or secondary chemical effects on the level of physiological resistance in the aphids due to feeding on clover or alfalfa, and therefore the difference in resistance on the two crops appears to be genetically based. (4) We assayed for genetic variation in resistance among individual pea aphid clones collected from clover fields and found no detectable genetic variation for resistance to parasitism within two populations sampled from clover. This is in contrast to Henter and Via's (1995) report of abundant genetic variation in resistance to this parasitoid within a pea aphid population on alfalfa. Low levels of genetic variation may be one factor that constrains the evolution of resistance to parasitism in the populations of pea aphids from clover, leading them to remain more susceptible than populations of the same species from alfalfa.     

Parasitism byAphidius ervi (Hym.: Aphidiidae): Preference for pea aphid and blue alfalfa aphid (Hom.: Aphididae) and competition withA. Smithi

The pea aphidAcyrthosiphon pisum (Harris) and the blue alfalfa aphidA. kondoi Shinji are pests of alfalfaMedicago sativa L. worldwide. The parasitoidAphidium ervi Haliday attacks both species andA. smithi is host specific to pea aphid Experiments were conducted to determine the preference ofA. ervi forA. pisum andA. kondoi at three densities of hosts using the percentages of parasitism and superparasitism, and the distribution of eggs among hosts as criteria. Also examined was the outcome of competition between the two parasitoids when exposed to the same hosts. A. ervi attacks moreA. pisum when provided alone (no choice test) and when provided together in equal numbers withA. kondoi (choice test). There was no significant difference in the number of progeny produced by either parasitoid when a female of each species was introduced simultaneously into the same test unit containingA. pisum hosts. The effects of the pathogenPandora neoaphidis on this competition is reviewed.     

Antipredator responses of aphids to parasitoids change as a function of aphid physiological state

Under predation risk, prey may prioritize antipredator behaviours and sacrifice feeding. However, energetically constrained animals may choose to sacrifice or change antipredator responses and accept relatively greater risk in order to secure food. In this last case, the antipredator tactics chosen must balance safety and feeding in such a way that costs are minimized and benefits maximized. We studied the antipredator behaviour of pea aphid, Acyrthosiphon pisum (Harris) (Hemiptera: Aphididae) subjected to different periods of food deprivation, against the parasitoid Aphidius ervi (Hymenoptera: Braconidae). As the energetic internal stress of aphids increased, the predominant antipredator response changed from walking away and dropping to kicking behaviour, and parasitization avoidance decreased. Parasitoids did not show preference between food-deprived and nonfood-deprived aphids. Dropping and walking away reduced parasitization from 50 to 33%. These results support the hypothesis that the antipredator behaviour of an aphid changes as a function of internal stress. By performing less costly behaviour such as kicking under energetically constrained conditions, aphids seem to minimize their probability of energy shortfall. Given that aphid antipredator behaviour is a function of nutritional state, its occurrence under natural conditions may match host quality spatial distribution. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Influence of “protective” symbionts throughout the different steps of an aphid–parasitoid interaction

Microbial associates are widespread in insects, some conferring a protection to their hosts against natural enemies like parasitoids. These protective symbionts may affect the infection success of the parasitoid by modifying behavioral defenses of their hosts, the development success of the parasitoid by conferring a resistance against it or by altering life-history traits of the emerging parasitoids. Here, we assessed the effects of different protective bacterial symbionts on the entire sequence of the host-parasitoid interaction (i.e., from parasitoid attack to offspring emergence) between the pea aphid, Acyrthosiphon pisum, and its main parasitoid, Aphidius ervi and their impacts on the life-history traits of the emerging parasitoids. To test whether symbiont-mediated phenotypes were general or specific to particular aphid–symbiont associations, we considered several aphid lineages, each harboring a different strain of either Hamiltonella defensa or Regiella insecticola, two protective symbionts commonly found in aphids. We found that symbiont species and strains had a weak effect on the ability of aphids to defend themselves against the parasitic wasps during the attack and a strong effect on aphid resistance against parasitoid development. While parasitism resistance was mainly determined by symbionts, their effects on host defensive behaviors varied largely from one aphid–symbiont association to another. Also, the symbiotic status of the aphid individuals had no impact on the attack rate of the parasitic wasps, the parasitoid emergence rate from parasitized aphids nor the life-history traits of the emerging parasitoids. Overall, no correlations between symbiont effects on the different stages of the host–parasitoid interaction was observed, suggesting no trade-offs or positive associations between symbiont-mediated phenotypes. Our study highlights the need to consider various sequences of the host-parasitoid interaction to better assess the outcomes of protective symbioses and understand the ecological and evolutionary dynamics of insect–symbiont associations.     

Impact of environmental stress on aphid clonal resistance to parasitoids: Role of Hamiltonella defensa bacterial symbiosis in association with a new facultative symbiont of the pea aphid

Resistance to endoparasitoids in aphids involves complex interactions between insect and microbial players. It is now generally accepted that the facultative bacterial symbiont Hamiltonella defensa of the pea aphid Acyrthosiphon pisum is implicated in its resistance to the parasitoid Aphidius ervi. It has also been shown that heat negatively affects pea aphid resistance, suggesting the thermosensitivity of its defensive symbiosis. Here we examined the effects of heat and UV-B on the resistance of A. pisum to A. ervi and we relate its stability under heat stress to different facultative bacterial symbionts hosted by the aphid. For six A. pisum clones harboring four different facultative symbiont associations, the impact of heat and UV-B was measured on their ability to resist A. ervi parasitism under controlled conditions. The results revealed that temperature strongly affected resistance, while UV-B did not. As previously shown, highly resistant A. pisum clones singly infected with H. defensa became more susceptible to parasitism after exposure to heat. Interestingly, clones that were superinfected with H. defensa in association with a newly discovered facultative symbiont, referred to as PAXS (pea aphid X-type symbiont), not only remained highly resistant under heat stress, but also expressed previously unknown, very precocious resistance to A. ervi compared to clones with H. defensa alone. The prevalence of dual symbiosis involving PAXS and H. defensa in local aphid populations suggests its importance in protecting aphid immunity to parasitoids under abiotic stress.     

Matching the supply of bacterial nutrients to the nutritional demand of the animal host

Various animals derive nutrients from symbiotic microorganisms with much-reduced genomes, but it is unknown whether, and how, the supply of these nutrients is regulated. Here, we demonstrate that the production of essential amino acids (EAAs) by the bacterium Buchnera aphidicola in the pea aphid Acyrthosiphon pisum is elevated when aphids are reared on diets from which that EAA are omitted, demonstrating that Buchnera scale EAA production to host demand. Quantitative proteomics of bacteriocytes (host cells bearing Buchnera) revealed that these metabolic changes are not accompanied by significant change in Buchnera or host proteins, suggesting that EAA production is regulated post-translationally. Bacteriocytes in aphids reared on diet lacking the EAA methionine had elevated concentrations of both methionine and the precursor cystathionine, indicating that methionine production is promoted by precursor supply and is not subject to feedback inhibition by methionine. Furthermore, methionine production by isolated Buchnera increased with increasing cystathionine concentration. We propose that Buchnera metabolism is poised for EAA production at certain maximal rates, and the realized release rate is determined by precursor supply from the host. The incidence of host regulation of symbiont nutritional function via supply of key nutritional inputs in other symbioses remains to be investigated.     

Growth and development of Cardiochiles nigriceps viereck (hymenoptera,braconidae) larvae and their synchronization with some changes of the hemolymph composition of their host,Heliothis virescens (F.) (Lepidoptera,Noctuidae)

Larval development of the parasitoid Cardiochiles nigriceps Viereck occurs in the last instar larva of its host, Heliothis virescens (F.). This allows the parasitoid to exploit the nutritional increase in the biosynthetic activity occurring in the host in preparation for metamorphosis. To understand the biochemical basis of this host parasitoid developmental synchrony, we undertook host ligation studies and analyzed host hemolymph for proteins and glycerol esters. Parasitization affected the biochemical profile of the host. The hemolymph protein concentration of parasitized last instar H. virescens larvae increased through time, whereas unparasitized (control) larvae were characterized by a decrease in the protein titer when they reached the prepupal stage. The effect of parasitism on glyceride titers of host hemolymph was not as pronounced as the effect on proteins. Ligation conducted on 5th instar hosts, which were parasitized as 4th instars, affected parasitoid development in a time-dependent way. The percentage of successfully developing C. nigriceps larvae increased with the increase of the time interval between parasitization and ligation. Ligation performed before day 2 of the 5th larval instar of H. virescens completely inhibited parasitoid development. Ligations that disrupted parasitoid developmentwere associated with a low host hernolymph protein concentration. Parasitoid development was successful when hernolymph protein titer was high, as occurred when ligations were performed after day 3 of the 5th host instar in both control and parasitized larvae. Ligations in both situations resulted in a slight increase in glyceride titers. The results suggest that host proteins and/or some factor(s) associated with them may play a role in parasitoid growth and development. © 1993 Wiley-Liss, Inc.     

The performance of an aphid parasitoid is negatively affected by the presence of a circulative plant virus

Plant viruses and aphids can interact via contest competition for plant resources and induce changes in plant physiology, which can have effects on a third trophic level. The aim of this study was to determine how the interactions between a circulative plant virus and its aphid vector may affect the performance of an endoparasitoid and how parasitism may affect the efficiency of virus transmission by its aphid vector. The timing when parasitized aphids were transferred to virus-infected lettuce leaves was critical for the performance of A. ervi. Higher parasitoid larvae mortality, longer developmental times and lower percentages of mummification were detected on viruliferous/parasitized aphid nymphs when the time lag between parasitism and exposure to the virus was less than 24 h. No significant differences were detected in virus transmission rate between parasitized and non-parasitized M. euphorbiae aphids.     

The influence of light environment on host colour preference in a parasitoid wasp

1. Visual chromatic cues and contrast effects are widely used by insects in behaviours involving host/prey/mate‐finding and recognition. However, naturally changing light conditions may challenge the visual perception of cues for these organisms. 2. We used the host/parasitoid system Acyrthosiphon pisum/Aphidius ervi to determine if apparent visual preference of the wasp for green over pink host aphids was a visually based choice or a post‐attack mechanism based on host susceptibility depending on anti‐parasitoid symbiotic bacteria. 3. The study tested the ability of the wasp to recognise and attack pea aphid clones expressing variation based on colour and/or symbionts under a broad range of LED‐controlled light environments mimicking natural variations. 4. Results showed that the amount of reflected light of pink morphs was about half that of the green morphs in the cyan‐green components. Both host colours were recognised and attacked under all tested light conditions, even red light (660 nm). The previously reported preference of A. ervi for green pea aphids, was clear only for naive females given a choice between two aphid colours under all light environments, but quickly disappeared. 5. Wasps showed no tendency of avoiding oviposition in clones with defensive symbionts. 6. These findings suggest that variable rates of pea aphid parasitism by A. ervi in fields do not depend on host colour discrimination, but rather on susceptibility variation among aphid clones in allowing larval development after egg‐laying. Further studies should consider deeper investigation of the impact of red lights used in modified light environments in greenhouses and the proportion of host colour morph available.     

The impact of co-infection by a nucleopolyhedrovirus and the endoparasitoid Microplitis pallidipes on the total hemocyte count and composition in larval beet armyworm,Spodoptera exigua

The physiological effects of nucleopolyhedrovirus (NPV) infection and parasitism by Microplitis pallidipes (Hymenoptera: Braconidae) on the hemocytes of Spodoptera exigua (Lepidoptera: Noctuidae) larvae were examined. We found that compared to healthy (control) larvae, the total hemocyte count (THC) and granulocyte count in parasitized larvae increased 1 day after parasitization and then decreased, while the plasmatocyte count was not significantly affected for the first 5 days but was significantly enhanced on day 6 after parasitization. In parasitized + infected larvae, both the THC and granulocyte counts began be lower from day 1 compared to parasitized larvae, while the plasmatocyte count was generally lower than in parasitized larvae. Compared to the control, THC, and granulocyte counts of virus-infected larvae were higher 1 day after infection. Compared to that in virus-infected larvae, THC and granulocyte counts in parasitized + infected larvae began to decrease from day 1 while the plasmatocyte count generally decreased. We concluded that the host immune response of cell communities to parasitization by M. pallidipes was elicited during the development of the parasitoid egg, but that immune response was inhibited during larval development of parasitoids in the host body. Meanwhile, we found that NPV infection impeded the regulatory effect of M. pallidipes on host cellular immune responses, and parasitization by M. pallidipes similarly inhibited the host cellular immune response caused by NPV infection.