Véronique Martel

Size-induced reproductive constraints in an egg parasitoid.

The size of adult parasitoids is influenced by the quantity and quality of resources available during immature development. Gregarious development of endoparasitoids results in scramble competition where the resources are shared among individuals developing into the same host. Individuals that developed gregariously are therefore smaller and that reduced size generally translates into lower fitness due to reductions in several life history traits including longevity, mobility and traits linked to reproduction. We measured the reproductive constraints induced by size in Anaphes listronoti (Hymenoptera: Mymaridae), a facultative gregarious egg parasitoid of Curculionidae. Size decreased with number of immatures developing in a host egg for both male and female A. listronoti. This reduction in size induced a reproductive cost as both males and females produced fewer gametes when developing gregariously. Contrarily to other egg parasitoids, A. listronoti males are not prospermatogenic and produced some sperm during their adult life. Their spermatogeny index is estimated at 0.6-0.7 that places this species as moderately synspermatogenic. Female A. listronoti have an ovigeny index of 0.70 and are therefore moderately synovigenic. Large females that developed singly received a full sperm complement only when mated by a large male that also developed singly. When mating with a small triplet male, a large female received less than half her sperm complement. Large males were able to mate three females before the number of sperm transferred started to decrease.

Seasonal Parasitism and Host Instar Preference by the Spruce Budworm (Lepidoptera: Tortricidae) Larval Parasitoid Tranosema rostrale (Hymenoptera: Ichneumonidae)

Abstract The seasonal pattern of parasitism by a parasitoid can be influenced by many factors, such as interspecific competition and host instar preference. We conducted field and laboratory experiments to describe the seasonal pattern of parasitism of spruce budworm Choristoneura fumiferana (Clemens) larvae by Tranosema rostrale (Brischke), and to investigate whether this pattern can be explained by interaction with other parasitoid species, or by host instar preference. Larval survival, developmental time, sex ratio, and adult size of T. rostrale developing in different host instars were also measured to further assess the potential importance of host instar on parasitoid life history. Parasitism by T. rostrale increased over the season, reaching the highest rate during the fourth-instar larva, and then decreased again until the sixth-instar. At the same time, parasitism by another parasitoid, Elachertus cacoeciae (Howard), increased over the season, and multiparasitism with T. rostrale suggests potential competition between these two parasitoids. Tranosema rostrale showed no host instar preference when third- to sixth-instar larvae were exposed simultaneously in a manipulative field experiment. The proportion of females emerging from spruce budworm larvae increased over the season; however, no difference in sex ratio was observed in the manipulative field experiment. Only male pupal development time and adult size were marginally increased in fifth-instar spruce budworm larvae. We conclude that T. rostrales seasonal phenology or competition with E. cacoeciae, but not host instar preference, were possibly responsible for the observed seasonal pattern of parasitism.

Developmental and reproductive responses of the spruce budworm (Lepidoptera: Tortricidae) parasitoid Tranosema rostrale (Hymenoptera: Ichneumonidae) to temperature

The temperature-dependent development and survival of immatures, as well as adult longevity and potential fecundity of the endoparasitoid Tranosema rostrale (Hymenoptera: Ichneumonidae) parasitizing spruce budworm Choristoneura fumiferana (Lepidoptera: Tortricidae) larvae was investigated under laboratory conditions at several constant temperatures ranging from 5 to 30°C. Maximum likelihood modeling approaches were used to estimate thermal responses in development, survival, and longevity. A model describing the effect of temperature on potential fecundity of the parasitoid was also developed taking oogenesis and oosorption into account. In-host and pupal development rates of the parasitoid increased with temperature up to 25°C, and decreased thereafter. Immature survival was highest below 20°C, and rapidly decreased at higher temperatures. Adult longevity decreased exponentially with increasing temperature for both males and females. Highest potential fecundity was reached at 10°C. Considering survival and potential fecundity, the parasitoid seems best adapted to cool temperatures below 20°C. Simulations of the life-history traits under variable temperature regimes indicate that temperature fluctuations decrease survival and increase realised fecundity compared to constant temperatures. The temperature-dependent fecundity model developed can be applied to other non-host-feeding synovigenic parasitoids. The equations and parameter estimates provided in this paper can be used to build comprehensive models predicting the seasonal phenology of this parasitoid and spruce budworm parasitism under changing climatic conditions.

High temperature induces downregulation of polydnavirus gene transcription in lepidopteran host and enhances accumulation of host immunity gene transcripts.

Endoparasitoids face the challenge of overcoming the immune reaction of their hosts, which typically consists of encapsulation and melanisation of parasitoid eggs or larvae. Some endoparasitic wasps such as the solitary Tranosema rostrale (Hymenoptera: Ichneumonidae) that lay their eggs in larvae of the spruce budworm, Choristoneura fumiferana (Lepidoptera: Tortricidae), have evolved a symbiotic relationship with a polydnavirus (PDV), which in turn helps them suppress the hosts immune response. We observed an increase in mortality of immature T. rostrale with increasing temperature, and we tested two hypotheses about the mechanisms involved: high temperatures (1) hamper the expression of T. rostrale PDV genes and (2) enhance the expression of spruce budworm immunity-related genes. Dissections of parasitized spruce budworm larvae reared at 30°C revealed that most parasitoid eggs or larvae had died as a result of encapsulation and melanisation by the host. A qPCR analysis of T. rostrale PDV (TrIV) gene expression showed that the transcription of several TrIV genes in host larvae was downregulated at high temperature. On the other hand, encapsulation, but not melanisation, of foreign bodies in spruce budworm larvae was enhanced at high temperatures, as shown by the injection of Sephadex™ beads into larvae. However, at the molecular level, the transcription of genes related to spruce budworms melanisation process (prophenoloxidase 1 and 2) was upregulated. Our results support the hypothesis that a temperature-dependent increase of encapsulation response is due to the combined effects of reduced expression of TrIV genes and enhanced expression of host immune genes.

Sperm-depleted males influence the reproductive behaviour of conspecifics

In many insect species, sperm-depleted males (SDMs, i.e. males that have exhausted their sperm after a given number of matings) remain sexually active and continue to mate females. Here, we investigated the behavioural modifications that occur in both sexes of the parasitoid Asobara tabida Nees (Hymenoptera: Braconidae), after matings by fertile males and sperm-depleted males. We show that (i) virgin females, mated females and females mated to a SDM exhibited different behaviours and that (ii) males responded differently to females depending on whether the females had previously mated with an SDM or not. Our findings demonstrate that SDM influenced the reproductive behaviour of both males and females, especially with regard to male responsiveness and female attractiveness. These findings are discussed in the context of adaptive behaviour and fitness maximization in both males and females.

Impact of rearing temperature on encapsulation and the accumulation of transcripts putatively involved in capsule formation in a parasitized lepidopteran host

Encapsulation and melanisation are innate immune reactions of insects against foreign intruders such as parasitoids. In an earlier study, we observed that immature life stages of the endoparasitoid Tranosema rostrale (Hymenoptera: Ichneumonidae) parasitizing Choristoneura fumiferana (Lepidoptera: Tortricidae) larvae experienced higher mortality due to encapsulation and melanisation when reared at high (30 °C) than at lower (10 °C, 20 °C) temperatures. Downregulation of T. rostrale polydnavirus genes in parasitized hosts and upregulation of two genes involved in the spruce budworms melanisation process were identified as likely contributors to parasitoid mortality at high temperature. However, levels of transcripts of genes involved in the spruce budworms cellular encapsulation process were not measured inasmuch as candidate genes, in the spruce budworm, had not yet been identified. In addition, our assessment of temperature-dependent encapsulation and melanisation of foreign objects in spruce budworm larvae was only partial. To fill these knowledge gaps, we injected Sephadex™ beads into unparasitized spruce budworm larvae and assessed their encapsulation/melanisation after the insects had been held at three different temperatures (10, 20, and 30 °C), and we identified spruce budworm genes putatively involved in the encapsulation process and quantified their transcripts at the same three temperatures, using a qPCR approach. As expected, both encapsulation and melanisation of Sephadex™ beads increased as a function of temperature. At the molecular level, three of the five genes examined (Integrin β1, Hopscotch, Stat92E) clearly displayed temperature-dependent upregulation. The results of this study further support the hypothesis that a temperature-dependent increase in the encapsulation response of C. fumiferana against T. rostrale is due to the combined effects of reduced expression of polydnavirus genes and enhanced expression of host immune genes.