Does the solitary parasitoid Microplitis pennatulae use a combinatorial approach to manipulate its host?

Host manipulation is a strategy used by some parasites to enhance their transmission. These parasites use a combination of neuropharmacological, psychoneuroimmunological, genomic/proteomic, or symbiont-mediated mechanisms to manipulate their hosts. Bodyguard manipulation occurs when parasitized hosts guard parasitoid pupae to protect them from their natural enemies. Bodyguard-manipulated hosts exhibit altered behaviours only after the egression of parasitoid prepupae. Behavioural changes in post-parasitoid egressed hosts could have resulted from their altered physiology. Previous studies have shown that gregarious manipulative parasitoids induce multiple physiological changes in their host, but the physiological changes induced by solitary manipulative parasitoids are unknown. Microplitis pennatulae Ranjith & Rajesh (Hymenoptera: Braconidae) is a larval parasitoid of Psalis pennatula Fabricius (Lepidoptera: Erebidae). After the egression of parasitoid prepupae, P. pennatula stops its routine activities and protects the parasitoid pupa from hyperparasitoids by body thrashes. In this study, we looked into the physiological changes induced by the solitary manipulative parasitoid, M. pennatulae, in its host, P. pennatula, during various stages of parasitization. We considered octopamine concentration and phenoloxidase (PO) activity as biomarkers of physiological change. We also examined whether M. pennatulae has a symbiotic virus and whether the wasp transfers it to the host during parasitization. We found that octopamine concentration was low in the pre-parasitoid egressed host, but it was elevated after the parasitoid egressed. Phenoloxidase activity was lower in the pre- and post-parasitoid egressed host than in the unparasitized host. We also detected symbiotic bracovirus (BV) in the wasp ovaries and isolated the BV virulence gene from the parasitised host. Our study suggests that solitary parasitoids also induce multiple physiological changes to influence the host behaviour to their advantage, as is the case with the gregarious parasitoids.     

Evolution of parasitoid host preference and performance in response to an invasive host acting as evolutionary trap

The invasion of a novel host species can create a mismatch in host choice and offspring survival (performance) when native parasitoids attempt to exploit the invasive host without being able to circumvent its resistance mechanisms. Invasive hosts can therefore act as evolutionary trap reducing parasitoids'' fitness and this may eventually lead to their extinction. Yet, escape from the trap can occur when parasitoids evolve behavioral avoidance or a physiological strategy compatible with the trap host, resulting in either host‐range expansion or a complete host‐shift. We developed an individual based model to investigate which conditions promote parasitoids to evolve behavioral preference that matches their performance, including host‐trap avoidance, and which conditions lead to adaptations to the unsuitable hosts. The model was inspired by solitary endo‐parasitoids attacking larval host stages. One important aspect of these conditions was reduced host survival during incompatible interaction, where a failed parasitization attempt by a parasitoid resulted not only in death of her offspring but also in host killing. This non‐reproductive host mortality had a strong influence on the likelihood of establishment of novel host–parasitoid relationship, in some cases constraining adaptation to the trap host species. Moreover, our model revealed that host‐search efficiency and genetic variation in host‐preference play a key role in the likelihood that parasitoids will include the suboptimal host in their host range, or will evolve behavioral avoidance resulting in specialization and host‐range conservation, respectively. Hence, invasive species might change the evolutionary trajectory of native parasitoid species, which is important for predicting biocontrol ability of native parasitoids towards novel hosts.     

寄生蜂多分DNA病毒的基因表达产物及对寄主昆虫的生理作用

最近有关由多分DNA病毒介导的寄生蜂与寄主关系的研究主要集中在多分DNA病毒基因的表达和伴随寄主生理功能失常的分子机理上 ,本文介绍在寄生蜂作用于寄主时一些重要的PDV基因表达产物 ,包括CsPDV ,MdPDV ,CcPDV ,TrPDV ,HdPDV ,CrPDV和其它PDV的基因表达产物 ,并简叙这些基因表达产物对寄主的生理影响。     

Nutritional ecology of endoparasitic insects and their hosts: An overview

The impact of insect endoparasites (parasitoids) on the physiology and behaviour of their hosts is reviewed within the context of the nutritional ecology of the parasitoids and their hosts. Alterations in the consumption, utilization and allocation of food by parasitized hosts are common, as are internal changes in their metabolic physiology. Gregarious parasitoid species frequently increase feeding by their host larvae whereas solitary parasitoid species often reduce feeding and growth of their hosts. Many parasitoid-associated changes in host physiology and behaviour are interpreted to be of adaptive significance to parasitoids. Substantial circumstantial evidence suggests, and a few direct tests of such adaptive significance indicate, that parasitoids alter their hosts in ways beneficial to their own fitness. However, most of the changes in parasitized hosts are of unknown cause and undocumented significance to the parasitoids. Several relevant hypotheses are presented, and these require extensive evaluation (often requiring novel experimental approaches) before a thorough understanding of parasitoid nutritional ecology is established.     

Host‐searching responses to herbivory‐associated chemical information and patch use depend on mating status of female solitary parasitoid wasps

1. In a tritrophic interaction system consisting of plants, herbivores, and their parasitoids, chemicals released from plants after herbivory are known to play important roles for many female parasitoids to find their hosts efficiently. On the plant side, chemical information associated with herbivory can act as an indirect defence by attracting the natural enemies of the host herbivores. 2. However, mated and virgin females of haplodiploid parasitoids might not necessarily respond to such chemical cues in the same way. Since virgin females can produce only sons, they might refrain from searching for hosts to invest eggs until copulation, in order to produce both sexes. 3. Here, we investigated differential host‐searching behaviours shown by mated and virgin females in the solitary parasitoid wasp, Cotesia vestalis, in response to herbivory‐associated chemical information from cruciferous plants infested by their host larvae, Plutella xylostella. 4. Mated females showed a significantly higher flight preference for host‐infested plants over intact plants, while no preference was observed with virgin females. Mated females also showed more intensive antennal searching and ovipositor probing behaviours to leaf squares with wounds caused by hosts than did virgin females. Furthermore, mated females stayed longer in host patches with higher parasitism rates than virgin females. 5. These results indicate that mating status of C. vestalis females clearly influences their host‐searching behaviour in response to herbivory‐associated chemical information and patch exploitation. Female parasitoids seem to forage for hosts depending on their own physiological condition in a tritrophic system.     

The roles of ecological fitting, phylogeny and physiological equivalence in understanding realized and fundamental host ranges in endoparasitoid wasps

Co-evolutionary theory underpins our understanding of interactions in nature involving plant-herbivore and host-parasite interactions. However, many studies that are published in the empirical literature that have explored life history and development strategies between endoparasitoid wasps and their hosts are based on species that have no evolutionary history with one another. Here, we investigated novel associations involving two closely related solitary endoparasitoids that originate from Europe and North America and several of their natural and factitious hosts from both continents. The natural hosts of both species are also closely related, all being members of the same family. We compared development and survival of both parasitoids on the four host species and predicted that parasitoid performance is better on their own natural hosts. In contrast with this expectation, survival, adult size and development time of both parasitoids were similar on all (with one exception) hosts, irrespective as to their geographic origin. Our results show that phylogenetic affinity among the natural and factitious hosts plays an important role in their nutritional suitability for related parasitoids. Evolved traits in parasitoids, such as immune suppression and development, thus enable them to successfully develop in novel host species with which they have no evolutionary history. Our results show that host suitability for specialized organisms like endoparasitoids is closely linked with phylogenetic history and macro-evolution as well as local adaptation and micro-evolution. We argue that the importance of novel interactions and 'ecological fitting' based on phylogeny is a greatly underappreciated concept in many resource-consumer studies.     

生物因子对寄生蜂滞育的影响

在影响寄生蜂滞育的生物因子中,寄主及亲代是主要影响因子。寄主可以从种类、类型、生理状态及丰富度等方面影响寄生蜂的滞育:寄生不同种类的寄主,寄生蜂的滞育反应、滞育进度、滞育比例以及光周期反应表现等均有不同;寄主类型的影响在蚜茧蜂类群中表现最为突出,不同寄主类型能对蚜茧蜂的滞育诱导产生不同影响;此外,寄主生理状态和寄主丰富度也是影响寄生蜂滞育的重要因子。亲代对寄生蜂滞育的影响则通过其所经历的环境条件以及自身因素等方面来表现:亲代经历的光周期和温度等能显著影响子代的滞育,甚至起完全决定作用;在亲代自身因素的影响中,亲代虫龄差异是主要因素。本文同时对寄生蜂滞育的研究及应用前景也一并作了展望。     

寄生蜂成虫在寄主搜索过程中的学习行为

综述了寄生蜂寄主搜索过程中的学习行为的概念、过程、适应性意义及影响因子。学习是寄生蜂的一种普遍特征,但学习基本上只发生在成虫期,其中联系性学习是一个主要特征。学习可产生启动和嗜好性学习二种效应,在寄生蜂搜索寄主的各个步骤中发挥重要作用,使其在复杂多变的生存环境中可高效地识别和利用各种有用信息,提高搜索效率。学习的潜力及效应可依寄生蜂和寄主的食性专化程度、寄主的发育阶段、寄生蜂自身的生理状态及环境中刺激的性质等因子而变化。对寄生蜂学习行为的了解有助于发展对其行为调控的技术,提高寄生蜂对害虫控制的效能。     

Does mother really know best? Oviposition preference reduces reproductive performance in the generalist parasitoid Aphidius ervi

The reproductive success of female parasitoids is dependent on their ability to accurately assess the suitability of a host for larval development. For generalist parasitoids, which utilize a broad range of species and instars as hosts, a set of assessment criteria determines whether a host is accepted or rejected. The suitability of a host, however, can only be imperfectly assessed by the female parasitoid, which can result in the selection of lesser quality hosts for oviposition. In this study we explored the disparity between host quality and host preference using the generalist koinobiotic parasitoid Aphidius ervi Haliday (Hymenoptera: Aphidiidae) and the host Aulacorthum solani (Harris) (Homoptera: Aphididae), the foxglove aphid. The second instar hosts produced the highest level of reproductive success, while third and fourth instars resulted in a substantially reduced reproductive performance. When given a choice of host instars, parasitoids preferred the older hosts for oviposition disregarding their reduced suitability for larval development. Results are discussed in context of mechanisms involved in A. ervi host selection and biases in the criteria used to assess hosts that may arise when parasitoids transfer host species between generations.     

Spatial and temporal variation in host–parasitoid interactions: leafcutter ant hosts and their phorid parasitoids

1. Parasitoid–host interactions are important components of ecological communities. Although parasitoid–host interactions are strongly shaped by evolutionary history, the abundance of both the parasitoid and the host may have a role in determining the nature of the interaction once phylogenetic relationships are considered. 2. Leafcutter ants are hosts of phorid parasitoids and represent a well‐defined and specialised module within a larger network of ant–symbiont interactions. A low specificity host taxa and a positive association between host abundance and parasitoid interaction frequency were expected due to the close phylogenetic relatedness of the hosts. 3. The interactions among all species of leafcutter ants and their parasitoids were quantified in two localities with different species richness. This study also characterised the spatial‐temporal variability of these interactions, determined the patterns of parasitoid specificity and host selection, and tested for an association between host abundance and parasitoid interaction frequency. 4. Contrary to expectation, most parasitoid species were highly specialised and interaction frequency for parasitoid species was not related to host abundance. All host ant species were attacked by more than one phorid species. Some phorid species used more than one host species and showed preference for the same species over space and time, suggesting that there are physiological and/or behavioural restrictions on host use. 5. These results show that there is a tendency for specialisation even when hosts are highly similar in their ecology. From a biological control perspective, these parasitoids may be effective candidates, due to the high specificity of some species and little host‐use variation through time.     

Parasitism rate, parasitoid community composition and host specificity on exposed and semi-concealed caterpillars from a tropical rainforest

The processes maintaining the enormous diversity of herbivore—parasitoid food webs depend on parasitism rate and parasitoid host specificity. The two parameters have to be evaluated in concert to make conclusions about the importance of parasitoids as natural enemies and guide biological control. We document parasitism rate and host specificity in a highly diverse caterpillar-parasitoid food web encompassing 266 species of lepidopteran hosts and 172 species of hymenopteran or dipteran parasitoids from a lowland tropical forest in Papua New Guinea. We found that semi-concealed hosts (leaf rollers and leaf tiers) represented 84 % of all caterpillars, suffered a higher parasitism rate than exposed caterpillars (12 vs. 5 %) and their parasitoids were also more host specific. Semi-concealed hosts may therefore be generally more amenable to biological control by parasitoids than exposed ones. Parasitoid host specificity was highest in Braconidae, lower in Diptera: Tachinidae, and, unexpectedly, the lowest in Ichneumonidae. This result challenges the long-standing view of low host specificity in caterpillar-attacking Tachinidae and suggests higher suitability of Braconidae and lower suitability of Ichneumonidae for biological control of caterpillars. Semi-concealed hosts and their parasitoids are the largest, yet understudied component of caterpillar—parasitoid food webs. However, they still remain much closer in parasitism patterns to exposed hosts than to what literature reports on fully concealed leaf miners. Specifically, semi-concealed hosts keep an equally low share of idiobionts (2 %) as exposed caterpillars.     

Abundance,not diversity,of host beetle communities determines abundance and diversity of parasitoids in deadwood

Most parasites and parasitoids are adapted to overcome defense mechanisms of their specific hosts and hence colonize a narrow range of host species. Accordingly, an increase in host functional or phylogenetic dissimilarity is expected to increase the species diversity of parasitoids. However, the local diversity of parasitoids may be driven by the accessibility and detectability of hosts, both increasing with increasing host abundance. Yet, the relative importance of these two mechanisms remains unclear. We parallelly reared communities of saproxylic beetle as potential hosts and associated parasitoid Hymenoptera from experimentally felled trees. The dissimilarity of beetle communities was inferred from distances in seven functional traits and from their evolutionary ancestry. We tested the effect of host abundance, species richness, functional, and phylogenetic dissimilarities on the abundance, species richness, and Shannon diversity of parasitoids. Our results showed an increase of abundance, species richness, and Shannon diversity of parasitoids with increasing beetle abundance. Additionally, abundance of parasitoids increased with increasing species richness of beetles. However, functional and phylogenetic dissimilarity showed no effect on the diversity of parasitoids. Our results suggest that the local diversity of parasitoids, of ephemeral and hidden resources like saproxylic beetles, is highest when resources are abundant and thereby detectable and accessible. Hence, in some cases, resources do not need to be diverse to promote parasitoid diversity.     

Models for pest control: Complementary effects of periodic releases of sterile pests and parasitoids

Four host-parasitoid models that incorporate the simultaneous or sequential release during each generation of sterile hosts and parasitoids for control or eradication of host species are presented. The models are based on two modifications of the Nicholson-Bailey model which incorporate density regulation either in the host larvae or via parasitoid oviposition. Parasitization of host larvae and adults forms another comparison. The models indicate that the release of sterile hosts alone is more efficient than release of parasitoids alone in controlling the hosts if population regulation is in the parasitoids; otherwise, the release of parasitoids alone is more efficient. The release of both steriles and parasitoids is much more efficient than the release of either alone for either suppressing or eradicating the hosts. This greater efficiency in combination rather than separately appears to be a special case of a more general principle, which is that two pest control methods will mutually complement each other if their optimal actions in reducing host numbers are at very different host densities. This is the case for sterile releases (optimal at low host densities) and parasitoid inundation (optimal at high host densities).     

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.     

Effect of fertilized,unfertilized, and UV‐irradiated hosts on parasitism and suitability for Trichogramma parasitoids

Trichogramma spp. (Hymenoptera: Trichogrammatidae) parasitoids have been commonly used as biological control agents in insect pest management. Host quality is believed to influence parasitism, host preference, and suitability for parasitoids. To date, limited studies have compared the parasitism of Trichogramma parasitoids on fertilized, unfertilized, and sterilized host eggs. Hence, we studied the performance of three Trichogramma egg parasitoids, Trichogramma japonicum Ashmead, Trichogramma chilonis Ishii, and Trichogramma leucaniae Pang & Chen, on fertilized, unfertilized, and ultraviolet (UV)‐irradiated fertilized (UVF) eggs of rice moth, Corcyra cephalonica (Stainton) (Lepidoptera: Pyralidae). In a no‐choice test, T. japonicum and T. leucaniae parasitized significantly more fertilized or UVF than unfertilized hosts, and T. chilonis parasitized significantly more UVF than either fertilized or unfertilized hosts. In a choice test, all three Trichogramma parasitoids parasitized UVF hosts the most, and unfertilized hosts the least. There were similar percentages of adult emergence and female progeny among fertilized, unfertilized, and UVF hosts for all three Trichogramma parasitoids, except that T. japonicum had significantly lower adult emergence on fertilized hosts. We also found that all three Trichogramma parasitoids developed slower on unfertilized hosts. Regardless of host treatments, T. leucaniae had the longest developmental time and T. chilonis had the shortest. We conclude that Trichogramma parasitoids prefer parasitizing UVF eggs of C. cephalonica without negative effects on their emergence and sex allocation.     

Effect of host instar on host discrimination of heterospecific‐parasitised hosts by sympatric parasitoids

1. Interspecific competition among hymenopteran parasitoids may shape their behavioural strategies for host resource exploitation. In order to reduce or prevent competition, many parasitoid species have evolved the ability to discriminate between unparasitised hosts and hosts parasitised by another parasitoid species (i.e. heterospecific host discrimination). However, discriminatory ability might be affected by host instar. 2. This study reports the first results on whether host instar can influence the use of heterospecific‐parasitised hosts by sympatric parasitoids of the genus Aphytis (Hymenoptera: Aphelinidae). 3. Aphytis melinus and Aphytis chrysomphali discriminated between unparasitised and heterospecific‐parasitised hosts when they found a third‐instar host (high quality), with a tendency to multi‐parasitise. However, this discrimination was not observed in the second instar (lower size). 4. The behavioural strategies adopted towards multi‐parasitise third‐instar hosts varied between both species. Aphytis chrysomphali reduced its clutch size in heterospecific‐parasitised hosts, whereas A. melinus tended to probe them for longer than healthy hosts. 5. Overall, our results highlight the importance of host instar in the study of intrinsic competition between parasitoids.     

Nectar provisioning close to host patches increases parasitoid recruitment,retention and host parasitism

In the adult stage, many parasitoids require hosts for their offspring growth and plant-derived food for their survival and metabolic needs. In agricultural fields, nectar provisioning can enhance biological control by increasing the longevity and fecundity of many species of parasitoids. Provided in a host patch, nectar can also increase patch quality for parasitoids and affect their foraging decisions, patch time residence, patch preference or offspring allocation. The aim of this study was to investigate the impact of extrafloral nectar (EFN) provisioning close to hosts on parasitoid aggregation in patches. The aphid parasitoid Diaeretiella rapae (M’Intosh) was released inside or outside patches containing Brassica napus L. infested by Brevicoryne brassicae L. aphids and Vicia faba L. with or without EFN. When parasitoids were released outside patches, more parasitoids were observed in patches with EFN than in patches deprived of EFN. This higher recruitment could be linked to a higher attraction of a combination of host and food stimuli or a learning process. A release–recapture experiment of labeled parasitoids released within patches showed the higher retention of parasitoids in patches providing EFN and hosts, suggesting that food close to the host patch affects patch residence time. Both attractiveness and patch retention could be involved in the higher number of parasitoids foraging in host patches surrounded by nectar and for the higher parasitism recorded. Nectar provisioning in host patches also affected female offspring allocation inside the patch.     

The parasitoid factor in the virulence and spread of lepidopteran baculoviruses

Insect parasitoids and baculoviruses play important roles in the natural and strategic biological control of insects. The two parasites are frequent competitors within common hosts and much research has focused on the negative impact that baculoviral host infections have on parasitoids. This review summarizes the impacts that parasitoids may have on the virulence and spread of lepidopteran baculoviruses. By changing host behavior and development, parasitoids have been shown to decrease baculovirus virulence and productivity within parasitized baculovirus-susceptible hosts; however, studies of the tools used by hymenopteran parasitoids to overcome their hosts' immune systems, suggest that parasitoids may, in some cases, facilitate baculoviral infections in less susceptible hosts. Laboratory and field research have demonstrated that parasitoids can mechanically transmit bacuioviruses between insects, and in this way, increase the efficacy of the viruses. Instances of new, more virulent isolates of baculoviruses have been recorded from specifically parasitoid-targeted hosts suggesting other possible benefits from the transmission or activation of baculoviruses by parasitoids.     

Influence of physiological state and experience on the responsiveness ofLysiphlebus testaceipes (Cresson) (hymenoptera: aphidiidae) to aphid honeydew and to host plants

The effect of physiological state (hunger) and experience on the responsiveness of the aphid parasitoid, Lysiphlebus testaceipes(Cresson), to clean and honeydew-contaminated host plants was investigated in laboratory bioassays. Both fed and unfed parasitoids spent significantly longer examining honeydew-contaminated plants compared to uncontaminated controls, but the presence of honeydew did not influence attack latency (i.e., the speed with which naive parasitoids found and attacked hosts). Hunger, however, had a significant negative effect on attack latency, presumably as a result of a physiologically based shift from host- to food-location behavior in starved parasitoids. The parasitoid 's response to clean plants was significantly increased as a result of classical conditioning procedures, whereas the response to honeydew-contaminated plants was not.