Comparative assessment of the thermal tolerance of spotted stemborer, Chilo partellus (Lepidoptera: Crambidae) and its larval parasitoid, Cotesia sesamiae (Hymenoptera: Braconidae).

Under stressful thermal environments, insects adjust their behavior and physiology to maintain key life-history activities and improve survival. For interacting species, mutual or antagonistic, thermal stress may affect the participants in differing ways, which may then affect the outcome of the ecological relationship. In agroecosystems, this may be the fate of relationships between insect pests and their antagonistic parasitoids under acute and chronic thermal variability. Against this background, we investigated the thermal tolerance of different developmental stages of Chilo partellus Swinhoe (Lepidoptera: Crambidae) and its larval parasitoid, Cotesia sesamiae Cameron (Hymenoptera: Braconidae) using both dynamic and static protocols. When exposed for 2 h to a static temperature, lower lethal temperatures ranged from -9 to 6 °C, -14 to -2 °C, and -1 to 4 °C while upper lethal temperatures ranged from 37 to 48 °C, 41 to 49 °C, and 36 to 39 °C for C. partellus eggs, larvae, and C. sesamiae adults, respectively. Faster heating rates improved critical thermal maxima (CTmax ) in C. partellus larvae and adult C. partellus and C. sesamiae. Lower cooling rates improved critical thermal minima (CTmin ) in C. partellus and C. sesamiae adults while compromising CTmin in C. partellus larvae. The mean supercooling points (SCPs) for C. partellus larvae, pupae, and adults were -11.82 ± 1.78, -10.43 ± 1.73 and -15.75 ± 2.47, respectively. Heat knock-down time (HKDT) and chill-coma recovery time (CCRT) varied significantly between C. partellus larvae and adults. Larvae had higher HKDT than adults, while the latter recovered significantly faster following chill-coma. Current results suggest developmental stage differences in C. partellus thermal tolerance (with respect to lethal temperatures and critical thermal limits) and a compromised temperature tolerance of parasitoid C. sesamiae relative to its host, suggesting potential asynchrony between host-parasitoid population phenology and consequently biocontrol efficacy under global change. These results have broad implications to biological pest management insect-natural enemy interactions under rapidly changing thermal environments.

Cost of Production of Telenomus remus (Hymenoptera: Platygastridae) Grown in Natural and Alternative Hosts.

The study of the technical and economic aspects of rearing natural enemies is essential for its effective use as a biological control agent in the field. The aim of this study was to determine the cost of production of the parasitoid, Telenomus remus (Nixon; Hymenoptera: Platygastridae), reared in eggs of its natural host, Spodoptera frugiperda (J. E. Smith; Lepidoptera: Noctuidae), and the alternative host, Corcyra cephalonica (Stainton; Lepidoptera: Pyralidae). The insects were reared in accordance with their respective methodologies of laboratory rearing. The cost of production of this biocontrol agent is US$ 0.0004 when reared with S. frugiperda eggs and US$ 0.0002 with C. cephalonica eggs.

Can the Use of Trichogramma ostriniae (Hymenoptera: Trichogrammatidae) to Control Ostrinia nubilalis (Lepidoptera: Crambidae) Be Economically Sustainable for Processing Sweet Corn?

European corn borer, Ostrinia nubilalis (Hübner) (Lepidoptera: Crambidae), is the main pest causing damage to sweet corn in North America. Conventional management with multiple use of insecticides is a common practice for processing sweet corn. In Canada, the use of Trichogramma spp. began in the 1990s, but the adoption of this approach for European corn borer management is still limited to the fresh market of sweet corn. Trichogramma ostriniae (Peng & Chen) has great potential as a biological control agent for large areas such as in processing sweet corn. The objective of this study was to evaluate an economically and environmentally sustainable alternative to insecticides for controlling European corn borer populations in processing sweet corn. During the growing season, the mean number of larvae decreased after insecticide (0.07 ± 0.04) and Trichogramma (1.32 ± 0.59) treatments compared with the control (2.42 ± 0.72). At harvest, damages associated with European corn borer were similar after Trichogramma (1.0 ± 0.7%) and insecticide (1.0 ± 0.6%) treatments, but significantly lower than the control (8.7 ± 3.3%). This study showed that the use of T. ostriniae can significantly decrease the pressure exerted by European corn borer and its damage on corn ears. This outcome is particularly interesting considering that it was achieved with lower doses of Trichogramma, a lower number of releases, and on large crop areas, compared with what is actually done to protect fresh market corn from European corn borer. Under these conditions, the use of Trichogramma is an economically and competitive alternative to insecticide applications.

A two-agent model applied to the biological control of the sugarcane borer (Diatraea saccharalis) by the egg parasitoid Trichogramma galloi and the larvae parasitoid Cotesia flavipes.

The paper is aimed at a methodological development in biological pest control. The considered one pest two-agent system is modelled as a verticum-type system. Originally, linear verticum-type systems were introduced by one of the authors for modelling certain industrial systems. These systems are hierarchically composed of linear subsystems such that a part of the state variables of each subsystem affect the dynamics of the next subsystem. Recently, verticum-type system models have been applied to population ecology as well, which required the extension of the concept a verticum-type system to the nonlinear case. In the present paper the general concepts and technics of nonlinear verticum-type control systems are used to obtain biological control strategies in a two-agent system. For the illustration of this verticum-type control, these tools of mathematical systems theory are applied to a dynamic model of interactions between the egg and larvae populations of the sugarcane borer (Diatraea saccharalis) and its parasitoids: the egg parasitoid Trichogramma galloi and the larvae parasitoid Cotesia flavipes. In this application a key role is played by the concept of controllability, which means that it is possible to steer the system to an equilibrium in given time. In addition to a usual linearization, the basic idea is a decomposition of the control of the whole system into the control of the subsystems, making use of the verticum structure of the population system. The main aim of this study is to show several advantages of the verticum (or decomposition) approach over the classical control theoretical model (without decomposition). For example, in the case of verticum control the pest larval density decreases below the critical threshold value much quicker than without decomposition. Furthermore, it is also shown that the verticum approach may be better even in terms of cost effectiveness. The presented optimal control methodology also turned out to be an efficient tool for the "in silico" analysis of the cost-effectiveness of different biocontrol strategies, e.g. by answering the question how far it is cost-effective to speed up the reduction of the pest larvae density, or along which trajectory this reduction should be carried out.

Review on Natural Enemies and Diseases in the Artificial Cultivation of Chinese Caterpillar Mushroom, Ophiocordyceps sinensis (Ascomycetes).

Ophiocordyceps sinensis (syn. Cordyceps sinensis), well known as DongChongXiaCao (DCXC), is one of the most valuable traditional Chinese medicinal species. In this article, we provide a systematic review of natural enemies and diseases encountered in artificial cultivation of DCXC. Unfortunately, DCXC has been endangered over the past decades due to overharvesting and a worsening ecological environment. Therefore, the artificial cultivation of DCXC has been extensively investigated in recent years. Complete indoor artificial cultivation and semi-field cultivation are the two most common strategies used to cultivate DCXC. However, cultured DCXCs are often attacked by various natural enemies and diseases, which have resulted in substantial loss of the valuable medicinal resource. In this study, we have summarized the species of natural enemies and types of diseases confronted by DCXC. Twenty reported natural enemy species are categorized into four classes, one of which is reported for the first time in this study. Moreover, six microbial pathogens are also discussed. The recapitulation of the natural enemies and diseases in DCXC artificial cultivation not only promote the development of integrated pest management of DCXC cultivation but also provide important information to help preserve and develop this valuable resource.

Costs and benefits of plant allelochemicals in herbivore diet in a multi enemy world.

Sequestration of plant defensive chemicals by herbivorous insects is a way of defending themselves against their natural enemies. Such herbivores have repeatedly evolved bright colours to advertise their unpalatability to predators, i.e. they are aposematic. This often comes with a cost. In this study, we examined the costs and benefits of sequestration of iridoid glycosides (IGs) by the generalist aposematic herbivore, the wood tiger moth, Parasemia plantaginis. We also asked whether the defence against one enemy (a predator) is also effective against another (a parasitoid). We found that the larvae excrete most of the IGs and only small amounts are found in the larvae. Nevertheless, the amounts present in the larvae are sufficient to deter ant predators and also play a role in defence against parasitoids. However, excreting and handling these defensive plant compounds is costly, leading to longer development time and lower pupal mass. Interestingly, the warning signal efficiency and the amount of IGs in the larvae of P. plantaginis are negatively correlated; larvae with less efficient warning signals contain higher levels of chemical defence compounds. Our results may imply that there is a trade-off between production and maintenance of coloration and chemical defence. Although feeding on a diet containing IGs can have life-history costs, it offers multiple benefits in the defence against predators and parasitoids.

Assessment of toxicity risk of insecticides used in rice ecosystem on Trichogramma japonicum, an egg parasitoid of rice lepidopterans.

Both chemical and biological methods are essential for control of insects, for example, lepidopterans, on rice. Thus, it is important to know the effect of chemicals on the biological control agents. In this study, we assessed the toxicity of commonly used insecticides on a biological control agent, Trichogramma japonicum Ahmead (an egg parasitoid of rice lepidopterans) by using a dry film residue method. Results showed that thirty insecticides from seven chemical classes exhibited various degree of toxicity to this parasitoid. Among the seven classes of chemicals tested, organophosphates (chlorpyrifos, fenitrothion, phoxim, profenofos, and triazophos) and carbamates (carbaryl, carbsulfan, isoprocarb, metolcarb, and promecarb) exhibited the highest intrinsic toxicity to T. japponicum, with an LC50 of 0.035 (0.029-0.044) to 0.49 (0.34-0.87) mg active ingredient (a.i.) L(-1), followed by antibiotics (abamectin, emamectin benzoate, and ivermectin), phenylpyrazoles (butane-fipronil, ethiprole, and fipronil), pyrethroids (cyhalthrin, cypermethrin, fenpropathrin, and lambda-cyhaothrin), and neonicotinoids (acetamiprid, imidacloprid, imidaclothiz, nitenpyram, thiacloprid, and thiamethoxam). Moreover, the insect growth regulator insecticides (chlorfluazuron, fufenozide, hexaflumuron and tebufenozide) exhibited the lowest toxicity to the wasps with an LC50 of 3,383 (2406-5499) to 30206 (23107-41008) mg ai. L(-1). Risk quotient analysis showed that phenylpyrazoles, pyrethroids, insect growth regulators, neonicotinoids (with the exception of thiamethoxam), and antibiotics (with the exception of abamectin) are classified as safe agents to the parasitoid, while organophosphates and carbamates are classified as slightly, moderately, or highly toxic agents to the parasitoid. The data presented in this paper provided useful information on the selection of compatible insecticides with T. japonicum.

Multi-objective behavioural mechanisms are adopted by foraging animals to achieve several optimality goals simultaneously.

1. Animals foraging for resources are under a variety of selective pressures, and separate optimality models have been developed predicting the optimal reproductive strategies they should adopt. 2. In most cases, the proximate behavioural mechanisms adopted to achieve such optimality goals have been identified. This is the case, for example, for optimal patch time and sex allocation in insect parasitoids. However, behaviours modelled within this framework have mainly been studied separately, even though real animals have to optimize some behaviours simultaneously. 3. For this reason, it would be better if proximate behavioural rules were designed to attain several goals simultaneously. Despite their importance, such multi-objective proximate rules remain to be discovered. 4. Based on experiments on insect parasitoids that simultaneously examine their optimal patch time and sex allocation strategies, it is shown here that animals can adopt multi-objective behavioural mechanisms that appear consistent with the two optimal goals simultaneously. 5. Results of computer simulations demonstrate that these behavioural mechanisms are indeed consistent with optimal reproductive strategies and have thus been most likely selected over the course of the evolutionary time.

Native range assessment of classical biological control agents: impact of inundative releases as pre-introduction evaluation.

Diadromus pulchellus Wesmael (Hymenoptera: Ichneumonidae) is a pupal parasitoid under consideration for introduction into Canada for the control of the invasive leek moth, Acrolepiopsis assectella (Zeller) (Lepidoptera: Acrolepiidae). Since study of the parasitoid outside of quarantine was not permitted in Canada at the time of this project, we assessed its efficacy via field trials in its native range in central Europe. This was done by simulating introductory releases that would eventually take place in Canada when a permit for release is obtained. In 2007 and 2008, experimental leek plots were artificially infested with pest larvae to mimic the higher pest densities common in Canada. Based on a preliminary experiment showing that leek moth pupae were suitable for parasitism up to 5-6 days after pupation, D. pulchellus adults were mass-released into the field plots when the first host cocoons were observed. The laboratory-reared agents reproduced successfully in all trials and radically reduced leek moth survival. Taking into account background parasitism caused by naturally occurring D. pulchellus, the released agents parasitized at least 15.8%, 43.9%, 48.1% and 58.8% of the available hosts in the four release trials. When this significant contribution to leek moth mortality is added to previously published life tables, in which pupal parasitism was absent, the total pupal mortality increases from 60.1% to 76.7%. This study demonstrates how field trials involving environmental manipulation in an agent's native range can yield predictions of the agent's field efficacy once introduced into a novel area.

A critical assessment of the effects of Bt transgenic plants on parasitoids.

The ecological safety of transgenic insecticidal plants expressing crystal proteins (Cry toxins) from the bacterium Bacillus thuringiensis (Bt) continues to be debated. Much of the debate has focused on nontarget organisms, especially predators and parasitoids that help control populations of pest insects in many crops. Although many studies have been conducted on predators, few reports have examined parasitoids but some of them have reported negative impacts. None of the previous reports were able to clearly characterize the cause of the negative impact. In order to provide a critical assessment, we used a novel paradigm consisting of a strain of the insect pest, Plutella xylostella (herbivore), resistant to Cry1C and allowed it to feed on Bt plants and then become parasitized by Diadegma insulare, an important endoparasitoid of P. xylostella. Our results indicated that the parasitoid was exposed to a biologically active form of the Cy1C protein while in the host but was not harmed by such exposure. Parallel studies conducted with several commonly used insecticides indicated they significantly reduced parasitism rates on strains of P. xylostella resistant to these insecticides. These results provide the first clear evidence of the lack of hazard to a parasitoid by a Bt plant, compared to traditional insecticides, and describe a test to rigorously evaluate the risks Bt plants pose to predators and parasitoids.

Ecological costs on local adaptation of an insect herbivore imposed by host plants and enemies.

Herbivore populations may become adapted to the defenses of their local hosts, but the traits that maximize host exploitation may also carry ecological costs. We investigated the patterns and costs of local adaptation in the pine processionary moth, Thaumetopoea pityocampa, to its host plants, Pinus nigra and P. sylvestris. The two hosts differ in needle toughness, a major feeding impediment for leaf-eating insects. We observed a west-to-east gradient of increasing progeny size in the Italian Alps, matching the pattern in toughness of their respective local host plant. Eastern populations that feed on the native P. nigra with tough needles had larger eggs, and neonate larvae with larger head capsules, than western populations that feed on the native P. sylvestris and the introduced P. nigra with softer foliage. In a reciprocal transfer experiment that involved the eastern-most and the western-most populations of T. pityocampa from this region, and excluded natural enemies, we found evidence for local adaptation to the host plant. Specifically, larvae from the western population only performed well when raised on their local hosts with soft needles, and they suffered near-complete mortality on the tough foliage at the eastern site. In contrast, larvae from the eastern population survived equally well at both sites. Local adaptation involved a trade-off between progeny size and the number of offspring. We hypothesized that an additional cost, imposed by natural enemies, may be associated with increased egg size: we also observed a west-to-east gradient of increased egg parasitism. We tested this hypothesis in a common garden by exposing eggs of both populations to parasitism by two native egg parasitoids, Ooencyrtus pityocampae and Baryscapus servadeii. The eastern population suffered a higher level of parasitoid attack by O. pityocampae than the western population, and performance of hatched adults of both parasitoids was enhanced in large eggs. Thus, increased neonate quality (larger eggs yielding larger larvae) confers an advantage on tough foliage but incurs the ecological cost of increased parasitism, which may constrain further adaptation by this herbivore.

Prophenoloxidases 1 and 2 from the spruce budworm, Choristoneura fumiferana: molecular cloning and assessment of transcriptional regulation by a polydnavirus.

Immune challenge in arthropods is frequently accompanied by melanization of the hemolymph, a reaction triggered by the activation of prophenoloxidase (PPO). Because their immature stages are spent inside the hemocoel of insect larvae, endoparasitoids have evolved strategies to escape or counter melanin formation. Very little molecular information is available on these endoparasitoid counterstrategies. We have sought to shed light on the inhibition of melanization in the spruce budworm, Choristoneura fumiferana, by the parasitic wasp Tranosema rostrale, by cloning two host PPO homologs and studying their transcriptional regulation after parasitization. The two polypeptides are encoded by transcripts of approximately 3.3 kb (for CfPPO1) and 3.0 kb (for CfPPO2) and possess structural features typical of other insect PPOs. While there appears to be a single CfPPO2 gene in the C. fumiferana genome, we detected three CfPPO1 mRNA variants displaying insertions/deletions in the 3' untranslated region, suggesting that there may be more than one CfPPO1 gene copy. Both CfPPO1 and CfPPO2 were expressed at high levels in C. fumiferana 6th instars, and parasitization by T. rostrale had no apparent impact on the level of their transcripts. Injection of a large dose (0.5 female-equivalent) of polydnavirus-laden calyx fluid extracted from T. rostrale, which is known to inhibit melanization in C. fumiferana, only caused a transient decrease in CfPPO1 and CfPPO2 transcript accumulation at 2-3 d post injection. It thus appears that transcriptional downregulation of C. fumiferana PPO by T. rostrale plays a minor role in the inhibition of hemolymph melanization in this host-parasitoid system.

Integrating biological and chemical controls in decision making: European corn borer (Lepidoptera: Crambidae) control in sweet corn as an example.

As growers switch to transgenic crops and selective insecticides that are less toxic to natural enemies, natural enemies can become more important in agricultural pest management. Current decision-making guides are generally based on pest abundance and do not address pest and natural enemy toxicity differences among insecticides or the impact of natural enemies on pest survival. A refined approach to making pest management decisions is to include the impact of natural enemies and insecticides, thereby better integrating biological and chemical control. The result of this integration is a dynamic threshold that varies for each product and the level of biological control expected. To demonstrate the significance of conserved biological control in commercial production, a decision-making guide was developed that evaluates control options for European corn borer, Ostrinia nubilalis (Hübner) (Lepidoptera: Crambidae), in sweet corn, Zea mays L., where the primary natural enemies are generalist predators. Management options are lambda-cyhalothrin (broad-spectrum insecticide), spinosad (selective insecticide), Trichogramma ostriniae (Peng and Chen) (Hymenoptera: Trichogrammatidae) (parasitoid), and Bacillus thuringiensis (Bt) sweet corn (transgenic variety). The key factors influencing thresholds for all treatments are the intended market, predator populations, and the presence of alternative foods for the predators. Treatment cost is the primary factor separating the threshold for each treatment within a common scenario, with the lowest cost treatment having the lowest pest threshold. However, when the impact of a treatment on natural enemies is projected over the 3-wk control period, the impact of the treatment on predators becomes the key factor in determining the threshold, so the lowest thresholds are for broad-spectrum treatments, whereas selective products can have thresholds > 6 times higher by the third week. This decision guide can serve as a framework to help focus future integrated pest management research and to aid in the selection of pest management tools.

Quantitative assessment of biotic mortality factors of the European corn borer (Lepidoptera: Crambidae) in field corn.

Five treatments were used to exclude naturally occurring predators and parasitoids, based on body size and flight ability, to assess their effect on Ostrinia nubilalis (Hübner) populations on corn plaits. Two initial O. nubilalis egg densities (one egg mass and three egg masses per plant) were assigned to each treatment. Egg predation was higher in uncaged treatments than in caged treatments. Flying insect predators, primarily Coleomegilla maculata DeGeer (Coleoptera: Coccinellidae), reduced egg densities by 50%. Thirty-five to 84% of O. nubilalis larvae were infected with Nosema pyrausta (Paillot) (Microspora: Nosematidae). The incidence of Beauveria bassiana (Balsamo) Vuillemin (Deuteromycotina: Hyphomycetes), ranged from 0 to 21%, whereas larval parasitism, mainly by Macrocentrus cingulum Reinhard (Hymenoptera: Braconidae) ranged from 0 to 31%. In contrast to previous studies, this 3-yr field study documents that egg predation and larval infections of O. nubilalis were significant and consistent biotic mortality factors.

Development of the one-on-one quality assessment assay for entomopathogenic nematodes.

The one-on-one bioassay was developed using Steinernema carpocapsae (All) nematodes against the wax moth larva, Galleria mellonella. The assay was used to develop and compare virulence profiles of both in vitro- and in vivo-produced nematodes and to provide a quality assessment 'standard' for in vitro-produced nematodes. The bioassay was subsequently used to develop virulence profiles for Steinernema carpocapsae (UK), S. feltiae (UK), S. feltiae (R1.5), S. feltiae (SN), S. glaseri (NJ-43), and S. riobrave (RGV). These profiles are unique for each species and isolate and are used as a standard of virulence in routine quality assessment of nematodes produced in liquid fermentation.

Neutral lipids and the assessment of infectivity in entomopathogenic nematodes: observations on four Steinernema species.

An 8-point visual index was developed for Oil Red O staining of neutral lipids in infective juveniles (IJs) of Steinernema carpocapsae (A11), S. riobravis (Biosys 355), S. feltiae (UK76) and S. glaseri (NC). The visual index was found to be a reliable and rapid method for determining the relative neutral lipid content of individual IJs and was validated quantitatively by gas chromatography. The relationship between neutral lipid utilization and infectivity of IJs stored in distilled water at 25 degrees C was also investigated and the first quantitative results on neutral lipid utilization in entomopathogenic nematodes are reported. Neutral lipid contents of freshly harvested IJs of S. carpocapsae, S. riobravis. S. feltiae and S. glaseri were 31, 31, 24 and 26% dry wt, respectively. Steinernema carpocapsae showed a sigmoidal pattern for neutral lipid utilization while S. riobravis used neutral lipids at an almost constant rate. Survivorship of these two species ranged between 120 and 135 days, whereas S. feltiae and S. glaseri lived > 450 days and had a slower rate of lipid utilization during a 260 day storage period. Oil Red O staining showed that individual IJs in each population utilized lipids at different rates, even though they had the same initial lipid index. The infectivity of S. riobravis, S. feltiae and S. glaseri declined with lipid utilization. In contrast, S. carpocapsae maintained a high level of infectivity even at relatively low lipid levels. Therefore, neutral lipid content was found to be a suitable indicator of infectivity for S. riobravis, S. feltiae and S. glaseri but not for S. carpocapsae.