Comparison of laboratory and field bioassays of laboratory‐reared Cydia pomonella (Lepidoptera: Tortricidae) quality and field performance

Maximum production and fitness of insect species that are mass‐reared for biological control programmes such as the sterile insect technique (SIT) have benefitted from the employment of quality control and quality management. With a growing interest in the use of SIT as a tactic for the suppression/eradication of key lepidopteran pests, such as the codling moth, Cydia pomonella L. (Lepidoptera: Tortricidae), there is a parallel interest in inexpensive bioassays that can accurately detect differences in insect quality and monitor insect field performance. In this study, we examined laboratory (mating and flight ability) bioassays and field (field cage and open field release) bioassays simultaneously to discern the ability of the different bioassays to predict quality and field performance of codling moths produced in a commercial mass‐rearing facility. Moth quality was degraded by different levels of radiation during the sterilization procedure. Both the laboratory flight bioassay and the field cage bioassay successfully detected quality and performance differences that were relevant to moth performance in the field. However, the study data suggest that the field cage bioassay was a better predictor of the daily performance of males that had been released in the orchard than the laboratory flight bioassay. Conversely, data suggest that the controlled climatic conditions of the laboratory allowed the flight cylinder bioassay to be more sensitive in detecting daily fluctuations in the quality of moths caused by factors within the mass‐rearing facility. Therefore, both laboratory and field bioassays may be required to provide feedback on quality and performance of mass‐reared moths in a SIT programme.     

Effect of rearing strategy and gamma radiation on fecundity and fertility of codling moth Cydia pomonella (Linnaeus) (Lepidoptera: Tortricidae)

The codling moth Cydia pomonella (Linnaeus) (Lepidoptera: Tortricidae) is a serious pest of pome fruit worldwide. In an effort to reduce the use of pesticides to control this pest, the sterile insect technique (SIT) is used or considered for use as a component of area‐wide integrated pest management programmes. Rearing codling moths through diapause has been shown to improve the competitiveness of sterile moths released in orchards, and provides management alternatives that would allow mass‐rearing facilities to increase their yearly production of sterile moths. Because radiosensitivity in insects can be influenced by numerous biological factors, laboratory tests were conducted to examine whether the response to increasing doses of radiation, as expressed in the fecundity and fertility of cohorts of moths, is similar for adult moths mass‐reared through diapause or through standard (non‐diapause) production protocols. Our data revealed that the effect of increasing doses of radiation on fecundity and fertility of codling moths reared through both rearing strategies was similar. In the case of fertility, this is a particularly important finding for the expanded application of codling moth SIT. If mass‐rearing facilities use year‐round diapause rearing, the dose required to treat the insects prior to release will be similar to that used when codling moths are reared through standard production protocols.     

Mating compatibility among populations of codling moth Cydia pomonella Linnaeus (Lepidoptera: Tortricidae) from different geographic origins

The codling moth Cydia pomonella (Linnaeus) (Lepidoptera: Tortricidae) is a serious pest of pome fruit worldwide and the sterile insect technique (SIT) provides an environmentally acceptable approach for its control. As the pest is present in both the southern and northern hemispheres it would be possible for a rearing facility in the northern hemisphere to supply sterile moths to an SIT programme in the southern hemisphere during the northern winter and vice versa. This could greatly improve the economics of moth production and the running costs of rearing facilities. However in order to develop this concept, it is important to assess if populations of codling moth from different geographical regions share mating compatibility. Twelve different laboratory and field populations from both hemispheres were sampled and field cage bisexual mating compatibility tests were carried out between selected combinations. The index of sexual isolation (ISI) and the female and male relative performance index (FRPI and MRPI, respectively) were calculated for each mating combination. In only two of the combinations was there a slight but significant deviation from random mating. There were also some significant differences in mating duration between the homotypic matings and the duration of a particular homotypic mating seemed to depend on the origin of the other population in the cage. It was concluded that there exist no barriers to mating between populations of codling moth from many parts of the world and that it would be feasible for sterile moths to be shipped from one rearing facility to SIT programmes in other parts of the world.     

Compatibility of codling moths Cydia pomonella (Linnaeus) (Lepidoptera: Tortricidae) from South Africa with codling moths shipped from Canada

The sterile insect technique (SIT) has been successfully applied against codling moth Cydia pomonella (Linnaeus) (Lepidoptera; Tortricidae) in British Columbia since 1992 where the mass‐rearing facility produces between 15 and 16 million moths per week. Due to the seasonality of this pest, the facility is only fully utilized for part of the year. The time and expense of implementing SIT against codling moth in South Africa may be substantially reduced if moths from Canada were sexually compatible with those from South Africa. In addition, because the pome fruit‐growing season in both countries is opposite, the programme in Canada might benefit by maintaining moth production year‐round and selling moths to South Africa. Semi‐field studies in small cages and release‐recapture studies were conducted in an unsprayed apple orchard in South Africa to assess mating compatibility of laboratory‐reared codling moth from Canada and wild codling moths from South Africa. The results suggest that Canadian codling moth males were equally attracted to calling Canadian and South African females despite the fact that Canadian moths had been transported (from Canada to South Africa) for 48 h as both pupae and adults. The data also suggest that at lower field temperatures Canadian moths were more active than South African moths. Results from the release‐recapture field trials indicated that Canadian and South African males were equally attracted to Canadian and South African females. These results suggest that codling moths from Canada and South Africa are fully compatible and indicate that Canadian moths can be used for SIT studies in South Africa. As these studies were conducted with moths from two very different climatic and time zones, it is proposed that populations of codling moth in other pome fruit production areas may also be compatible with the Canadian moths.     

Effect of radiation on fecundity and fertility of codling moth Cydia pomonella (Linnaeus) (Lepidoptera: Tortricidae) from South Africa

Codling moth Cydia pomonella (Linnaeus) (Lepidoptera: Tortricidae) is the key pest of pome fruit in South Africa, and it’s control in apple and pear orchards relies on the application of insecticides and pheromone‐mediated mating disruption. Development of resistance to insecticides and placement of restrictions on the use of certain insecticides has made control of codling moth in South Africa increasingly problematic. The use of the sterile insect technique (SIT) as a control tactic for codling moth is under investigation as a potential addition to the current control strategy. We investigated the radiosensitivity of a laboratory strain of codling moth that was established from moths collected from commercial and organic orchards in the Western Cape, South Africa. Fecundity and fertility of this strain following radiation were consistent with values for the codling moth strain in the Canadian rearing facility in British Columbia. For both strains, the female codling moth was considerably more radiosensitive than the male. At a radiation dose of 100 Gy or higher, treated females were 100% sterile. The fertility of the South African strain was higher (86.3%) than for the Canadian strain (71.9%). This difference in fertility between the two strains was maintained when the dose of radiation was 100 Gy. However, the level of fertility was very similar between the two strains for doses ≥150 Gy. Therefore, based upon previously published work and the data from this study, an operational dose of 150 Gy is recommended for future codling moth SIT programmes in South Africa.     

Improved quality management to enhance the efficacy of the sterile insect technique for lepidopteran pests

Lepidoptera are among the most severe pests of food and fibre crops in the world and are mainly controlled using broad spectrum insecticides. This does not lead to environmentally sustainable control and farmers are demanding alternative control tools which are both effective and friendly to the environment. The sterile insect technique (SIT), within an area‐wide integrated pest management (AW‐IPM) approach, has proven to be a powerful control tactic for the creation of pest‐free areas or areas of low pest prevalence. Improving the quality of laboratory‐reared moths would increase the efficacy of released sterile moths applied in AW‐IPM programmes that integrate the (SIT). Factors that might affect the quality and field performance of released sterile moths are identified and characterized in this study. Some tools and methods to measure, predict and enhance moth quality are described such as tests for moth quality, female moth trapping systems, ‘smart’ traps, machine vision for recording behaviour, marking techniques, and release technologies. Methods of enhancing rearing systems are discussed with a view to selecting and preserving useful genetic traits that improve field performance.     

Cold treatment enhances low‐temperature flight performance in false codling moth,Thaumatotibia leucotreta (Lepidoptera: Tortricidae)

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Quality of mass-reared codling moth (Lepidoptera: Tortricidae) after long-distance transportation: 1. Logistics of shipping procedures and quality parameters as measured in the laboratory

The sterile insect technique (SIT) is a proven effective control tactic against lepidopteran pests when applied in an areawide integrated pest management program. The construction of insect mass-rearing facilities requires considerable investment and moth control strategies that include the use of sterile insects could be made more cost-effective through the importation of sterile moths produced in other production centers. For codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), this is an attractive option because mating studies have confirmed the absence of mating barriers between codling moth populations from geographically different areas. To assess the feasibility of long-distance transportation of codling moths, pupae and adult moths were transported in 2004 from Canada to South Africa in four shipments by using normal commercial transport routes. The total transport time remained below 67 h in three of the consignments, but it was 89 h in the fourth consignment. Temperature in the shipping boxes was fairly constant and remained between -0.61 and 0.16 degrees C for 76.8-85.7% of the time. The data presented indicate that transporting codling moths as adults and pupae from Canada to South Africa had little effect on moth emergence, longevity, and ability to mate, as assessed in the laboratory. These results provide support to the suggestion that the STT for codling moth in pome fruit production areas might be evaluated and implemented by the importation of irradiated moths from rearing facilities in a different country or hemisphere.     

Mass‐rearing conditions do not affect responsiveness to sex pheromone and flight activity in sweetpotato weevils

For ensuring the effectiveness of sterile insect technique (SIT) programmes, maintaining the reproductive competitiveness and dispersal ability of mass‐reared sterile males is essential. Inadvertent selection is an important genetic process that frequently occurs during mass rearing to produce sterile males. We investigated the effect of mass‐rearing conditions on the responsiveness to sex pheromones and spontaneous flight activity of males of the sweetpotato weevil Cylas formicarius (Coleoptera: Brentidae). There were no significant differences in the responsiveness to sex pheromones and spontaneous flight activity between wild and mass‐reared strains. These results indicate that mass‐reared strains of C. formicarius might not cause serious problems for implementing SIT programmes.     

Potential for controlling codling moth Cydia pomonella (Linnaeus) (Lepidoptera: Tortricidae) in Argentina using the sterile insect technique and egg parasitoids

Codling moth is the main pest affecting apples and pears worldwide. Most pest control strategies used against this insect have relied on the use of broad‐spectrum insecticides which have led to non‐desirable effects like pesticide resistance, residues in the environment, human health concerns and the reduction of access to international markets. Therefore, alternative pest control strategies that would result in sustainable fruit production systems while taking care of the environment are strongly promoted. The use of the sterile insect technique has proven to be a valuable pest control tactic within area‐wide integrated pest management strategies, and its synergistic effect for Lepidoptera pests when combined with other biological control tactics such as parasitoids has been documented. The purposes of this research were to evaluate the response of an Argentinean codling moth strain to a sub‐sterilizing radiation dose of 100 Gy and to assess the acceptability and suitability of sterile codling moth eggs by the egg parasitoids, Trichogramma cacoeciae (Marchal) and Trichogramma nerudai (Pintureau and Gerding). Irradiated female moths survived better than irradiated male moths and non‐irradiated male and female moths. Also, the fecundity of irradiated female moths was reduced by more than 30% as compared to non‐irradiated ones whereas their fertility was close to zero. The F₁ generation was male biased with a lower fertility (inherited sterility) than the parental generation. Trichogramma cacoeciae and T. nerudai parasitized both fertile and sterile eggs. However, there was a significant reduction in acceptability for sterile eggs. Trichogramma nerudai parasitized more eggs than T. cacoeciae, but egg acceptability for this species was proportionally lower than for T. cacoeciae especially on eggs oviposited by irradiated females. Development to adult of both parasitoids species was not substantially affected by the origin of the eggs and the wasps had acceptable levels of adult emergence, survival and fecundity. These results provided useful information on the potential for controlling the codling moth using egg parasitoids and the sterile insect technique in Argentina.     

Improvement of the sterile insect technique for codling moth Cydia pomonella (Linnaeus) (Lepidoptera Tortricidae) to facilitate expansion of field application

The codling moth Cydia pomonella (L.) (Lepidoptera Tortricidae) is a key pest of pome fruit (apple, pear and quince) and walnut orchards in most temperate regions of the world. Efforts to control the codling moth in the past mostly relied on the use of broad spectrum insecticide sprays, which has resulted in the development of insecticide resistance, and the disruption of the control of secondary pests. In addition, the frequent reliance and use of these insecticides are a constant threat to the environment and human health. Consequently, there have been increased demands from the growers for the development of codling moth control tactics that are not only effective but also friendly to the environment. In that respect, the sterile insect technique (SIT) and its derivative, inherited sterility (IS), are, together with mating disruption and granulosis virus, among the options that offer great potential as cost‐effective additions to available control tactics for integration in area‐wide integrated pest‐management approaches. In support of the further development of the SIT/IS for codling moth control, the Joint FAO/IAEA Programme of Nuclear Techniques in Food and Agriculture implemented a 5‐year Coordinated Research Project (CRP) entitled ‘Improvement of codling moth SIT to facilitate expansion of field application’. Research focussed on sterile codling moth quality and management (e.g. mobility and life‐history traits in relation to rearing strategy, dispersal, flight ability, radiosensitivity and mating compatibility) and a better understanding of the basic genetics of codling moth to assist the development of genetic sexing strains (e.g. cytogenetics, the development of dominant conditional lethal mutations, molecular characterization of the sex chromosomes, sex identification in embryos and cytogenetic markers). The results of the CRP are presented in this special issue.     

Insecticidal resistance and cross-resistance in populations of Cydia pomonella (Lepidoptera: Tortricidae) in central Europe

Insecticide bioassays were used to investigate resistance of Cydia pomonella (L.) (Lepidoptera: Tortricidae) to insecticides with various types of active ingredients. The efficacy baselines of selected insect growth regulators (fenoxycarb), insect growth inhibitors (diflubenzuron and teflubenzuron), organophoshorous insecticides (phosalone), and neonicotinoids (thiacloprid) against the eggs and first and fifth instars of sensitive laboratory strains of codling moth were determined. According to concentration-mortality baseline, 50% lethality concentration values and 90% lethality concentration values were determined for all the tested insecticides. The lethal concentration ratio quantified the relation between the efficacy of selected insecticides against fifth instars found by topical application and against first instars found by diet-treated bioassay. No difference was detected when the efficacy of technical grade diflubenzuron diluted in tetrahydrofuran and diflubenzuron in the formulated product Dimilin 48 SC diluted in water was compared. However, just before the application of insecticide, the integument of larvae must be treated with acetone. Two bioassays were used to monitor the resistance of codling moths collected in 2003-2005 in two apple (Malus spp.) orchards with different intensities of chemical control. Resistance ratios (RRs) to the tested insecticides were determined for both field populations of codling moth. For the population of codling moth from a commercial apple orchard in Velké Bílovice, cross-resistance to fenoxycarb, teflubenzuron, and phosalone was detected after the topical application of insecticides to fifth instars. The population of codling moth from Prague-Ruzyne was slightly resistant to phosalone and teflubenzuron. No resistance to diflubenzuron was detected in either tested population.     

Integrating chemical control with sterile insect releases in an integrated pest management programme for Thaumatotibia leucotreta

False codling moth Thaumatotibia leucotreta (Meyrick) (Lepidoptera: Tortricidae) is an important indigenous pest of citrus in southern Africa. Successful control is dependent upon integration of area‐wide sterile insect releases and other suppression methods. The aim of this work was to test pyrethroid and organophosphate‐based insecticides (tau‐fluvalinate and chlorpyrifos) for their residual effect on mortality of released irradiated T. leucotreta male moths. Both of these insecticides were effective in killing irradiated T. leucotreta for 7 days after application on leaves, after which degradation of the active ingredient resulted in a marked reduction in efficacy after 14 days and rendering them harmless. Mortality was found to be similar for irradiated and non‐irradiated male T. leucotreta after treatment. Consequently, even though these insecticides might have an effect on moths in the field, ratios of sterile:wild moths should not be altered. Supporting field data from six sites in the Sundays River Valley over a season of sterile insect releases showed the conventional chemical crop protection programme to be as effective as an integrated pest management programme in facilitating effective control of T. leucotreta through sterile insect releases. The study also confirmed that the ratios of sterile:wild male moths in the commercial citrus orchards were not affected by the application of insecticides. These findings confirm the high potential of sterile insect releases for control of T. leucotreta in citrus.     

Response of Cydia pomonella to selection on mobility: laboratory evaluation and field verification

1. The codling moth Cydia pomonella (L.), largely regarded as a sedentary species, shows great variation in flight capacity among individuals in the laboratory, as has been reported for tethered flight. The occurrence of individuals with the ability to fly over long distances is considered ecologically significant for the colonisation of new habitats in response to deteriorating environmental conditions. The work reported here was designed to investigate the importance of the genetic component in regulating dispersal in C. pomonella. 2. Bi‐directional selection on mobility measured by actograph was carried out in the laboratory. Both male and female codling moths responded significantly to the upward and downward selections. Divergence between the two selected strains occurred after one generation of selection and increased as further selections continued. The realised heritabilities for mobility were estimated as 0.43 in males and 0.29 in females when averaged from the two selection directions. 3. The dispersal capacity of the selected mobile and sedentary strains was verified by mark–release–recapture experiments in the field, though only the released male moths were captured effectively. Flight distances by males differed significantly between the two strains, with the mobile strain showing a greater dispersal capacity than the sedentary strain. 4. This study demonstrated a positive correlation between mobility measured by actograph and field dispersal capacity in the codling moth. Furthermore, it provides the first experimental data to show field performances of different genotypes in insect dispersal.     

Evaluation of dyes for marking painted apple moths (Teia anartoides Walker, Lep. Lymantriidae) used in a sterile insect release program

Abstract  Two methods of marking painted apple moth ( Teia anartoides Walker, Lepidoptera: Lymantriidae), internal and external dyes, were investigated for use with the sterile insect technique, to ensure that wild and released moths could be distinguished absolutely during the eradication program. Calco Red dye was used to mark all moths internally. It was added to artificial diet and was detectable in the adults. Dye was incorporated into the larval diet at four rates. No differences were observed in mortality of the larvae, fertility of the adult female or response of male moths to virgin females in a flight tunnel. Fluorescent powder dyes were used throughout the sterile insect program to identify irradiated moths, as well as their release location and colony. However, the powdered dyes reduced the ability of male moths to detect a component of the sex pheromone in electro-antennogram tests and to respond to a calling virgin female in flight tunnel trials. Field trials suggest that moths may subsequently overcome these negative effects and respond to female moths, as moths heavily coated in dye were recaptured.     

Factors influencing flight potential of Choristoneura conflictana

Abstract Factors affecting the flight potential of Choristoneura conflictana, an insect that undergoes cyclical changes in population density, are investigated using computer‐linked flight mills. Female and male moths are flown for a 12‐h period and the longest single flight and total flight distance of each moth is recorded. After flight bioassays, moths are subjected to lipid extraction with a soxhlet apparatus to determine the effect of body lipid content on flight. Larger C. conflictana moths fly farther than smaller moths. Female C. conflictana fly farther than male moths but the effects of both mating status and moth age on the distance flown are dependent on sex. Mated females fly farther than virgins and older females fly farther than young females but these relationships do not occur in males. Body lipid content affects the distance flown by both females and males through a significant interaction with sex and age. The factors examined in this study will vary with density in natural populations and are predicted to be important indicators of flight potential and dispersal in this species.     

Overwintering of codling moth (Lepidoptera: Tortricidae) larvae in apple harvest bins and subsequent moth emergence.

Codling moths, Cydia pomonella (L.), have long been suspected of emerging from stacks of harvest bins in the spring and causing damage to nearby apple and pear orchards. With increased use of mating disruption for codling moth control, outside sources of infestation have become more of a concern for growers using pheromone based mating disruption systems. Studies were designed to provide information on bins as a source of codling moth and the pattern of codling moth emergence from stacks of bins. In these studies, codling moth larvae colonized wood harvest bins at a much higher frequency than harvest bins made of injection molded plastic (189 moths emerged from wood compared with five from plastic). There was no statistical difference in the number of moths infesting bins that had been filled with infested fruit compared with bins left empty at harvest. This suggests that codling moth enter the bins during the time that the bins are in the orchard before harvest. Emergence of laboratory reared adult codling moth from wood bins placed in stacks was found to be prolonged compared with field populations. Temperature differences within the bin stacks accounted for this attenuated emergence pattern. Covering bin stacks with clear plastic accelerated codling moth development in the upper levels of the stack. Codling moth emergence patterns from plastic-covered stacks more closely coincided with male flight in field populations. This information could be important in developing a technique for neutralizing codling moth-infested bins, and in understanding how infested bins may influence pest management in fruit orchards that are located near bin piles. Implications for control of codling moth in conventional orchards and in those using mating disruption as the principal component of an integrated pest management system include increased numbers of treatments directed at areas affected by infested bins.     

苹果蠹蛾不育昆虫释放技术研究进展

不育昆虫释放技术(sterile insect technique,SIT)是一种环境友好、可作为大面积害虫综合治理(AW-IPM)的防治技术,是以压倒性比例释放不育昆虫来减少田间同种害虫繁殖量的害虫治理方法。苹果蠹蛾Cydia pomonella(L.)是世界重要的梨果类害虫,现已入侵世界5洲71国。本文综述了苹果蠹蛾大规模饲养技术、辐射不育技术、释放技术3个关键环节的研究与技术进展,主要包括:苹果蠹蛾人工饲料、实验种群建立、饲养设备与条件、收集和质量评估、长距离运输、辐射源与设备、辐射剂量与敏感性、释放方法、释放标记和释放量等,并介绍了各国采用SIT技术的应用效果。苹果蠹蛾在我国新疆、甘肃、宁夏、内蒙、黑龙江、吉林6个省区发现,对我国苹果产业安全生产构成严重威胁,我国很有必要引进并建立苹果蠹蛾SIT防治技术体系。     

Male oriental fruit moth response to a combined pheromone-based attracticide formulation targeting both oriental fruit moth and codling moth (Lepidoptera: Tortricidae)

Combined attracticide formulations targeting Oriental fruit moth, Grapholita molesta (Busck), and codling moth, Cydia pomonella (L.), were tested in a field trapping experiment. Capture of male codling moths in traps baited with the combined formulation was reduced compared with traps baited with the codling moth formulation alone, whereas capture of male Oriental fruit moth was increased compared with traps baited with the Oriental fruit moth formulation alone. Subsequent wind tunnel experiments showed that a single locus of the mixed attracticide formulation or close parallel presentation of the two formulations enhanced source contact by male Oriental fruit moths but did not influence earlier behaviors. However, the two formulations presented in a serial arrangement to Oriental fruit moth males in the wind tunnel resulted in enhanced lock-on, upwind flight, and source contact behaviors. In addition, male Oriental fruit moths remained on mixed pheromone droplets of the paste matrix longer than on droplets of the Oriental fruit moth formulation alone. The increased time spent on the mixed droplet was correlated with a more rapid poisoning and a greater proportion of poisoned males compared with males exposed to the Oriental fruit moth attracticide alone. These results demonstrate that a combined attracticide formulation will have different effects on each of the targeted species. It is anticipated that, due to decreased attractiveness, a combined formulation would be less effective against the codling moth. However, a mixed formulation, due to increased attractiveness and toxicity, could be more effective against the Oriental fruit moth under field conditions.     

Molecular technologies to improve the effectiveness of the sterile insect technique

The application of the Sterile Insect Technique (SIT) in area-wide integrated pest management (AW-IPM) programmes continues to increase. However, programme efficiency can still be considerably enhanced when certain components of the technology are improved, such as the development of improved strains for mass rearing and release. These include strains that (1) produce only male insects for sterilization and release and (2) carry easily identifiable markers to identify released sterile insects in the field. Using both classical and modern biotechnology techniques, key insect pests are targeted, where SIT programmes are being implemented. The pests include mosquitoes, the Mexican fruit fly, the codling moth, the oriental fruit fly and the pink bollworm. This special issue summarizes the results of research efforts aimed at the development and evaluation of new strains to a level where a decision can be made as to their suitability for use in large scale SIT programmes. Major beneficiaries will be operational AW-IPM programmes that apply the SIT against major insect pests.