Is lack of mating competitiveness in spring linked to mating asynchrony between wild and mass‐reared codling moths from an operational sterile insect programme?

Mating competitiveness and pheromone trap catches of mass‐reared, male codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), from the Osoyoos, British Columbia, Canada, mass‐rearing facility operated by the Okanagan‐Kootenay Sterile Insect Release Board, were compared to wild males using mark–release–recapture field experiments in spring, summer, and autumn at Summerland, British Columbia. In spring, significantly more wild diapause males mated with tethered, wild females than did non‐irradiated (0 Gy) or irradiated (100 or 250 Gy) non‐diapause, mass‐reared males. A lower dose of radiation did not improve mating competitiveness, nor catches of mass‐reared males released in spring. Median mating time for wild males was approximately 45 min earlier than mass‐reared males with most wild males (70.5%) mating before sunset and mass‐reared males mating at or shortly after sunset in spring. Superior mating competitiveness of wild males in spring was mirrored by greater recapture rates in pheromone‐baited traps. In summer, mating competitiveness of mass‐reared moths improved relative to wild males and there was a significant inverse relationship between radiation dose (0, 100, and 250 Gy) and competitiveness of mass‐reared males. In autumn, untreated, wild males were significantly more responsive to pheromone traps than non‐diapausing mass‐reared males receiving 250 Gy of radiation. Mass‐reared males, subjected to diapause‐inducing conditions as larvae and emerged from diapause before this irradiation treatment, were recaptured significantly more often than similarly irradiated, non‐diapause, mass‐reared males, but not more than untreated, diapause wild males. We hypothesize that differences between wild and mass‐reared males in daily timing or speed of responses to natural or synthetic pheromone sources under montane weather patterns typical of spring in British Columbia may partially explain poor activity of sterile males, and low sterile : wild overflooding ratios during spring when measured using pheromone traps by the sterile insect release programme in British Columbia.     

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。     

Influence of juvenile hormone and mating on oogenesis and oviposition in the codling moth,Cydia pomonella

Oogenesis in the codling moth, Cydia pomonella, and the role of juvenile hormones (JHs) were addressed. Rudimentary ovarian structures were recognisable in day 3–4 pupae, when haemolymph JH was still undetectable by coupled gas chromatography‐mass spectrometry in the selected ion mode (GC‐MS/SIM). The presence of developing oocytes was observed by light microscopy on day 8, coincident with very low JH titres (0.74 ± 0.05 ng/ml JH II). Chorionation was only evident upon emergence, following an increase in JH in the pharate adult (0h old: 4.71 ± 0.34 ng/ml JH II). Analysis of haemolymph from virgin and mated females indicated that JH II was predominant, with approximately equal and lower quantities of JHs I and III (3.3‐ to 5.0‐fold less). When pupae or newly emerged adults were treated with JH homologues, no alteration in ovarian protein content was apparent, but the JH mimetic, fenoxycarb, depressed the number of oocytes filling ≥ 50% follicular volume. Chorion deposition was stimulated by JHs I, II, or III (10 μg), but not by fenoxycarb (0.05 μg, 10 μg). Mating provided correct stimuli for enhanced choriogenesis and egg laying, and, since haemolymph JH titres were concomitantly elevated (approximately 2‐fold), it was postulated that the rise in JH elicited both these events. Application of JHs to virgin females, however, could not mimic mating; only increases in choriogenesis were induced: JH‐treatment of virgins (or mated insects) significantly decreased oviposition rates over 24 and 48 h and markedly reduced the life‐time total number of eggs. Arch. Insect Biochem. Physiol. 41:186–200, 1999. © 1999 Wiley‐Liss, Inc.     

转基因在苹果蠹蛾不育昆虫释放技术中的应用

使用不育昆虫释放技术是一项新兴的苹果蠹蛾Cydia pomonella(L.)防治方法,使用转基因得到不育雄蛾具有比传统辐射方法更多的优点。转基因技术通过使用基于piggyBac等转座子的质粒载体,并插入显性条件致死基因以培育遗传性别品系,同时插入荧光蛋白等标记基因来显示转基因的效果;在人工控制的发育条件,建立成熟的稳定苹果蠹蛾品系,用以最终的田间释放以达到防治的目的。     

Efficacy and mode of action of female‐equivalent dispensers of pheromone for mating disruption of codling moth

1 We evaluated the efficacy and mode of action of commercially available female‐equivalent dispensers of pheromone for mating disruption of codling moth Cydia pomonella (L.) (Lepidoptera Tortricidae). 2 Scentry fibres, but not Hercon flakes, were consistently as or more effective than Isomate C Plus dispensers when applied by hand at 50 dispensers per tree. 3 Individual Scentry fibres were slightly more attractive to male codling moth than Hercon flakes. 4 Efficacy of aerially applied Scentry fibres was equivalent to that of Isomate dispensers in disrupting male codling moth in 4‐ha commercial apple plots. 5 Initial deposition and retention of aerially applied fibres were inefficient with approximately 44% formulation loss at application, poor rainfastness and a gradual loss of dispensers from tree canopies after application. 6 Male codling moths were captured in traps baited with lures containing 0.1 mg of pheromone and mated with tethered virgin females that were surrounded by eight fibres placed 30 cm away or 16 fibres placed 45 cm away in untreated plots and plots treated with a background of 50 Isomate dispensers per hectare. 7 A plausible explanation for mating disruption of codling moth by female‐equivalent dispensers is competitive attraction without associated habituation and thus improving the effectiveness of these technologies will depend on maximizing the attractiveness of individual dispensers as well as the application density of dispensers per area of crop.     

Attraction of codling moth males to apple volatiles

The attraction of the codling moth, Cydia pomonella, to apple volatile compounds known to elicit an antennal response was tested both in the field and in a wind tunnel. In the field, (E)‐β‐farnesene captured male moths. The addition of other apple volatiles, including (E,E)‐α‐farnesene, linalool, or (E,E)‐farnesol to (E)‐β‐farnesene did not significantly augment trap catch. Few females were caught in traps which also caught male moths, but female captures were not significantly different from blank traps. In the wind tunnel, males were attracted to (E,E)‐farnesol, but not to (E)‐β‐farnesene. The addition of (E,E)‐α‐farnesene to (E)‐β‐farnesene had a synergistic effect on male attraction. The male behavioural sequence elicited by plant volatiles, including upwind flight behaviour, was indistinguishable from the behaviour elicited by sex pheromone.     

Juvenile hormone catabolism and oviposition in the codling moth, Cydia pomonella, as functions of age, mating status, and hormone treatment.

In vitro catabolism of juvenile hormone (JH) in haemolymph of adult female Cydia pomonella was ascribed mainly to juvenile hormone esterase (JHE) activity. No significant differences were noted between virgin and mated females 0-96 h post-emergence. Changes in JHE activity did not appear dependent upon fluctuations in JH titre; conversely, changes in JHE activity could not explain the changes in JH titres. Maximal JHE activity was recorded at 24 h (331.47 +/- 47.25 pmol/h/microl; 355.93 +/- 36.68 pmol/h/microl, virgin; mated insects, respectively) and preceded the peak in JH titres at 48 h. Topical application of JH II (10 ng-10 microg) or fenoxycarb (50 ng) enhanced JHE activity up to 640 and 56%, respectively. Treatment upon emergence with 10 microg JH II induced enzymic activity for less than 24 h, and when 10 microg JH II or 50 ng fenoxycarb were applied, circulating JH titres returned to control levels within 24 h. Oviposition was highly sensitive to exogenous JH and declined significantly with dosages >100 pg. To allow a degree of oocyte maturation before JH treatment, the hormone was administered at 6, 12, 24, or 48 h post-emergence and/or females were mated. Neither measure "protected" the system; oviposition declined immediately after JH application.     

Antennal and behavioural response of codling moth Cydia pomonella to plant volatiles

Identification of host volatile compounds attractive to codling moth Cydia pomonella, a most important insect of apple, will contribute to the development of safe control techniques. Synthetic apple volatiles in two doses were tested for antennal and behavioural activity in codling moth. Female antennae strongly responded to (Z)3‐hexenol, (Z)3‐hexenyl benzoate, (Z)3‐hexenyl hexanoate, (±)‐linalool and E,E‐α‐farnesene. Two other compounds eliciting a strong antennal response were the pear ester, ethyl (E,Z)‐2,4‐decadienoate, and its corresponding aldehyde, E,E‐2,4‐decadienal, which is a component of the larval defence secretion of the European apple sawfly. Attraction of codling moth to compounds eliciting a strong antennal response was tested in a wind tunnel. Male moths were best attracted to a blend of (E,E)‐α‐farnesene, (E)‐beta‐farnesene and ethyl (E,Z)‐2,4‐decadienoate. The aldehyde E,E‐2,4‐decadienal had an antagonistic effect when added to the above mixture.     

Development of a direct test of baculovirus resistance in wild codling moth populations

The occurrence of resistance of codling moth (CM, Cydia pomonella L.) to Cydia pomonella granulovirus (CpGV) used as biological control agent revealed the need for fast and reliable resistance monitoring methods. Here, we describe the development of a laboratory resistance test that is directly performed on larvae extracted from infested apples. This test is based on a 14‐day bioassay at a discriminative CpGV concentration of 2 × 10⁵ occlusion bodies/ml diet and can be applied to L1–L4 larvae. Information on virus resistance can be obtained within <4 weeks. In a survey, CM larvae were isolated and tested from 6698 apples from 10 different orchards in Germany, Austria, Switzerland, Italy and the Netherlands. We identified seven CM populations resistant or partly resistant to CpGV isolates. Although some of the orchards were treated with commercial CpGV products, this method allowed us to obtain reliable information about the resistance status of the examined populations.     

Flight tunnel response of codling moth Cydia pomonella to blends of codlemone,codlemone antagonists and pear ester

Upwind orientation flights of codling moth males Cydia pomonella L. to a single source of sex pheromone (E,E)‐8,10‐dodecadienol (codlemone) are significantly reduced when blending it with pheromone antagonists, either with codlemone acetate, (E,E)‐8,10‐dodecadienyl acetate, or with the codlemone isomer (E,Z)‐8,10‐dodecadienol. However, once activated by a pheromone stimulus, males no longer distinguish between a pheromone source and these antagonistic blend sources. This shows that the pheromone stimulus required for the initiation of an upwind flight response differs from the stimulus for maintaining upwind flight and landing at the source. In contrast to pheromone antagonists, males discriminate between pheromone alone and a blend source of pheromone and the plant volatile pear ester, ethyl (2E,4Z)‐2,4‐decadienoate. This indicates a difference in the detection and neural integration of pheromone and plant volatile stimuli.     

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.     

Effect of plant architecture and hail nets on temperature of codling moth habitats in apple orchards

Plant architecture of apple trees in commercial orchards was rapidly changed from traditional tall trees to dwarf trees to optimize yield and fruit quality. Additionally, hail nets are widely used to prevent yield loss by hail. These changes are expected to considerably influence the orchard microclimate and thus the developmental rates of pest insects in apple. However, these relationships have not yet been fully elucidated. The present study was conducted over the seasonal cycle to investigate the influence of plant architecture and hail nets on the habitat temperatures of the codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), in apple, Malus domestica Borkh. (Rosaceae). Within the canopies, leaf area index (LAI) and global site factor (GSF) were quantified using hemispherical photography. Temperature was analysed for the main habitats of the different codling moth stages, i.e., air within the canopy, bark of tree stems, and apple fruit. In dwarf trees, LAI was lower, leading to a higher GSF than in tall trees. Hail nets did not influence LAI and GSF. Results for dwarf trees compare as follows with those for tall trees: Average air temperatures within the canopy were 0.7 °C higher during daytime, whereas 0.4 °C lower at night. Mean surface temperatures of bark were 0.9 °C higher on sunny and 0.4 °C on overcast days. Mean surface temperatures of apple fruits were 1.8–2.7 °C higher on sunny days, but 0.6 °C cooler on overcast days. The effect of hail nets was confined to a reduction of the air temperature within the canopy by approximately 0.2–0.8 °C. Bark and apple surface temperatures were not significantly affected. Based on the temperature differences in the habitats considered, the calculated development of the codling moth in dwarf trees was on average 3 days faster than in tall trees. The calculations imply a negligible effect of hail nets on codling moth development.     

A test of fruit varieties on entry rate and development by neonate larvae of the codling moth,Cydia pomonella

The rate of entry by neonate larvae of the frugivorous codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), into fruit material was investigated. We used no‐choice bioassays in climate‐controlled rooms to assay larval entry across four host plant species (apple, pear, quince, walnut) and three varieties within a single fruit species (apple). Larvae successfully entering apples were reared to adulthood, and we collected tissue samples from apples which were successfully colonized in order to determine sucrose concentrations. This information was used to evaluate differences in adult moth size, development time, and pulp sucrose concentration due to apple variety. Four important findings emerged: (1) neonate larvae had the highest frequency of entry (86% of larvae) into apple fruits, compared with pear (78%), quince (56%), and walnut (32%); (2) the frequency of larval entry into immature apples differed across apple varieties, and larval entry rate was highest in variety Golden Delicious (72%), compared with Granny Smith (46%) and Red Delicious (64%); (3) on average, adult moths were larger and development times were shorter on the variety with the highest entry frequency (Golden Delicious); and (4) apple pulp sucrose concentrations were higher for Golden Delicious (17.5 μg mg^?1) than for either Granny Smith (15.9 μg mg^?1) or Red Delicious (15.1 μg mg^?1) varieties, which correlates positively with entry and development data. We conclude that host fruit species and varietals within a species affect the entry rate of neonate codling moth larvae in no‐choice assays. We hypothesize that larval development is influenced by mean sucrose concentrations or other phytochemical differences associated with host fruit varieties.     

The occurrence and distribution of resistance of codling moth to Cydia pomonella granulovirus in Europe

Thirty‐five codling moth (CM, Cydia pomonella L., Lepidoptera, Tortricidae) populations collected in different commercial orchards in six European countries were tested for their susceptibility to Cydia pomonella granulovirus (CpGV‐M). Including previously published data on CpGV‐M resistance, a total of 38 CM colonies showed considerably elevated LC₅₀ values, independent of the country origin. When only few test individuals are available, determination of mortality of neonate larvae at a discriminating concentration range of 10⁴ to 10⁶ OB/ml (>log4) as a direct measure of percentage susceptible individuals in a CM population is more advisable than calculation of LC₅₀ values. The >log4 mortality alone or in combination with the LC₅₀ value can be used for identification of resistance in a population. Results indicated a locally separated but widely spread occurrence of CM populations with low susceptibility to CpGV‐M. The most plausible hypothesis for the emergence of CpGV‐M resistance is its selection by repeated use of CpGV products.     

Mating disruption for control of damage by codling moth in Virginia apple orchards

Polyethylene dispensers (Shin Etsu) containing 172 ml of the sex pheromone, (E, E)-8,10-dodecadien-1-ol (63%), dodecenol (31%) and tetradecenol (6%), of codling moth (CM),Cydia pomonella (L.), were placed in apple orchards in Virginia. Two blocks of about 2 ha each were treated in 1989, and three in 1990. Dispensers were placed in trees at a density of 1000/ha shortly after apple bloom. Male orientation to pheromone traps was almost totally disrupted (a few males were captured at high population densities). In 1989, the Daleville pheromone-treated block had 0.9% and 0.8% CM-injured fruit in the center and edge, respectively; 0% and 39.5% injured fruit were found in the conventional control and abandoned blocks, respectively. The Criglersville orchard (‘organically’ managed, with high CM density) CM harvest injury was 16.0%, 16.5%, 34.5%, and 26.5% in the pheromone-treated center and edge, organic control and abandoned blocks, respectively. In 1990, the Daleville CM harvest injury was 4.7%, 7.3%, 1.1%, 0.3% and 58%, in the pheromone-treated center and edge, control center and edge, and abandoned blocks, respectively (possible reasons for the high injury in this block are discussed). Harvest injury in the Fincastle pheromone-treated and control blocks were 0.7% and 0%, respectively. The Criglersville orchard yielded 17%, 19% and 20% CM-injured fruit at harvest in the pheromone-treated, organic control and abandoned blocks, respectively. Pheromone release rate was calculated as 37 mg/ha/h in 1989.     

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.     

Parasitoid exploitation of the seasonal variation in host plant volatile emission for herbivore location

Volatile compounds from the apple, Malus domestica Borkh. (Rosaceae), change considerably as the season progresses, and this is successfully exploited by the female codling moth Cydia pomonella L. (Lepidoptera, Tortricidae), as it searches for oviposition sites. In this study, we investigated the effect of seasonal emissions of apple fruit volatiles on the host location behaviour of a parasitoid of the codling moth larvae, Hyssopus pallidus (Askew) (Hymenoptera: Eulophidae). In dual choice olfactory bioassays, the behaviour of the parasitoid in response to apple cues was observed over the complete 2003 growing season. Our results show that codling moth infested apples evoked a strong response from the parasitoid at the beginning of the season, until July. Then, attraction dropped drastically, increasing again at the beginning of August. At the end of the growing season, just prior to harvest, infested apples hardly evoked any behavioural response. Interestingly enough, mid‐season emissions of healthy apples were per se attractive to the parasitoid, and even preferred over volatiles from infested apples. Simultaneous volatile collections from healthy apples on twigs in the field were analysed throughout the season, showing that the overall quantity of headspace volatiles peaks at the beginning of June and mid‐August. The seasonal volatile emission is correlated with the behaviour of the parasitoid during the fruit ripening stage. The results are discussed in relation to the use of H. pallidus as a potential biocontrol agent, in order to enhance current integrated pest management (IPM) programs.     

Aedes aegypti lines for combined sterile insect technique and incompatible insect technique applications: the importance of host genomic background

Aedes aegypti L. (Diptera: Culicidae), being the primary vector of pathogenic arboviruses, is a target for the development of novel genetic approaches to complement current conventional vector control strategies such as the combined sterile insect and incompatible insect technique (SIT/IIT). A transinfected line of Ae. aegypti carrying the wAlbB Wolbachia strain (WB2) was introgressed into two genomic backgrounds, Brazil and Mexico, producing two new Ae. aegypti strains (WB2-BRA and WB2-MEX). These strains were evaluated with respect to several life-history traits such as fecundity, fertility, longevity, pupa size, pupation curve, and male mating competitiveness, as well as their response to irradiation. Our results show that the impact of Wolbachia infection depends on the genomic background and that the Brazilian one had no significant effect, whereas the Mexican one negatively affected fertility, longevity, and pupal size. Interestingly, Wolbachia-infected Ae. aegypti lines required a lower irradiation dose to achieve complete female sterility than the uninfected ones. The present findings are discussed given the potential use of Wolbachia-infected Ae. aegypti lines in combined SIT/IIT population suppression programs.     

Colonization of a hybrid strain to restore male Anastrepha ludens (Diptera: Tephritidae) mating competitiveness for sterile insect technique programs

To restore male mating competitiveness of Mexican fruit flies, Anastrepha ludens (Loew) (Diptera: Tephritidae), reared for sterile insect releases by the Mexican Fruit fly Eradication Campaign, two strain replacement techniques were evaluated. Field cage male competitiveness tests revealed that laboratory males of the Metapa strain mated 3 times less often with wild females than field-collected wild males. A strain developed from the cross of wild males and laboratory females (hybrid strain) was similar to a strain developed from the cross of laboratory males and females (laboratory strain) in that its females produced similar amounts of eggs and the eggs displayed similar levels of hatch and egg-to-pupa transformation in artificial diet. By contrast, a strain developed from the cross of wild males and females (wild strain), forced into artificial rearing, experienced a series of bottlenecks involving reduced egg laying and extremely poor development in diet. The male F1 progeny of the hybrid strain and field-collected wild males outcompeted Fl laboratory males in field cage tests for matings with field-collected wild females. In conclusion, we found that strains developed from the cross of wild males and laboratory females allowed us to restore male mating competitiveness of F1 Mexican fruit flies without compromising mass-rearing production.