Pleiotropy of insecticide resistance in the codling moth, Cydia pomonella

The codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), has developed resistance to various insecticides. Relative fitness of one susceptible strain (Sv) and two strains selected for resistance to diflubenzuron (Rt) and deltamethrin (Rv), respectively, was measured in the absence of insecticide selection pressure. Mating rate, fecundity, fertility, developmental time, fifth instar weight, and adult longevity were compared. Both resistant strains were less fecund and fertile, developed more slowly, weighed less, and had shorter life-spans than the susceptible strain. These results indicate that biological constraints are associated with insecticide resistance in the codling moth. We also found that fitness estimates of the Rv strain did not differ statistically from those of the Rt strain. Enhanced mixed-function oxidase and glutathione-S-transferase activities have been shown to be involved in insecticide resistance in both Rt and Rv strains. This suggests that the fitness cost described in both resistant strains was mainly associated to metabolic resistance. The impact of such deleterious pleiotropy of insecticide resistance in C. pomonella in terms of resistance management in the field is discussed.     

Diapause induction in the codling moth, Cydia pomonella: effect of prediapause temperatures

The effect of four prediapause temperatures (18, 22, 26 and 30°C) on the photoperiodic response of the codling moth, Cydia pomonella (L.), was studied under controlled conditions. The highest rates of diapause were recorded, for all day-lengths, at temperatures of 22 and 26°C while relatively lower rates of diapause were elicited at 18 and 30°C. The same trend was demonstrated by projecting the values of the critical photoperiod which induces 50% diapause (=CPhP₅₀) over the prediapause temperature. The change in diapause incidence as a function of photoperiod, at all prediapause temperatures, exhibited a response characteristic of long-day insects, i.e. high rates of diapause at short days (12–13.5 h) and a decrease in diapause incidence at long days (14–15 h). The results for temperatures 22, 26 and 30°C support the view that lower prediapause temperatures enhance diapause induction, at a give photoperiod, while higher temperatures tend to avert or diminish the process. On the other hand, the low rates of diapause obtained at 18°C contradict this view. Nevertheless, high correlation was found between the laboratory evidence and field data, indicating the adaptability of the Israeli codling moth to subtropical climate.     

Diapause induction in the codling moth, Cydia pomonella: effect of larval diet

The effect of larval diet on diapause induction in the Israeli strain of the codling moth, Cydia pomonella (L.), was studied in a field trial using intact apple fruits of two varieties: Ana (early-ripening, in the end of June) and Granny Smith (late-ripening, in October). Diapause incidence increased as fruit age (determined as days from fruit-set) progressed. These results corroborate former studies on other strains of the codling moth, where excised fruits were used.The combination of 80-day-old, fully ripe, Ana fruit treatment with the longest days of the year, yielded 38% diapause. This result demonstrates that mature fruit (inducing diapause) cannot completely override the effect of long day (averting diapause), but does confirm that larval diet modifies the photoperiodic induction of diapause in the codling moth.Deceased, October 1988     

Field resistance of codling moth against Cydia pomonella granulovirus (CpGV) is autosomal and incompletely dominant inherited

The current appearance of local codling moth populations with resistance to Cydia pomonella granulovirus (CpGV) is an impediment to continuous CpGV application. Therefore, crossing experiments have been performed in order to gain information about the inheritance of resistance. Evidence is presented that the observed field resistance is stably inherited even under non-selective conditions in the laboratory. Offspring of reciprocal F(1) crosses between a susceptible ('S') and a resistant ('R') strain and backcrosses between F(1) and S were bioassayed at different CpGV concentrations. The resistant strain showed 100 times lower susceptibility in 7-day bioassays. The responses of the reciprocal crosses (male S x female R and female S x male R) did not differ significantly, indicating that resistance is autosomally inherited. The median lethal concentration for the F(1) progeny was intermediate between those of its parental strains. Mortality data obtained from the backcrosses suggested that inheritance of resistance is due to a non-additive, polygenic trait.     

Selection of suitable trichogramma strains to control the codling mothCydia pomonella and the two summer fruit tortrix mothsAdoxophyes orana,Pandemis heparana [Lep.: Tortricidae]

Simple laboratory methods to select candidateTrichogramma strains for use in biological control were tried. 17 strains were screened for their suitability against the codling mothCydia pomonella L. as well as the 2 summer fruit tortrix mothsAdoxophyes orana F.R. andPandemis heparana Schiff. In one set of experiments, the capacity ofTrichogramma to parasitize each of these target pests was examined, in another set, the preference ofTrichogramma to contact and parasitize the target pests compared to the standard mass rearing hostSitotroga cerealella was assessed in choice experiments. A strain ofTrichogramma dendrolimi from the People's Republic of China was found to have the highest fecundity with all the 4 hosts tested. In choice experiments, however, this strain was shown to have a near equal preference between the target tortrix pestsC. pomonella, A. orana and the replacement hostS. cerealella. 11 localTrichogramma strains collected from fruit orchards and vineyards in different locations in the Federal Republic of Germany were reared separately. Four of these local strains, one identified asT. embryophagum, showed clear preference to the tortrix pestsC. pomonella andA. orana compared toS. cerealella. The preference of one of these strains was particularly strong and amounted to a near total rejection ofS. cerealella eggs in the presence of any of the 2 tortrix pests. The remaining 12 strains had weaker parasitization capacity and/or less preference to the target pests. Four of the strains tested in these experiments, includingT. dendrolimi andT. embryophagum, were mass reared and released in apple orchards between 1984 and 1986. The results of these field experiments are planned to be published in this journal. Mit Unterstützung der Deutschen Forschungsgemeinschaft.     

New method for testing solar sensitivity of commercial formulations of the granulovirus of codling moth (Cydia pomonella, Tortricidae: Lepidoptera)

A method for screening codling moth granulovirus (CpGV) formulation sensitivity to sunlight using specially prepared half apples and a solar simulator is described. The half apple preparation allows an even coverage of virus over the surface of the fruit that would not be possible using whole apples. Leaves and artificial medium were not usable for extended periods of exposure in the solar simulator due to excess drying. Fruit was sprayed with 10⁻³ and 10⁻⁵ dilutions of three commercial formulations of CpGV (Carpovirusine, Cyd-X, and Virosoft) and infested with codling moth neonates. Half of the sprayed fruit was exposed to 650 W/m² for 4 h in an Atlas Suntest CPS solar simulator resulting in an accumulated radiant energy of 9.36 × 10⁶ J/m² before they were infested with neonate codling moth larvae. Spraying non-irradiated fruit with the 10⁻³ dilution of Cyd-X and Virosoft resulted in nearly 100% mortality of neonate larvae. Irradiation reduced viral activity by 71-98% at the 10⁻³ dilution and by up to 32% at the 10⁻⁵ dilution relative to non-irradiated fruit. The procedures utilized enabled good preservation of the fruit throughout the incubation period and minimized invasion of the fruit by plant pathogens and saprophytic organisms. This laboratory method for screening candidate formulations and potential UV protectants could conserve time and resources by eliminating adjuvants with less potential in laboratory tests and field testing only the most promising candidates. It also enables year-round testing.     

Host preferences ofTrichogrammatoidea bactrae fumata (Hym.: Trichogrammatidae) an egg parasitoid of leafrollers (Lep.: Tortricidae)

Trichogrammatoidea bactrae fumata Nagaraja is a naturally occurring egg parasitoid of some leafrollers in New Zealand kiwifruit orchards. The parasitoid showed a distinct preference for younger host-eggs ofEpiphyas postvittana, in which it achieved a higher rate of successful development. Such a preference is not as apparent withCtenopseustis obliquana. Host-species preference experiments found that the parasitoid rarely attackedPlanotortrix octo, and successful wasp emergence from parasitised eggs of this host was low.Epiphyas postvittana was the preferred host whenT. bactrae fumata was reared on this species. When reared onC. obliquana, the parasitoid showed no preference forE. postvittana orC. obliquana.     

Incidence and transmission of a granulovirus in a large codling moth [Cydia pomonella L. (Lepidoptera: Tortricidae)] rearing facility

Incidences of potential per os Cydia pomonella granulovirus (CpGV) transmission within a large codling moth colony were identified. CpGV was detected in the water which is used to wash egg sheets. When pre-neonates were extracted from eggs prior to emergence and tested for the presence of CpGV, 40% were found to carry amounts of CpGV detectable by a polymerase chain reaction (PCR) assay, suggesting possible transovarial transmission of the virus. Although symptoms typical of virus infection and larval death were found infrequently within communal rearing trays, the frequency with which CpGV DNA was detected by PCR assays increased from a mean of 31% of 10-day-old larvae to 94% of 25-day-old larvae. CpGV in codling moth cadavers remained virulent after being held at 60 degrees C for 3 days under conditions similar to the treatment of spent diet at the rearing facility before its disposal. PCR tests of surface samples taken from air filters and rearing rooms of the rearing facility were found to contain CpGV. Bioassays of surface samples from the diet trash bin and a filter through which outside air is passed before entering the rearing chambers resulted in significant codling moth neonate mortality. The virulence of CpGV in dust from the spent diet and the original inadvertent positioning of the diet trash bin directly below one of the air intake ducts are suggested as a possible additional source of CpGV contamination within the facility.     

Orchard trials in Canada on control ofCydia pomonella (Lep: Tortricidae) by granulosis virus

Cydia pomonella (L.), a major pest of apples in Canadian orchards, is susceptible to a granulosis virus (CpGV). Orchard trials with different formulations of CpGV in Ontario, whereC. pomonella has two generations per year, indicated that as many as seven applications of virus would be needed to maintain the percentage of fruit with deep larval entries below 4%. In Nova Scotia, where there is only one generation ofC. pomonella per year, two applications of CpGV usually were needed although, in some cases, one sufficed. The protection of fruit conferred by CpGV equalled that by the organophosphate insecticides, azinphosmethyl or phosmet, in some tests. The advantages of a selective treatment, such as CpGV, over use of broad-spectrum insecticides are discussed.

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Résumé Cydia pomonella (L.), un ravageur important du pommier dans les vergers canadiens, est sensible au virus de la granulose. Des essais en vergers utilisant deux préparations de ce virus en Ontario, oùC. pomonella présente deux generations par année, indiquent qu'il faudrait jusqu'à sept pulvérisations pour maintenir à moins de 40% la proportion de fruits portant des infestations larvaires. En Nouvelle Ecosse, oùC. pomonella ne compte qu'une seule génération par an, deux pulvérisations furent nécessaires, quoique dans certains cas une seule a suffi. Dans certains essais, le niveau de protection des fruits égalait celui fourni par les esters phosphoriques, l'azinphos-méthyle ou le phosmet. On discute des avantages des traitements sélectifs tels que le virus de la granulose en comparaison avec l'emploi des insecticides de grande polyvalence.

     

Isolation of microsatellite loci in the codling moth,Cydia pomonella (Lepidoptera: Tortricidae)

Cydia pomonella L. is an important insect pest of pome fruits worldwide. We have isolated and characterized 17 microsatellite loci from the enriched genomic libraries constructed using a biotin/streptavidin capture protocol. Among these loci, 11 were scored polymorphic when 50 individuals from a laboratory population were screened. Their alleles numbered two to four, with the observed heterozygosity (H_O) ranging from 0.114 to 0.404. Successful amplification was obtained for all these loci when the designed primers were tested, showing the promise of use in genetic mapping and population studies.     

Parasitization of potato tuber moth eggs (Lep., Gelechiidae) from irradiated adults by Trichogramma (Hym., Trichogrammatidae) and control of moth population with combined releases of sterile insect and egg parasitoid

Abstract:  Eggs of the potato tuber moth (PTM) Phthorimaea operculella (Zeller) obtained from 150 or 300 Gy-irradiated moths, either inbred or out-crossed to untreated counterparts, were offered as host material to three generalist Trichogramma species; Trichogramma cacoeciae , Trichogramma evanescens , and Trichogramma principium , to determine the ability of these egg parasitoids to develop in F₁ sterile eggs. All eggs laid by the different parental crosses at 150 or 300 Gy, were acceptable and suitable for Trichogramma development. However, in no choice and paired comparison tests, eggs from 300 Gy-irradiated parents were significantly less preferred compared with eggs laid by non-irradiated moths. No intra-specific differences for PTM sterile eggs were recorded among the tested Trichogramma species. The mean number of wasps emerging from each egg types was not affected by an interaction between parental crosses and applied doses. Overall, this study demonstrates that release of irradiated and non-irradiated moths at a 5 : 1 over-flooding ratio and Trichogramma over potatoes in small plexiglass boxes, is complementary and more effective in reducing PTM F₁-emerged progeny than either technique used alone. The results provide baseline data for developing an integrated control approach by using inherited sterility technique in conjunction with Trichogramma for PTM population suppression.     

Attract and kill: a new method for control of the codling moth Cydia pomonella

The attract and kill technique has been formulated in a product under the trade name `Sirene CM®'. It consists of a viscous paste containing 0.16% codlemone to attract the male moths and 6.0% permethrin to kill them. The formulation is applied by hand twice per season using a specially developed system which can be calibrated for application of the paste to the host plant in small droplets of either 100 l or 50 l. Between 1995 and 1997, 15 trials on control of the codling moth were conducted in isolated orchards in the Lake Geneva region. In each plot, depending on tree size, two applications varying between 52 and 537 g ha^–1 of Sirene CM were made. In 14 trials, the larval attack of the codling moth on fruit was below the economic threshold of 1% and the hibernating population stayed at a low level. One single plot (0.4 ha) had to be treated with a curative spray in 1995, because the initial population was much too high. According to the reductions in trap catch and of mating frequency measured by tethered codling moth females, efficiency of the attract and kill droplets lasted 5–7 weeks, after which it decreased slowly.     

Overcoming the resistance of codling moth against conventional Cydia pomonella granulovirus (CpGV-M) by a new isolate CpGV-I12

Recently, codling moth (CM, Cydia pomonella L.) populations with a significantly reduced susceptibility to C. pomonella granulovirus (CpGV) products have been observed in Germany. A novel CpGV isolate, designated CpGV-I12, is able to overcome the CpGV resistance. CpGV-I12 originated from Iran and showed superior efficacy in laboratory bioassays against a resistant CM strain (CpR), which has a 100-fold reduced susceptibility to commercially used isolate CpGV-M. Determination of the median lethal concentration (LC₅₀) indicated that CpGV-I12 is nearly as efficient in resistant CpR as CpGV-M in a susceptible CM strain (CpS). Beyond, CpGV-I12 caused superior mortality in CpS. Infection experiments showed that the resistance breaking effect can be observed in all instars of CpR. CpGV-I12 is a promising alternative control agent of CM in orchards where conventional CpGV products fail. In addition, we demonstrate in bioassays with recombinant expressed Cry1Ab that cross-resistance to CpGV and Bacillus thuringiensis products is not likely.     

Activity of Bacillus thuringiensis delta-endotoxins against codling moth (Cydia pomonella L.) larvae

Solubilized protoxins of nine Cry1 and one hybrid Cry1 delta-endotoxin from Bacillus thuringiensis were tested for their activity against larvae of the codling moth (Cydia pomonella L). Cry1Da was the most toxic, followed by Cry1Ab, Cry1Ba, and Cry1Ac, while Cry1Aa, Cry1Fa, Cry1Ia, and SN19 were still less active. Cry1Ca and Cry1Cb showed no activity. In vitro trypsin activation increased activity of all eight active delta-endotoxins, and dramatically enhanced toxicity of hybrid SN19, Cry1Aa, Cry1Ac, and Cry1Fa. The differences between toxicity of proteins before and after trypsin digestion suggests that proteolytic activation in the C. pomonella digestive tract plays a critical role for the activity of Cry proteins against this insect.     

Evaluation of spray-dried lignin-based formulations and adjuvants as solar protectants for the granulovirus of the codling moth, Cydia pomonella (L)

Commercial formulations of the codling moth, Cydia pomonella L., granulovirus (CpGV) are limited by their short residual activity under orchard conditions in the Pacific Northwest. We evaluated spray-dried lignin-encapsulated formulations of CpGV for improved solar stability based on laboratory bioassays with a solar simulator and in field tests in an infested apple orchard. In laboratory tests, aqueous lignin formulations containing a high dosage of 3 x 10(10) occlusion bodies (OB)/L, with and without the additives titanium dioxide (TiO(2)) and sugar, provided significant solar protection of virus, i.e., mortality of codling moth exposed to lignin formulations that had been irradiated with 9.36 x 10(6) joules/m(2) was 92-94%, compared with 66-67% from a glycerin-stabilized product (Cyd-X) or suspension of pure unformulated virus at the same rates. By comparison, a lower dosage of the lignin formulation (3 x 10(8)OB/L) did not provide significant solar protection. Equivalent dosage-dependent patterns in solar protection were observed in further tests with the lignin formulation, when an intermediate (3 x 10(9)OB/L) as well as the low dosage provided no solar protection. Equivalent rates of a blank lignin formulation (containing no virus) did not affect larval mortality, suggesting a protective effect of the lignin on the virus at the high rate. The use of several spray adjuvants, 'NuFilm-17' and 'Organic Biolink' (sticker-spreaders at 0.06% v/v), 'Raynox' (sunburn protectant at 5% v/v), and 'Trilogy'(neem oil at 1% v/v) did not provide solar protection of a commercial CpGV preparation in laboratory tests. In season long orchard tests (Golden Delicious), the lignin formulation of CpGV applied at 6.57 x 10(12)OB/ha did not significantly improve control of codling moth or protection of fruit compared with Cyd-X at equivalent rates. Our studies show that lignin-based CpGV formulations provided solar protection at relatively high virus dosages. The testing of lignin formulations containing reduced virus concentrations may allow virus solar protection to be achieved at more economical rates.     

Efficacy of entomopathogenic nematodes (Rhabditida: Heterorhabditidae and Steinernematidae) against codling moth,Cydia pomonella (Lepidoptera: Tortricidae) in temperate regions

The biocontrol potential of South African isolates of Heterorhabditis zealandica, Steinernema citrae, S. khoisanae, S. yirgalemense, and Steinernema sp., was evaluated against codling moth, Cydia pomonella. Codling moth was susceptible to all six nematode isolates at a concentration of 50 infective juveniles/insect (78–100% mortality). Low temperatures (10 h at 17°C; 14 h at 12°C) negatively affected larvicidal activity (≤3%) for all isolates. All tested isolates were most effective at higher levels of water activity (a _w=1). The average a _w50-values for all isolates tested was 0.94 (0.93–0.95), except S. khoisanae 0.97 (0.97–0.98). Regarding host-seeking ability, no positive attraction to host cues could be detected amongst isolates, except for H. zealandica. Three of the isolates, H. zealandica, S. khoisanae, and the undescribed Steinernema sp., were selected for field-testing and proven to be effective (mortality >50%). Insect containment methods used during field experimentation was shown to influence larvacidal activity, as different levels of mortality were obtained using various containment methods (wooden planks vs. pear tree logs vs. mesh cages). Pear tree logs were impractical. Predictive equations were subsequently developed, enabling future trials to be conducted using either planks or cages, enabling the prediction of the expected level of control on tree logs. All tested isolates therefore showed a certain degree of biological control potential, however, none of the experiments showed clear efficacy-differences amongst isolates. The study highlighted the importance of environmental factors to ensure the successful application of these nematodes for the control of diapausing codling moth larvae in temperate regions.     

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.     

苹果蠹蛾性信息素的研究和应用进展

苹果蠹蛾Cydia pomonella (L.)是我国重要的检疫害虫, 在我国仅分布于新疆和甘肃以西局部地区, 但一直保持向我国东部扩张的趋势。在国际上, 利用性信息素监测和迷向防治苹果蠹蛾已经成为一种切实可行并广泛应用的害虫管理技术。本文综述了苹果蠹蛾性信息素的成分鉴定、人工合成和应用情况的研究进展, 指出了目前存在的问题和应用前景, 以期为我国苹果蠹蛾的防控策略的研究和应用提供参考。