Resistance monitoring in codling moth: a need for standardization

BACKGROUND: A wide range of methods has been used for monitoring resistance in Cydia pomonella L. However, the results from specific tests are not always concordant and they generate doubts over the extent to which the data represent the field situation. Therefore, the variability in the expression of resistance mechanisms and in resistance ratios between developmental stages and their reliability in a field situation were studied. RESULTS: Neonate larvae showed the highest insecticide resistance, except to spinosad. Resistance ratios were from 2 to 9.5, depending on the insecticide and the strain tested, when compared with the susceptible strain. Spinosad exhibited the highest resistance ratio for diapausing larvae (4.4, 12.2 and 4.4 for Raz, Rdfb and RΔ strains, respectively). Enzymatic ratios were higher in adults than in neonates or diapausing larvae, and the highest enzymatic ratios were observed for mixed‐function oxidases (7.3–36.7 for adults of resistant strains). CONCLUSIONS: The sensitivity of different instars depends on the insecticides used, and there is a differential expression of resistance mechanisms between stages. For the population considered in this study, tests on neonates provided the most consistent results to the field situation, but adult response to chlorpyrifos‐ethyl and azinphos‐methyl in the laboratory was also consistent with the field results. Copyright © 2008 Society of Chemical Industry     

Diversity of insecticide resistance mechanisms and spectrum in European populations of the codling moth, Cydia pomonella

Only a few of the registered insecticides against Cydia pomonella L. are still effective in areas where insecticide resistance has emerged in this pest. Resistance mechanisms are multiple, and their lone or cumulative effects in a single population are not completely understood. A detailed estimation of resistance spectrum is still required to define the suitable insecticides to use against a given population. The efficacy of ten insecticides was therefore investigated together with the resistance mechanisms expressed in four laboratory strains and 47 field populations of C. pomonella from five countries. Bioassays were performed using topical applications of diagnostic concentrations on diapausing larvae, and resistance mechanisms were analysed on adults emerging from control insects. All populations exhibited a reduced susceptibility to at least one insecticide when compared with the susceptible laboratory strain. Cross-resistances were observed between azinphos-methyl or phosalone and more recent compounds such as spinosad and thiacloprid. Resistances to azinphos-methyl, diflubenzuron, spinosad, tebufenozide and thiacloprid were significantly correlated with mixed-function oxidase activity, while increased glutathione-S-transferase and reduced non-specific esterase activities were correlated with resistance to azinphos-methyl and emamectin, respectively. Conversely, resistances to azinphos-methyl, tebufenozide and thiacloprid were negatively correlated with increased esterase activity. None of the observed mechanisms explained the loss of susceptibility of populations to chlorpyrifos-ethyl, and no significant correlation was detected between resistance to deltamethrin and the presence of the kdr mutation. The suitability of such non-target instars to monitor insecticide resistance in field populations is discussed.     

Resistance of codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), larvae in Michigan to insecticides with different modes of action and the impact on field residual activity

BACKGROUND: The codling moth is one of the principal pests of apple in the world. Resistance monitoring is crucial to the effective management of resistance in codling moth. Three populations of codling moth in neonate larvae were evaluated for resistance to seven insecticides via diet bioassays, and compared with a susceptible population. In addition, apple plots were treated with labeled field rate doses of four insecticides. Treated fruit were exposed to neonate larvae of two populations from commercial orchards. RESULTS: Two populations of codling moth expressed two‐ and fivefold resistance to azinphos‐methyl, seven‐ and eightfold resistance to phosmet, six‐ and tenfold resistance to lambda‐cyhalothrin, 14‐ and 16‐fold resistance to methoxyfenozide and sixfold resistance to indoxacarb, but no resistance to acetamiprid and spinosad. The impact of the resistance to azinphos‐methyl, measured as fruit damage, increased as the insecticide residues aged in the field. In contrast, fruit damage in methoxyfenozide‐ and lambda‐cyhalothrin‐treated fruit was observed earlier for resistant codling moth. No differences in efficacy were found for acetamiprid. CONCLUSIONS: Broad‐spectrum insecticide resistance was detected for codling moth. Resistance to azinphos‐methyl, lambda‐cyhalothrin and methoxyfenozide was associated with reduced residual activity in the field. Broad‐spectrum resistance presents serious problems for management of the codling moth in Michigan. Copyright © 2008 Society of Chemical Industry     

Effectiveness of twelve insecticides applied topically to diapausing larvae of the codling moth, Cydia pomonella L

Dose-mortality curves were established for 12 insecticides administered by topical application to diapausing larvae from a susceptible codling moth strain. Toxicity varied greatly among the insecticides tested. LC50 values ranged from 0.1 mg kg(-1) for fenoxycarb to over 2800 mg kg(-1) for diflubenzuron and indoxacarb. Discriminating dose levels were determined from dose-mortality reference curves for the detection of resistance in field-collected diapausing larvae.     

Modelling the interactions between phenology and insecticide resistance genes in the codling moth Cydia pomonella

In the codling moth Cydia pomonella (L), insecticide resistance genes have been associated with pleiotropic effects affecting phenology. In this paper, we investigated whether an increase in the frequency of insecticide resistance in field populations of C pomonella was likely to entail significant divergences in the temporal occurrence of both susceptible and insecticide-resistant individuals. For this purpose, we built a phenological model that provided suitable predictions of the distinct and diverging seasonal evolutions of populations of a susceptible and two insecticide-resistant (at two and three loci) homozygous genotypes of C pomonella. Model simulations for each genotype were further compared with pheromone trap catches recorded in a field insecticide-treated population over an 8-year period (from 1992 to 2000), which reflected the progressive annual increase in the frequency of resistance in southeastern France. We found a significant delay in field adult emergence relative to those predicted by the homozygous susceptible model, and the magnitude of such a delay was positively correlated with increasing frequencies of insecticide resistance in the sampled field population of C pomonella. Adult emergence predicted in the theoretical population that was homozygous for resistance at two loci converged with those recorded in the field during the investigated 8-year period. This suggested that the pleiotropic effects of resistance were likely to result in a significant phenological segregation of insecticide-resistant alleles in the field. The results of this study emphasized the potential for pest populations exposed to chemical selection to evolve qualitatively with respect to phenology. This may raise critical questions regarding the use of phenological modelling as a forecasting tool for appropriate resistance management strategies that would take into account the diverging seasonal evolutions of both insecticide resistance and susceptibility.     

Assessment of methoxyfenozide exposure on the sexual attractiveness and responsiveness of adult codling moth, Cydia pomonella L., in small orchard blocks

BACKGROUND: The sexual attractiveness of female and the responsiveness of male codling moth, Cydia pomonella L., exposed to surfaces treated with the ecdysteroid agonist methoxyfenozide were investigated in small orchard block experiments. The main purpose of the study was to determine whether the reported sublethal effects of methoxyfenozide affecting moth behavior also occur under field conditions. RESULTS: Data from the first experiment showed that in some trials untreated females were significantly more attractive to wild males than methoxyfenozide-treated females. The second experiment revealed that some released methoxyfenozide-treated males were not as responsive to calling females as untreated males, and that some untreated females were more attractive to males than methoxyfenozide-treated females. The third experiment revealed that wild males exposed to treated tree surfaces were significantly less responsive to traps than were wild males exposed to untreated trees. However, there were no significant differences in mean recaptures of released males between the untreated and treated orchard blocks. CONCLUSION: Exposure of adult C. pomonella to methoxyfenozide-treated surfaces resulted in a negative impact on male responsiveness to calling females and synthetic pheromone lures. However, the level of impact on the sexual attractiveness of treated females did not appear to be as explicit.     

Disruption of mating in codling moth (Lepidoptera: Tortricidae) by chlorantranilipole, an anthranilic diamide insecticide

The influence of the anthranilic diamide insecticide chlorantranilipole (DPX-E2Y45; Rynaxypyr against the adult stage of codling moth, Cydia pomonella L., was examined. Insecticide residues in plastic cups sprayed with 56 mg AI L(-1) (equivalent to the recommended field rate) had a minimal effect on adult survival or fecundity, but significantly fewer female moths were mated in treated than in untreated cups. Similar results were found in field studies with adults placed in screened cages on treated apple foliage for up to 3 weeks. Mating was disrupted when either sex was topically dosed with technical material (0.01-0.1 microg). Males were more sensitive to the disruptive effects of chlorantranilipole than females. A 2 h exposure significantly reduced male activity (walking and wing fanning), but not female calling. A 25 h exposure significantly reduced the activity of both sexes. The proportion of males landing near a sex pheromone source in a flight tunnel was significantly reduced following exposures of 2-4 h and completely eliminated following a 17 h exposure. Male moths partially recovered from either topical or residual exposure to chlorantranilipole, but the extent of their recovery was dose and time dependent. Male response to sex pheromone recovered within 2 h following a 4 h exposure, but following a 17 h exposure only a partial recovery occurred after 48 h. Mating by moth pairs exposed to chlorantranilipole residues for 24 h did not significantly increase during recovery periods of 24-96 h, except in cups treated with 10% of the standard concentration. The proportion of field-collected female codling moths caught in pear ester-baited traps that were mated was significantly lower in apple orchards treated season-long with chlorantranilipole compared with females caught in either untreated orchards or those treated with sex pheromones to cause mating disruption.     

Toxicity of azinphos-methyl to various development stages of the codling moth Cydia pomonella (Lepidoptera: Tortricidae)

The response to azinphos-methyl of different life-stages of the codling moth Cydia pomonella (L.) was studied. A similarity in response to azinphos-methyl (LC(50), LC(90)) was observed in neonate larvae obtained from the first and second generations of field populations. Mortality rates for neonate larvae of a field population cross-bred with a laboratory colony were lower (by a factor of 1.2-2.2) in comparison with field neonate larvae. The mortality rate of larvae from a laboratory colony exposed to artificial diet containing azinphos-methyl gradually decreased at older instars. The mortality rates of fifth-instar larvae were remarkably low when exposed to artificial diet mixed with azinphos-methyl or when topically treated with the insecticide. One- to three-day-old females were more sensitive than males of the same age, whereas the opposite was observed in 14-to 17-day-old adults. Mortality rates of 14- to 17-day-old adults were higher than those under 10 days old. No significant difference in sensitivity to the organophosphorus compound was noticed between the sexes of 7- to 10-day-old adults. Neonate larvae of the codling moth can serve as a target life-stage for various groups of pesticides, and the importance of using such a stage as a standardized methodology for monitoring resistance in the codling moth is discussed.     

Monitoring resistance to diflubenzuron and deltamethrin in French codling moth populations (Cydia pomonella)

French populations of codling moth have developed resistance to several insecticide classes. The susceptibility of susceptible and resistant laboratory strains to diflubenzuron and deltamethrin was evaluated using different exposure methods against various life stages. The tarsal contact method for adults was found to be an appropriate method for testing neurotoxic compounds such as deltamethrin. Insect growth inhibitors, like diflubenzuron, need to be analysed on juvenile instars. Monitoring methodologies were developed and evaluated (a) on neonates obtained by crossing field‐collected males with virgin females from the susceptible laboratory strain so as to overcome the problem of reduced fertility of wild females and (b) on diapausing larvae. All 36 populations analysed exhibited significant levels of resistance to both diflubenzuron and deltamethrin. Resistance to the two compounds was linked, regardless of the cultivation methods used in the monitored area. The monitoring methodologies will be implemented to evaluate the evolution of resistance according to the resistance management strategies that have been adopted. © 2000 Society of Chemical Industry     

苹果蠹蛾的发生现状、监测技术及防治方法研究进展

苹果蠹蛾(Cydia pomonella L.)是一种严重危害苹果、梨、桃、杏的检疫性害虫,目前在我国局部分布,严重威胁我国苹果主产区。本文综述了苹果蠹蛾的发生现状、化学生态学研究进展、常用的监测技术方法以及主要防治方法,对苹果蠹蛾综合防控存在的问题及改进方向进行了讨论,以期为减缓苹果蠹蛾的扩散传播提供理论基础。     

Efficacy of the pear ester as a monitoring tool for codling moth Cydia pomonella (Lepidoptera: Tortricidae) in New Zealand apple orchards

BACKGROUND: The behavioural response of both sexes of codling moth, Cydia pomonella to the pear-derived kairomone (ethyl (2E,4Z)-2,4-decadienoate), codling moth sex pheromone (E,E-8,10-dodecadien-1-ol), and sex pheromone combined with the pear derived kairomone loaded into red rubber septum were investigated in trapping experiments in New Zealand apple orchards. A range of 0.01-10.0 mg of pheromone loading in rubber septum dispensers was tested and the highest catch of males was in traps baited with 1.0 mg. No dose response in trap catch of males was seen in traps baited with different amounts of pear-derived kairomone (0.01-10.0 mg). RESULTS: The number of females caught was significantly affected by the amount of pear derived kairomone used to bait traps, with the highest catch obtained at 10 mg loading. The attractiveness of sex pheromone was not enhanced by the addition of the kairomone either when used in the same bait or in a separate bait. The mean number of males captured in traps was reduced by 44% when the pheromone and kairomone were combined at ratio of 1:1 (0.1 mg pheromone: 0.1 mg kairomone) in separate sources. CONCLUSION: Kairomone baited traps showed some potential for monitoring the flight activity of female C. pomonella in apple orchards in two locations (Canterbury and Hawke's Bay). However, the number of male moths caught was low as compared to the number of male moths caught in pheromone-baited traps, and therefore the sex pheromone should continue to be used for monitoring male activity.     

Plant volatile compounds shorten reaction time and enhance attraction of the codling moth (Cydia pomonella) to codlemone

BACKGROUND: The codling moth is the most serious pest of deciduous tree fruit (apples, pears, crabapples, walnuts, quince) worldwide. The high frequency of insecticide treatments per season has resulted in breakdown of codling moth control owing to insecticide resistance. As an alternative, integrated pest management includes mating disruption to achieve population suppression in orchards. Under this scheme, the sex pheromone of the codling moth, (E, E)‐8,10‐dodecadien‐1‐ol (codlemone), is released from dispensers in crops to hinder mating by luring males. Increasing the attractiveness of codlemone formulations to codling moth males can be regarded as a key to increasing the efficacy of mating disruption. With this aim, the effects of adding plant volatiles on the behavioural responses of codling moth males to codlemone were tested. RESULTS: Adding R(+)‐limonene, linalool, (E)‐β‐farnesene or ethyl (E, Z)‐2,4‐decadienoate to codlemone significantly increases the proportion of males flying to the pheromone source in a wind tunnel. The response level is equivalent to that of males responding to females releasing codlemone. Using real‐time recordings, it is shown how these four plant products also shorten the response time of males to codlemone under the behavioural criteria time to activation, time till upwind flight is induced and time to pheromone source contact. CONCLUSION: Shortening the response time and increasing source location by males of dispensers releasing codlemone with R(+)‐limonene, linalool, (E)‐β‐farnesene or ethyl (E, Z)‐2,4‐decadienoate added would enhance mating disruption through better engagement of males with dispensers, to the detriment of females. Copyright © 2011 Society of Chemical Industry     

Potential of Heterorhabditis zealandica (Rhabditida: Heterorhabditidae) for the control of codling moth,Cydia pomonella (Lepidoptera: Tortricidae) in semi-field trials under South African conditions

Infection of codling moth, Cydia pomonella (L.), larvae by the entomopathogenic nematode, Heterorhabditis zealandica, was studied under semi-field conditions. Late-instar diapausing codling moth larvae, coccooned in perforated cardboard strips, were used in all trials to evaluate the nematode treatments involved. A morning application of 0.5, 1.0 and 1.5 MM infective juveniles (IJs) per tree resulted in 85%, 95% and 100% codling moth larval infection, respectively. Contrasting results were obtained for an evening application, with low levels of codling moth infection (<50%). Nematode treatment strips placed in direct sunlight resulted in almost no infection (<10%), opposed to strips attaining 67% infection placed in the shade.     

Involvement of a sodium channel mutation in pyrethroid resistance in Cydia pomonella L, and development of a diagnostic test

Populations of the codling moth, Cydia pomonella L (Lepidoptera, Tortricidae) have developed resistance to several classes of insecticide such as benzoylureas, juvenile hormone analogues, ecdysone agonists and pyrethroids, but the corresponding resistance mechanisms have not been extensively studied. Knockdown resistance (kdr) to pyrethroid insecticides has been associated with point mutations in the para sodium channel gene in a great variety of insect pest species. We have studied two susceptible strains (S and Sv) and two resistant strains (Rt and Rv) of C pomonella that exhibited 4- and 80-fold resistance ratios to deltamethrin, respectively. The region of the voltage-dependent sodium channel gene which includes the position where kdr and super-kdr mutations have been found in Musca domestica L was amplified. The kdr mutation, a leucine-to-phenylalanine replacement at position 1014, was found only in the Rv strain. In contrast, the super-kdr mutation, a methionine-to-threonine replacement at position 918, was not detected in any C pomonella strain. These data allowed us to develop a PCR-based diagnostic test (PASA) to monitor the frequency of the kdr mutation in natural populations of C pomonella in order to define appropriate insecticide treatments in orchards.     

苹果蠹蛾对高效氯氟氰菊酯的抗性现状、机制及治理策略

苹果蠹蛾Cydia pomonella属鳞翅目卷蛾科,是全球仁果类水果种植地区最重要的果树害虫之一,也是我国重大农业入侵物种,对全球水果生产造成严重威胁。当前主要采用高效氯氟氯氰菊酯等杀虫剂对苹果蠹蛾进行防治,杀虫剂的频繁使用导致其对高效氯氟氰菊酯产生了抗性。针对苹果蠹蛾对高效氯氟氰菊酯的抗性问题,该文综述了全球苹果蠹蛾种群对高效氯氟氰菊酯的抗性现状,指出由编码细胞色素P450（cytochrome P450,P450）、羧酸酯酶（carboxylesterase,CarE）和谷胱甘肽S-转移酶（glutathione S-transferase,GST）的解毒酶基因过表达导致代谢能力增强是其最普遍的抗性机制,提出合理的抗性治理策略,并对苹果蠹蛾的绿色防控进行展望。     

Management of insecticide resistance in Oriental fruit moth (Grapholita molesta; Lepidoptera: Tortricidae) populations from Ontario

The development of resistance in the Oriental fruit moth, Grapholita molesta (Busck) to organophosphorus (OP) insecticides (azinphos-methyl and phosmet) is a serious threat to the tender fruit industry in Ontario (50% crop losses in 1994). Resistance to carbamate insecticides and increased survival of field-collected moths at diagnostic concentrations of pyrethroids were widespread. As a result, four different treatment regimes were tested to manage resistance in G molesta, and the changes in resistance frequencies under each treatment regime were monitored from 1996 to 1999. The data indicated that the levels of resistance were significantly influenced by the various treatment regimes. The seasonal pattern of resistance was similar for all treatment regimes, in that resistance peaked in mid-season and declined in the late season. Levels of resistance in G molesta to OPs decreased from 55% to 14% and that to pyrethroids declined from 30% to 10% from 1996 to 1999 under a treatment regime consisting of endosulfan-organophosphate-pyrethroid rotation. Similarly, under a treatment regime implemented in commercial orchards (organophosphate-pyrethroid rotation), resistance to OP insecticides declined from 50% to 12% and resistance to pyrethroids evolved to around 16%. The overall data indicated that resistance was unstable; a strategy based on rotation of insecticides by class for each generation of G molesta was successful in managing resistance to both OP and pyrethroid insecticides. The rotational strategy has been widely adopted by growers and is applied to ca 85% of the acreage.     

一种监测棉铃虫对Bt杀虫晶体蛋白抗性的技术

研究了以诊断试剂盒监测棉铃虫对苏云金杆菌(Bt)杀虫晶体蛋白(ICP)抗性的技术。将1日龄幼虫在含诊断剂量的饲料中(每毫升饲料含Bt ICP0.01mg)取食7天,以生长至3龄以上的个体作为抗性个体的判断指标;筛选出人工饲料中防腐剂剂量的最佳值为每350毫升饲料中含丙酸2.5ml。经对多个棉铃虫不同抗性种群的验证试验,证明诊断试剂盒检测抗性个体频率较为精确且应用简便。     

Azinphos-methyl and carbaryl resistance in adults of the codling moth (Cydia pomonella (L.), Lepidoptera: Tortricidae) from Northeastern Spain

The resistance of Cydia pomonella (L.) to organophosphates is widespread throughout the pome fruit growing areas. The lethal effects of two insecticides inhibitors of the acetylcholine esterase, azinphos-methyl and carbaryl, were evaluated in adults of five and four field populations of the codling moth, respectively. The lethal concentrations (LC₅₀ and LC₉₀) of these insecticides were determined in a susceptible strain from Spain (S_Spain). Topical bioassays using the approximate LC₉₀ values (3000 mg (a.i.)/L of carbaryl and 2000 mg (a.i.)/L of azinphos-methyl) that were obtained in S_Spain were tested as diagnostic concentrations. The enzymatic activities of mixed-function oxidases (MFO), glutathione S-transferases (GST) and esterases (EST) were measured to investigate their potential role in the detoxification of these insecticides.Carbaryl and azinphos-methyl caused ?53% and ?39% corrected mortality, respectively, in field populations, although the diagnostic concentrations applied were twofold and fourfold higher than the maximum concentration registered in Spain, respectively. The activities of MFO and GST were 7.3- to 16.1-fold higher and 2.5- to 3.7-fold higher in all the field populations compared to those in S_Spain, respectively.