苹果蠹蛾滞育特性研究进展

苹果蠹蛾是苹果、梨等仁果类果树上的重要害虫,也是重要的入侵害虫。其具有寄主广泛、适生性强、蛀果为害等特点,一旦侵入很难彻底根除。老熟幼虫脱果后寻找隐蔽场所滞育的习性是其适应不利环境和维持种群繁衍的重要生存策略。本文就苹果蠹蛾幼虫滞育的诱导和终止因素、滞育场所的选取及聚集习性、滞育幼虫对环境的适应能力及内在生理生化特性以及针对滞育幼虫的防治技术等研究进行了综述,为探明苹果蠹蛾的灾变机理、预测预报和防治提供依据。     

Control of Codling Moth,Cydia pomonella (Lepidoptera: Tortricidae), with Steinernema carpocapsae: Effects of Supplemental Wetting and Pupation Site on Infection Rate

Infection of cocooned codling moth (cydia pomonella) larvae by the entomopathogenic nematode Steinernema carpocapsae was studied in three field experiments. Factors that varied within or between experiments included method of application, type of substrate containing cocooned larvae, time when nematodes were applied, seasonal effects, and supplemental wetting before or after nematode application. Conventional air-blast sprayer applications of 0.5–5.0 million infective juveniles (IJs)/tree in fall resulted in ca. 30% mortality of larvae in cardboard trap bands, whereas hand-gun application (2 million IJs/tree) produced mortality of ca. 70%. Application in the evening caused higher larval mortality than application in the morning when no supplemental wetting was used after treatments. Morning and evening applications caused equivalent larval mortality when a postwetting treatment was included. In a trial conducted in midsummer, supplemental wetting, either before or after hand-gun application of 1 million IJs/tree, enhanced nematode-produced mortality. Mortality approached 100% if both pre- and postwetting was used. Larvae in exposed cocoons on apple wood were infected at a higher rate (86%) than those on wood in less exposed positions (73%) or in nonperforated cardboard (72%). Mortality rates for larvae in perforated cardboard were intermediate (77%). Application volumes used to deliver nematodes slightly enhanced infection rate of larvae in some substrates but not others. In one trial, parasitism of codling moth by the wasp Mastrus ridibundus (Ichneumonidae) was negatively correlated with nematode infection of codling moth larvae. Dissections showed that ca. 10% of larvae infected by nematodes had been attacked by the wasp.     

Trade-off between mobility and fitness in Cydia pomonella L. (Lepidoptera: Tortricidae)

Abstract.  1. The codling moth Cydia pomonella , one of the most important pest insects of apple fruit orchards worldwide, is regarded as a 'sedentary' insect, but the natural populations consist of both mobile and sedentary genotypes that display different dispersal capacity in the field. The current study investigated whether there is a fitness consequence of being mobile in this species by comparing two strains with contrasting levels of mobility obtained through bi-directional selection.
2. Female moths of the sedentary strain were significantly larger, laid more eggs during their lifespan, and both male and female moths of this strain lived longer, when compared with the mobile strain.
3. The age-specific fecundity of female moths from the sedentary strain was significantly greater, in particular between 3 and 5 days after emergence when the peaks of both oviposition and flight capacity occur, relative to the mobile moths, thereby leading to a greater reproductive function.
4. These differences resulted in different demographic parameters between the two strains. The sedentary strain had a significantly greater net reproductive rate ( R _o ) and a higher intrinsic rate of natural increase ( r _m ), although the generation time ( T ) was similar, hence requiring a shorter time to double the population size, in comparison to the mobile strain.
5. This study provides empirical evidence for the cost of being mobile in C. pomonella . Trade-offs between mobility and fitness might have contributed to the existence of genetic variance for dispersal behaviour in the natural populations and the maintenance of relative sedentariness in the species.     

Costs of insecticide resistance in Cydia pomonella (Lepidoptera: Tortricidae)

The obvious benefits associated with insecticide resistance for pest species may come at a cost to other life-history traits. In this study, we compared the larval and pupal developmental times, pupal mass wing surface area and wing fluctuating asymmetry (FA) in insecticide resistant and control strains of codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), collected from apple (Malus spp.) orchards in central Canada. Resistant strains had significantly longer larval developmental times and lower pupal masses compared with the susceptible strain. Although the forewings of resistant moths were smaller in resistant than control strain, no difference in wing FA was detected. Longer developmental times could increase exposure time to natural enemies, and reduced adult size could affect longevity and total reproductive output.     

Mating behaviors of Cydia pomonella (Lepidoptera: Tortricidae) as influenced by sex pheromone in electrostatic powder

Entostat is an electrostatically charged wax powder that can adhere strongly to insect cuticle, making it an ideal carrier to deliver pheromone for pheromone-based confusion techniques. We investigated the attractiveness of Cydia pomonella (L.) (Lepidoptera: Tortricidae) males treated with pheromone-laden Entostat powder to naive conspecifics as well as mating behaviors of males after such treatment in a laboratory flight tunnel. Male moths exposed to Entostat containing 1% E,E-8,10-dodecadien-1-ol (codlemone) acquired and retained the largest amount of the powder and became more attractive as point sources to naive moths compared with those treated with powder containing 5 or 10% codlemone. All Entostat-exposed males remained as attractive as a 0.1-mg codlemone lure for up to 24 h in flight tunnel investigations. Male moth orientation to normally attractive sources of codlemone was completely disrupted directly after treatment with Entostat powder. Males' ability to orient to 0.1-mg lures recovered progressively over a 6-d postexposure interval; however, their responses never reached the levels observed with untreated control moths. Entostat-exposed moths retained detectable amounts of codlemone up to 4 d. Our laboratory flight tunnel results suggest that the mode of action of Entostat powder as an autodissemination control tactic may be due to creating both attractive false point sources after exposure to the powder as well as directly inhibiting contaminated males' capability to orient to pheromone.     

Overwintering Strategy and Mechanisms of Cold Tolerance in the Codling Moth (Cydia pomonella)

Background

The codling moth (Cydia pomonella) is a major insect pest of apples worldwide. Fully grown last instar larvae overwinter in diapause state. Their overwintering strategies and physiological principles of cold tolerance have been insufficiently studied. No elaborate analysis of overwintering physiology is available for European populations.

Principal Findings

We observed that codling moth larvae of a Central European population prefer to overwinter in the microhabitat of litter layer near the base of trees. Reliance on extensive supercooling, or freeze-avoidance, appears as their major strategy for survival of the winter cold. The supercooling point decreases from approximately −15.3°C during summer to −26.3°C during winter. Seasonal extension of supercooling capacity is assisted by partial dehydration, increasing osmolality of body fluids, and the accumulation of a complex mixture of winter specific metabolites. Glycogen and glutamine reserves are depleted, while fructose, alanine and some other sugars, polyols and free amino acids are accumulated during winter. The concentrations of trehalose and proline remain high and relatively constant throughout the season, and may contribute to the stabilization of proteins and membranes at subzero temperatures. In addition to supercooling, overwintering larvae acquire considerable capacity to survive at subzero temperatures, down to −15°C, even in partially frozen state.

Conclusion

Our detailed laboratory analysis of cold tolerance, and whole-winter survival assays in semi-natural conditions, suggest that the average winter cold does not represent a major threat for codling moth populations. More than 83% of larvae survived over winter in the field and pupated in spring irrespective of the overwintering microhabitat (cold-exposed tree trunk or temperature-buffered litter layer).     

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.     

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.     

Genetic structure and diversity of a newly invasive species, the codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae) in China

Cydia pomonella (L.) was firstly reported in China in the 1950s and considered as one of the most serious invasive pest in fruit orchards of China. It spread rapidly from the original site in Xinjiang to other northwestern regions. The pest has further penetrated northeastern China since 2006. With its rapid invasion rate, most pome fruit production areas of China are being threatened. As yet there has been no research into the genetic diversity and structure of the codling moth population in China. We investigated the genetic variations of 12 C. pomonella populations collected from the main distribution regions (Xinjiang, Gansu and Heilongjiang Provinces) in China and compared them with one German and one Swiss population using eight microsatellites loci to infer the characteristics of genetic diversity and genetic structure. We observed sequential loss of genetic diversity and significant structuring associated with distribution but no significant correlation between genetic distance and geographic distance among northwestern populations. There was no genetic evidence for bottleneck effects in any of the populations. The results suggest that the loss of genetic diversity in C. pomonella populations resulted from the successive colonization of founder populations. Recent invasion history led to the lack of any bottleneck effect. The high level of population genetic structuring is related to the weak flight capacity of the codling moth and the human-aided dispersal rather than to geographic distance. These genetic data not only provide us with an understanding of the micro-evolutionary processes related to successful biological invasions, but also provide guidance for pest management strategies.     

Effects of chlorpyrifos on enzymatic systems of Cydia pomonella (Lepidoptera: Tortricidae) adults

The control program of codling moth (Cydia pomonella L.) in the Río Negro and Neuquén Valley is intended to neonate larvae. However, adults may be subjected to sublethal pesticide concentrations generating stress which might enhance both mutation rates and activity of the detoxification system. This study assessed the exposure effects of chlorpyrifos on target enzyme and, both detoxifying and antioxidant systems of surviving adults from both a laboratory susceptible strain (LSS) and a field population (FP). The results showed that the FP was as susceptible to chlorpyrifos as the LSS and, both exhibited a similar chlorpyrifos‐inhibitory concentration 50 (IC₅₀) of acetylcholinesterase (AChE). The FP displayed higher carboxylesterase (CarE) and 7‐ethoxycoumarine O‐deethylase (ECOD) activities than LSS. Both LSS and FP showed an increase on CarE activity after the exposure to low‐chlorpyrifos concentrations, followed by enzyme inhibition at higher concentrations. There were no significant differences neither in the activities of glutathione S‐transferases (GST), catalase (CAT) and superoxide dismutase (SOD) nor in the reduced glutathione (GSH) content between LSS and FP. Moreover, these enzymes were unaffected by chlorpyrifos. In conclusion, control adults from the FP exhibited higher CarE and ECOD activities than control adults from the LSS. AChE and CarE activities were the most affected by chlorpyrifos. Control strategies used for C. pomonella, such as rotations of insecticides with different modes of action, will probably delay the evolution of insecticide resistance in FPs from the study area.     

Uptake of Granulovirus from the Surface of Apples and Leaves by First Instar Larvae of the Codling Moth Cydia pomonella L. (Lepidoptera: Olethreutidae)

Cydia pomonella granulovirus (CpGV) has received considerable attention as a potential microbial insecticide for the control of the codling moth (Cydia pomonella L.) , a worldwide pest of apples. Laboratory experiments were established to investigate virus uptake by first instar larvae, using a novel leaf disc bioassay technique. Virus uptake was found to be independent of active feeding and larvae became infected simply by walking or browsing on sprayed leaf disc surfaces in as little time as 3.5 min. Infection increased as a function of time spent on the leaf disc surface and a linear log time/probit mortality relationship could be fitted. The bioassay technique used has potential for the realistic laboratory testing of virus spray formulations. A field experiment showed that virus infection could be contracted by newly hatched codling moth larvae both from the surface of sprayed leaves and sprayed fruit. The potential for exploiting this knowledge for improving spray formulations is discussed.     

Genetic analysis of Cydia pomonella (Lepidoptera: Tortricidae) populations with different levels of sensitivity towards the Cydia pomonella granulovirus (CpGV)

Microsatellite (simple sequence repeats, SSR) and mitochondrial DNA markers were used to assess the structure of European codling moth populations showing different levels of susceptibility towards one of the most important biocontrol agents used in apple production, the Cydia pomonella granulovirus CpGV-M. In 638 C. pomonella individuals from 33 different populations a total of 92 different alleles were scored using six SSR loci. The global estimate of genetic differentiation for all 33 populations was not significantly different from zero, thus indicating a lack of genetic differentiation. AMOVA analysis revealed a very weak but significant variance among C. pomonella populations from different geographic regions, however, no significant variation was evident between CpGV-M resistant or susceptible C. pomonella populations. Sequence analysis of a fragment of the cytochrome oxidase subunit 1 in eight C. pomonella populations resulted in 27 haplotypes, which were grouped in two distinct clusters. Again, no genetic differentiation between CpGV-M resistant and susceptible codling moth populations was detectable. In addition, Structure analysis using microsatellites and association tests with mtDNA haplotypes found neither population-level nor individual correlations associated with CpGV-M resistance. Accordingly, this lack of population structure does not allow discriminating between one or several, separate origins of CpGV-M resistance.     

Control of Cydia pomonella L. and Cydia molesta (Busck) (Lepidoptera Tortricidae) in pome-fruit orchards with Ecodian sex pheromone dispensers

A mating disruption approach using high densities of pheromone dispensers, has been recently proposed for codling moth, Cydia pomonella (L.), and oriental fruit moth, Cydia molesta (Busck.), (Lepidoptera Tortricidae), control. Ecodian Star dispensers, made of low-cost biodegradable material and easy to apply, were formulated with 10 mg of codlemone (E8,E10-12OH) and 10 mg of grapamone (Z8-12OH) and placed at a rate of 1,400-2,000 dispensers/ha. The pheromone release rates from new and field aged dispensers were evaluated by hexane extraction of the residual attractant (indirectly) and gas-chromatographic analysis. The release rate of field-aged dispensers decreased over time with a good linearity; they released a significant amount of synthetic sex pheromones over the entire season. Dispensers elicited close-range approaches of codling moth males in wind tunnel irrespective of their age. Field trials carried out from 2003 to 2004 confirmed the efficacy of Ecodian Star dispensers for codling moth and oriental fruit moth control, regardless the size of the treated area. Our results demonstrate that Ecodian dispensers achieved a good level of activity and longevity over the season. The potential of this strategy for the control of the moths is discussed.     

苹果蠹蛾不同虫态体征及雌雄个体的快速鉴别方法

【目的】通过对苹果蠹蛾Cydia pomonella (L.)不同虫态的形态特征进行识别描述和定量观测,比较分析并研究提出苹果蠹蛾老熟幼虫、蛹及成虫雌雄个体的快速、准确鉴别特征与方法。【方法】利用体式显微镜拍照、测量记录苹果蠹蛾不同虫态雌雄个体间的形态特征、体征参数等。【结果】苹果蠹蛾老熟幼虫雌雄个体的体征差异:雄性腹节背面有明显肾形斑,雌性无斑。蛹雌雄个体的体征差异:雄蛹腹面翅尖后有4条明显体节线,第9腹节腹面中央有一纵裂缝,裂缝两侧各有一半圆形的瘤状突起;而雌蛹翅尖后有3条明显体节线,第8腹节腹面前缘有一纵裂缝,与腹节线相连形成类似"Y"形,裂缝两侧平坦无突起。成虫雌雄个体的体征差异:雄虫前翅反面中室后缘有一黑褐色条斑,雌虫无此条斑;雄虫腹部狭长,抱器瓣常开张,呈钳状,雌虫腹部圆筒状,末端较细。【结论】研究结果揭示了苹果蠹蛾老熟幼虫、蛹及成虫等不同虫态雌雄个体间的体征差异,比较研究并提出了便于快速鉴别雌雄个体的典型特征和识别方法。     

The influence of humidity on the effect of Steinernema feltiae against diapausing codling moth larvae (Cydia pomonella L.) (Lepidoptera: Tortricidae)

Codling moth (CM) is a serious and global pest of pome fruit. It overwinters in cryptic habitats as cocooned diapausing larvae. Field trials with the entomopathogenic nematode Steinernema feltiae (Rhabditida: Steinernematidae) report control of diapausing CM of up to 70%, but results are variable. The objective of this study was to define environmental conditions favouring the performance of the nematodes. Cocooned larvae were more susceptible than non-cocooned larvae. S. feltiae was unable to infect CM at a water activity (aw-values) < or = 0.9. Mortality of cocooned larvae was reported at lower aw-values than of non-cocooned larvae. Exposure time and impact of external relative humidity (RH) was studied. Mortality of cocooned larvae did not further increase after half an hour of exposure, whereas the mortality increased with increasing exposure time in non-cocooned larvae. LC50 and LC90 considerably decreased with increasing RH. The influence of the relative humidity was less pronounced when surpassing 80% than the effect of the volume of applied water. When S. feltiae was formulated in a surfactant-polymer-formulation (SPF), mortality significantly increased when compared to application in water.     

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.     

Attract and kill for codling moth Cydia pomonella (Linnaeus) (Lepidoptera: Tortricidae) control in Syria

The possibility of controlling the codling moth Cydia pomonella (Linnaeus) using an attract and kill approach as an alternative to chemical sprays with contact insecticides was investigated in widely separated orchards. The results of a 4‐year study have shown that, using an attract and kill approach, three applications/season kept infestation rates in treated orchards below the economic injury level except in one with a too high codling moth population density. The mean number of male codling moths/trap/week in attract and kill‐treated orchards was much lower in comparison with control orchards which were treated with the usual cover sprays of insecticides. The results also showed that the efficacy of attract and kill under orchard conditions decreased with time and the relationship between time effect and codling moth death rate was very strong. These data indicate that the attract and kill technique applied at a rate of three application per season resulted in good control of codling moth in well managed orchards in Syria.