Behaviour of flying oriental fruit moth males during approach to sex pheromone sources

Abstract. The pheromone-modulated upwind flight tracks of Grapholita molesta (Busck) males were video recorded as they approached a point-source of pheromone in a wind tunnel. The field of view of the video recording was divided longitudinally into 33 cm sections and the flight behaviour of the males in these sections was measured and compared as they approached from 233 cm to 50 cm downwind of the pheromone source. As the males approached the source, their mean ground speeds decreased. The mean values of their track angles increased with respect to due upwind (0), indicating movement more across the wind. These changes resulted mainly from the males decreasing their air speeds as they progressed up the plume toward the source. They did not change the average direction of their steering (course angle). Thus, the increase in track angles resulted from the males allowing themselves to drift more in the wind as they approached the odour source. The males also increased their average rate of counterturning as they approached the source. The net result of all these behavioural changes was a track that slowed and grew narrower, giving the impression that the males were 'homing-in' on the pheromone source as they approached. Causes of these systematic changes in behaviour are considered with respect to the known systematic changes in pheromone plume structure as the distance to the source decreases.     

Optomotor anemotaxis polarizes self-steered zigzagging in flying moths

ABSTRACT. Experiments with oriental fruit moth males, Grapholita molesta (Buck), provide evidence that a pheromone plume in zero wind elicits an endogenous, self-steered programme of counterturning (zigzagging) flight, and that wind experienced in flight establishes the polarity of the counterturns; they become aligned so that displacement occurs toward the source, even after the wind is stopped. In zero wind, males located a pheromone source more frequently when they had experienced a wind after having already taken flight before the wind was stopped (46%) compared with those that took flight later and therefore only experienced wind while they were in contact with the ground (14%). Furthermore, males placed in a stationary pheromone plume in zero wind located the source, eventually, on 21% of occasions. The flight tracks of these males, as well as those having experienced a wind only while on the ground, often exhibited repetitive counterturns (zigzags) of c. 180–200. However, the counterturns meandered around the flight tunnel, the inter-reversal track angles having no consistent direction. Sometimes the males displaced down-tunnel in the stationary plume, sometimes up, eventually locating the source and performing a courtship display. The inter-reversal track angles of males counterturning in wind, on the other hand, displayed a consistent orientation of c. 60 to either side of the wind line, resulting in consistent upwind displacement toward the source. With no pheromone present, with or without wind, counterturns were not observed.     

Manoeuvres used by flying male oriental fruit moths to relocate a sex pheromone plume in an experimentally shifted wind-field

ABSTRACT. In a wind-field experimentally shifted in direction by 35d?, flying male Grapholita molesta (Busck) zigzagging upwind either maintained contact with a pheromone plume and followed it across during the shift or lost it and commenced casting at c. 90d? across the shifting windline to locate it eventually in its new position. Males accomplished both of these results by integrating the previously described systems of optomotor anemotaxis and self-steered counterturning, but with faster reaction-times to pheromone on and off than heretofore calculated for this species. We found no evidence that males following the plume across used chemotaxis as proposed for another species, Rather, the sawtoothed-shaped tracks were a result of the anemotactic and counterturning systems responding rapidly and reiteratively to each loss and gain of pheromone along the plume in the shifting wind. The response to an increase or decrease in pheromone concentration by males was to change their course angle to more upwind or more crosswind, respectively, on the very first reversal (within c. 0.15 s) after the concentration changed. Because males adjusted their airspeeds more slowly to changes in concentration, the groundspeeds along the more upwind-orientated legs were lower than those along cross-wind legs, contributing to the sawtoothed shape of tracks of plume-followers. The self-steered counterturning programme also reacted quickly to concentration changes, the reversal intervals tending to be shorter following each contact with pheromone than after each excursion into cleaner wind. Following casting after losing the plume, males relocating the pheromone plume exhibited an upwind ‘surge’ of narrow zigzagging flight because on the first leg in the plume they steered a course more directly upwind than on the previous leg and increased the frequency of counterturning to its highest value while maintaining the relatively high airspeed acquired while casting.     

Pheromone-mediated upwind flight of male gypsy moths, Lymantria dispar, in a forest

Abstract Lymantria dispar L. males flying upwind in a pheromone plume in a forest were video-recorded at 2.5, 10 and 20 m from the source of pheromone. Males flew slower and steered more across the wind as they approached the source. In concert, their ground speed decreased and track angles increased. In contrast to these changes, their drift angles were fairly constant and the transverse component of image flow, above and/or below the moths eyes, showed almost no change. The inter-turn duration (time between sequential turns), a temporal aspect of the male flight manoeuvres, showed a consistent but relatively small increase as the distance from the source increased. The flight tracks narrowed as the males approached close (2.5 m) to the source. Because of unpredicted correlations between physical variables (i.e. temperature, wind velocity) and the distance from the source, we used principal components analysis to generate a set of completely independent variables. Greater than 90% of the variability in the data could be explained by four principal factors which corresponded well with known relationships in the flight manoeuvres. All four of these factors showed a significant regression against distance to the source. Although uncontrolled factors such as temperature and wind velocity may have contributed to changes in flight behaviour, recent data indicate that, in addition to concentration, certain temporal and spatial characteristics (i.e. burst period, burst return period) of plumes in wind vary systematically with distance from the source. We propose that L.dispar males might adjust their flight manoeuvres in response to these changes.     

Mating disruption barriers in pome fruit for improved control of oriental fruit moth Grapholita molesta Busck (Lep., Tortricidae) in stone fruit under mating disruption

Oriental fruit moth (OFM) Grapholita molesta Busck (Lep., Tortricidae) is a very important pest in commercial orchards in Victoria, Australia. Pheromone‐mediated mating disruption (MD) applied in stone fruit orchards successfully controlled OFM populations for many years, but damage to shoot tips and fruit at the edge of peach blocks located adjacent to pear blocks under insecticide treatments has become problematic. To improve protection of stone fruit against edge damage and outbreaks of OFM, all orchards were treated with sex pheromone dispensers for MD on an area‐wide basis. Area‐wide MD treatment, including all pome and stone fruit orchards in a discrete area, successfully controlled edge infestations of OFM, but was expensive. To reduce the cost of OFM control, sex pheromone dispensers for MD were applied as barrier treatments to 54–60 m of neighbouring pears adjacent to peaches under MD. Detailed monitoring of the OFM population, shoot tip and fruit damage assessments indicated that application of MD barriers on pears during two consecutive seasons provided sufficient control of OFM on peaches. This MD barrier treatment was able to reduce the number of OFM caught in all experimental peach blocks, with damage to shoot tips and fruit giving similar results to MD treatment of the whole neighbouring pear block. Extending the MD treatment area for 54–60 m into the neighbouring pear block significantly reduced the edge damage in MD‐treated peaches in the first season and almost eliminated OFM damage in the second season.     

Effects of intermittent and continuous pheromone stimulation on the flight behaviour of the oriental fruit moth, Grapholita molesta

ABSTRACT. When male oriental fruit moths, Grapholita molesta (Busck) (Tortricidae), casting in clean air entered an airstream permeated with pheromone their flight tracks changed immediately on initial contact with pheromone, but after a few seconds returned to casting as if in clean air. The degree of change in the flight track was directly related to the concentration of pheromone. Although little net uptunnel movement occurred in response to the continuous stimulation provided by a uniformly permeated airstream, when an intermittent stimulus provided by a point-source plume was superimposed onto the permeated airstream moths were able to 'lock on' and zigzag uptunnel in the plume. The percentage of moths doing so corresponded to the difference between the peak concentration within the plume and the background concentration of pheromone permeating the airstream. Moths also locked onto, and flew upwind along the pheromone-clean-air boundary formed along a pheromone-permeated side corridor. Because a similar response was observed along a horizontal edge between a pheromone-permeated floor corridor and clean air, we conclude that the intermittent stimulation at the edge perpetuated the narrow zigzagging response to pheromone.     

Field and laboratory electroantennographic measurements of pheromone plume structure correlated with oriental fruit moth behaviour

Abstract. Peak-to-trough electroantennogram amplitudes (bursts), caused by the individual filaments of a plume of female pheromone, diminish as high-emission-rate sources are approached by male Grapholita molesta , and this reduction is correlated with in-flight arrestment (ceasing to advance upwind). These findings are consistent with the hypothesis that one cause of in-flight arrestment in response to high-concentration point sources is the attenuation of the peak-to- trough amplitudes close to the source. High burst frequency, high pheromone flux, or low levels of continuous neuronal activity all are less well correlated with arrestment. Rather, arrestment appears due to a reduction of chemosensory input to the CNS during flight up the plume, even though the actual molecular concentration continues to increase. In a laboratory wind tunnel, upwind flight initiation by more than 20% of males was elicited only by pheromone source concentrations evoking significant fluctuations in EAG amplitudes at downwind release points. The burst frequencies that evoked high levels of upwind flight initiation ranged from a mean of 0.4-2.2 bursts/s. Because a previous study revealed that flying male G. molesta change their course angle within 0.15 s of losing or contacting pheromone, these EAG burst frequencies indicate that during flight in a pheromone plume, many manoeuvres are probably made in response to contact with individual plume filaments. Thus, upwind flight tracks may be shaped by hundreds of steering reactions in response to encounters with individual pheromone filaments and pockets of clean air. Field-recorded EAGs reveal that burst amplitudes diminish from 3 to 30 m downwind of the source, whereas burst frequencies do not, averaging c. 1/s at 3, 10 and 30 m downwind.     

Volatile compounds from young peach shoots attract males of oriental fruit moth in the field

Abstract

The oriental fruit moth (OFM) is one of the most serious pests of commercial fruit orchards worldwide. Newly planted peach trees in particular, can be very attractive for mated OFM females for oviposition. Samples of airborne host plant volatiles from intact young peach shoot tips and old leaves of the same potted plants were collected and analyzed with a gas chromatograph-mass spectrometer. Chemicals present in young shoot tips, but not in old leaves, were used for field trials. Moth capture by traps with the synthetic chemicals was compared to that of the standard terpinyl acetate (TA) food trap. The TA food trap caught OFM males and mated females, but tested synthetic chemicals trapped only OFM males. We observed that the mixture of (Z)-3-hexenyl acetate: (E)-β-ocimene: (E)-β-farnesene in proportion 1:2:2 attracted OFM males. Further, 1 mg of (E)-β-ocimene, and that of (E)-β-farnesene also attracted OFM males.     

A non-anemotactic mechanism used in pheromone source location by flying moths

ABSTRACT. Male oriental fruit moths, Grapholitha molesta (Busck) (Tortricidae), continue to zigzag along a pheromone plume to the source in zero wind, if they have started flight with wind on. If the pheromone source is removed and the plume is hence truncated, moths flying in zero wind out of the end of the plume into clean air increase the width of their reversals and the angles of the straight legs of the tracks so they are more directly across the former wind line. Such moths reach the source less often than do those flying along a continuous plume. The males continue to zigzag up a plume in zero wind, apparently by a combination of sequential sampling of concentration along their path and the performance of an internal, self-steered programme of track reversals (zigzags) whose frequency increases with concentration. Visual feedback may aid in the still-air performance of the zigzags. We propose that both the sequential sampling (longitudinal klinotaxis) and self-steered counter-turning programme also are used in wind as well; anemotaxis apparently polarizes the direction of the zigzags to result in upwind displacement, and the narrow zigzags caused by the higher concentration in the plume keep the male 'locked on' to the odour.     

The effects of pheromone concentration on the flight behaviour of the oriental fruit moth, Grapholitha molesta

ABSTRACT. Male oriental fruit moths, Grapholitha molesta (Busck) (Tortricidae), flew at lower overall and net ground velocities when they flew toward higher concentration pheromone sources. Turning frequency was greater with increased pheromone concentration, while the distance of turns from the plume axis back towards the axis decreased. Turning magnitude and inter-reversal track angles remained constant at all concentrations tested. Concomitant with the changes in ground velocity but constant inter-reversal angles, were decreases in airspeed, decreases in the moths' course angles and increases in their drift angles. The significance of these changes is discussed in relation to their possible role in a longitudinal chemoklinotactic programme of turning operating in conjunction with anemotaxis to allow location of a pheromone source in wind.     

Structured populations of the oriental fruit moth in an agricultural ecosystem

Intercontinental trade has led to multiple introductions of invasive pest species at a global scale. Molecular analyses of the structure of populations support the understanding of ecological strategies and evolutionary patterns that promote successful biological invasions. The oriental fruit moth, Grapholita (=Cydia) molesta, is a cosmopolitan and economically destructive pest of stone and pome fruits, expanding its distribution range concomitantly with global climate warming. We used ten newly developed polymorphic microsatellite markers to examine the genetic structure of G. molesta populations in an agricultural ecosystem in the Emilia‐Romagna region of northern Italy. Larvae collected in eight sampling sites were assigned to a mosaic of five populations with significant intra‐regional structure. Inferred measures of gene flow within populations implicated both active dispersal, and passive dispersal associated with accidental anthropogenic displacements. Small effective population sizes, coupled with high inbreeding levels, highlighted the effect of orchard management practices on the observed patterns of genetic variation within the sampling sites. Isolation by distance did not appear to play a major role at the spatial scale considered. Our results provide new insights into the population genetics and dynamics of an invasive pest species at a regional scale.     

梨小食心虫生物防治研究进展

梨小食心虫Grapholita molesta(Busck)是世界性分布的果树主要害虫之一,可危害多种果树。多年来,过度依赖化学农药防治梨小食心虫效果并不理想,且杀伤天敌、污染环境、导致农药残留。利用自然天敌防治梨小食心虫高效、无毒、无污染,符合当前社会对环保的要求。本文结合前人工作,从病原微生物、寄生性天敌、捕食性天敌、性信息素、化学信息物质等方面,综述了梨小食心虫生物防治的研究进展,并对其应用前景进行了展望。     

Distribution of the oriental fruit moth Grapholita molesta Busck (Lep., Tortricidae) infestation on newly planted peaches before and during 2 years of mating disruption

Abstract:  Oriental fruit moth (OFM) Grapholita molesta Busck (Lep., Tortricidae) is one of the most important pests of commercial stone fruit orchards in the Goulburn-Murray Valley region of Victoria, Australia. OFM populations have been successfully controlled by the use of the mating disruption (MD) technique for many years, but damage to shoot tips and fruit has now started to increase. The most severe damage under MD is found at the edge of peach blocks, adjacent to the pear blocks under insecticide treatment. In 1997–98, OFM infestation levels were examined in a newly planted peach block surrounded by older peaches, pears, apples and pasture. The infestation distribution was followed up for four consecutive years. No treatments were used against OFM for the first 2 years in the newly planted peaches, but in years 3 and 4 the whole block was treated with MD. At the end of year 2, shoot tip damage was randomly distributed throughout the newly planted peach block with no 'edge effect'. After MD was applied in year 3, the damage was confined to the edges of the block adjacent to insecticide-sprayed apples and pears. No 'edge effect' was detected along the border with an older peach block treated with MD or on the border with pasture. Extending the MD treated area for 25–30 m into the neighbouring apples and pears in year 4 reduced the 'edge effect'.     

Comparative analysis of peach and pear fruit volatiles attractive to the oriental fruit moth,Cydia molesta

Cydia molesta is an important pest of peach and pear fruits late in growing season. We identified and quantified volatiles from immature and mature fruits of peach and pear using gas chromatography-mass spectrometry. Antennally, active compounds were identified by gas chromatography-electroantennogram detection and were further tested in the field. Consistent electroantennographic activity was obtained for nine compounds from headspace collections of the mature fruits of two peach and two pear varieties. Esters were the major components, but qualitative and quantitative differences were found among four odor profiles. Blends mimicking pear fruit volatiles were more attractive to both sexes than blends mimicking peach fruit volatiles in both orchards. Our finding indicated that mixtures mimicking peach and pear fruit volatiles attracted both females and males of C. molesta, and these host volatiles may also be involved in the host switch behavior from peach to pear in mid-late season.     

梨小食心虫性信息素田间应用技术研究

梨小食心虫Grapholitha molesta(Busck)是梨园、桃园重要的果树害虫。本文比较研究了梨小食心虫性信息素在梨园、桃园悬挂不同高度、设置不同密度对梨小食心虫雄性成虫诱捕效果的影响,结果表明:在梨树树冠范围内,梨小食心虫性信息素最佳设置高度为2 m;在桃树树冠范围内,梨小食心虫性信息素最佳设置高度为1.5 m;同时,田间诱捕效果与性信息素设置密度密切相关,梨园中,每0.067 hm2设置密度为6¹0个诱芯时,对梨小食心虫的诱捕效果影响不大,而桃园中每0.067 hm2设置8个诱芯时,有最好的诱集效果。本研究可为梨小食心虫性信息素田间应用提供指导。     

A pulsed cloud of sex pheromone elicits upwind flight in male moths

ABSTRACT. Male oriental fruit moths do not fly upwind in a continuous uniform cloud of pheromone, but readily do so when the cloud is pulsed at 1 or 0.5/s or when a plume from a point source of pheromone is placed within the continuous cloud. It is suggested that males of moth species that require such fluctuating pheromone stimulation for upwind flight will normally receive it from a filamentous, point-source-produced plume. However, we hypothesize that upwind progress may cease close to the source due to excessively high emission rates or inappropriate blend ratios, when fluctuating sensory output becomes attenuated, despite higher actual molecular concentration fluctuations.     

Dual pheromone dispenser for combined control of codling moth Cydia pomonella L. and oriental fruit moth Grapholita molesta (Busck) (Lep., Tortricidae) in pears

Codling moth (CM) and oriental fruit moth (OFM) are very important orchard pests worldwide, and particularly in Victoria, Australia, where both species damage pome fruit. Individually CM and OFM have been controlled successfully by pheromone‐mediated mating disruption, but treatment of pome fruit with full registered rates of two individual hand‐applied dispensers for CM and OFM could be uneconomical for growers. Field trials conducted over three seasons in plots sprayed with insecticides consistently demonstrated that dual Isomate C/OFM TT dispensers, designed to disrupt both CM and OFM, were as effective as Isomate CTT and Isomate OFM Rosso dispensers applied individually in pears for control of CM and OFM respectively. The dual‐ and individual‐species dispensers reduced moth catches and fruit damage to a similar degree. The results suggest that combined control of CM and OFM in pears by applying dual Isomate C/OFM TT dispensers at the full‐recommended rate of 500 dispensers per hectare will be more economical than use of individual species dispensers, because the price and application cost of dual dispenser is about half that for individual dispensers.     

An analysis of anemotactic flight in female moths stimulated by host odour and comparison with the males' response to sex pheromone

ABSTRACT. The flight pattern of mated female navel orangeworm moths, Amyelois transitella (Walker), responding to odour from potential larval hosts is zigzagging upwind flight. However, at times these moths are capable of flying nearly directly upwind towards the odour source (track angles near 0). This response indicates that these females are capable of very accurate anemotactic control of their heading or course angle, since small angular errors in this measure would translate into larger deviations from direct upwind flight. Males of this species exhibit flight patterns similar to those of females, including track angles clustered about 0 when flying upwind to a source of the female-produced pheromone, but under these experimental conditions they flew with a higher average airspeed than the females. When females lose contact with an odour plume they initiate a well-defined programme of cross-wind counterturning or casting, which may normally increase their chances of retrieving contact with that plume when the wind direction shifts. The resultant track angles of females increase significantly by 0.8 s after plume loss, indicating that the female has initiated changes in both her course angle and airspeed. By 1 s after plume loss the females' track angles are no longer unimodally distributed about 0, but are bimodally distributed about -90 and +90. Males responded more rapidly to the loss of a pheromone plume, demonstrating a significant change in track angle 0.4 s after plume loss. Overall, female and male A.transitella exhibited remarkably similar anemotactic flight manoeuvres during upwind flight to odour sources as well as after plume loss.     

A comparison of various mating disruption technologies for control of two internally feeding Lepidoptera in apples

Sex pheromone mating disruption (MD) is an approach used to control several moth pest species of pome fruit by disrupting the ability of the males to find females and consequently prevent mating. The following experiments were performed to determine the effectiveness of several new and experimental sex pheromone MD technologies, and dispenser densities for simultaneous control of the codling moth (CM), Cydia pomonella (L.), and the oriental fruit moth (OFM), Grapholita molesta (Busck) (both Lepidoptera: Tortricidae), in Pennsylvania apple orchards. In one study, three MD approaches to control CM and oriental fruit moth – CM and OFM Disrupt Micro‐Flakes, Isomate CM/OFM TT, and both a CideTrak OFM and a CideTrak CM dispenser containing both codlemone and pear ester – and an insecticides‐only treatment were compared over the course of 2 years. In the other studies, the efficacy of several CheckMate Duel dispenser densities (i.e., 250, 375, 425, and 500 dispensers ha^?1) were compared against Isomate CM/OFM TT, and an insecticides‐only treatment. The CideTrak CM/pear ester combination and Isomate CM/OFM TT treatments both substantially reduced CM captures in traps in 2007 and 2008. Meanwhile, OFM trap shutdown was highest in the CheckMate Duel densities of 375 (99.9 ± 0.08%) and 500 dispensers ha^?1 (98.9 ± 0.07%) and the Isomate CM/OFM TT treatment (98.0 ± 1.13%), and lowest in the 250 dispensers ha^?1 density treatment (94.3 ± 3.23%). In orchards where OFM is the dominant pest species, a CheckMate Duel dispenser density of 375 ha^?1 is necessary for effective control, whereas higher densities are needed to control CM.     

Effect of varying proportions of the alcohol component on sex pheromone blend discrimination in male Oriental fruit moths

ABSTRACT. The flight response of individual male Oriental fruit moths, Grapholitha molesta (Busck), was observed in a sustained-flight tunnel to 100 blend–dosage combinations of the three sex pheromone components: (Z)- and (E)-8-dodecenyl acetate and (Z)-8-dodecen-l-ol (1, 3, 10, 30 and 100 μg of Z8-12: AC with, 2, 6, 10, 20% E and, 0, 3, 10, 30 or100% OH alcohol added). Complete flights to the source were observed only to blend combinations containing all three components. Males exhibited highest response levels to two dosages (3 and 10μg) of the natural 6% E blend and these levels were relatively unaffected by changes in the proportion of Z8-12: OH. Certain treatments surrounding the peak area also elicited high response levels compared to the 6% E treatments, but these were strongly dependent on the proportion of OH in the blend. Hierarchical cluster analysis was utilized to compare and group treatments that elicited similar levels of response over the entire flight sequence and to derive an area of blend-dosage combinations within the matrix tested that elicited peak levels of attraction. Analysis of the response patterns for suboptimal treatments adjacent to the area of optimal attraction showed that response specificity was controlled by two major effects on flight behaviour, one occurring early in the flight sequence affecting male orientation to the odour plume, and the other occurring later in the sequence as an arrestment of upwind flight. These effects were strongly influenced by changes in the OH component and the E isomer, with the latter playing the critical role in effecting flight behaviour. Temporal analysis of the flight response to treatments within the optimal area showed that whereas the % OH did not significantly affect the magnitude of response, increasing the level of Z8-12: OH in the blend did significantly increase the duration of each phase of the behavioural sequence. Considering both the magnitude and temporal aspects of male response, optimal attraction in male OFM was best characterized by a small area of treatments around the peak 6% E blends that contained 10% OH. Finally, field tests showed a high degree of correlation between trends in response with changing proportion of Z8-12: OH as observed in the flight tunnel. Peak dosages were generally higher in the field, however, compared to those in the flight tunnel.