Attractants for the gravid Queensland fruit fly Dacus tryoni

The vapours of certain pure chemicals, typical of ripe fruits, elicited characteristic components of ovipositional behaviour from gravid Dacus tryoni (Froggat) in an olfactometer: the flies walked and flew upwind to the source of the vapour and then probed with their ovipositors. A range of alcohols, acids, ketones and esters having 2–6 carbon atoms were effective (1 and 10% of iso-butyric acid, n-butyric acid, methyl butyrate, ethyl butyrate, 2-butanone, ethyl lactate and ethyl acetate; and 10% concentrations of ethanol and 2-propanone). The most effective were 4–6 carbon acids, esters and ketones. Behavioural threshold for n-butyric acid vapour at 26°C was obtained from a 5×10^–3% dilution in paraffin oil; maximum fly response occurred at about 200 times this concentration. Low concentrations of the 15-carbon sesquiterpene, -farnesene, were also very effective, despite its lower volatility. These results suggest that at least three different types of alfactory sensory neurones are involved in the identification of fruit attractants by gravid D. tryoni.     

The effect of photoperiod on the circadian rhythm of mating responsiveness in the fruit fly, Dacus tryoni

ABSTRACT. The daily fluctuation in mating responsiveness of Dacus tryoni was recorded through a range of light cycles. Evidence was obtained of strict control of the fluctuation by a circadian clock. The phase setting of the fluctuation in the different light cycles was in conformity with Aschoff's (1965) generalizations about circadian rhythms, and had no relationship to overt sexual behaviour occurring during the cycles. The amplitude of the fluctuation in mating responsiveness varied between cycles, suggesting possible variation in the amplitude of the oscillation of the circadian clock.     

The dynamics of ovarian maturation and oocyte resorption in the Queensland fruit fly, Dacus tryoni, in daily-rhythmic and constant temperature regimes

The rate of oocyte development in constant regimes corresponded to the rates predicted by Pritchard's (1970) relationship which indicates that the lower temperature threshold is 13.5dgC. Rates in fluctuating regimes indicated that development occurs at certain temperatures below the constant threshold if these are alternated with higher temperatures on a daily basis
In certain fluctuating regimes, oocyte development occurred at a rate in excess of 3.5% per day and maturation proceeded to completion. In such regimes resorption occurred only if egg laying was prevented; when this happened the terminal oocytes usually remained intact and the penultimate ones were resorbed. In other fluctuating regimes, oocyte development proceeded at a rate of less than 3.5% per day and no maturation was achieved because the most advanced oocytes were resorbed-either before or during the vitellogenic phase.
The 3.5% threshold corresponds to a value of 2 day-degrees per day (2DDPD) above 13.5C. Resorption, but no development, was observed in partly mature flies around O DDPD, whereas neither occurred in a still colder regime. These results are shown to be in accord with Fletcher's (1975) field data which are also used to discuss the significance of resorption. Finally, the DDPD relationship is used to define those daily temperature profiles which may permit maturation in the field.     

Experimental analysis of the pattern of tethered flight in the Queensland fruit fly, Dacus tryoni

ABSTRACT. The tethered flight of the Queensland fruit fly, Dacus tryoni Frogg. (Diptera, Tephritidae), was investigated, and the duration of each flight during a 2-h experimental period was recorded. The pattern of flight was analysed, and related to the age, sex and origin of the specimens, and to the availability of host fruit during the rearing of the adults. The effect of adult crowding on the pattern of flight was also briefly examined. The results indicated that the origin of the flies had little effect on the pattern of flight; male and female flies showed different trends with respect to the proportion of short flights undertaken as the flies matured; and the availability of fruit had a marked effect on the pattern of flight in recently mature flies. These data are discussed with respect to the dispersive/non-dispersive movements of the flies postulated from previously documented field data. It is suggested that there is a characteristic pattern of tethered flight, which can be related to the absence of hosts in the immediate environment, and would be likely to lead to greater dispersal under natural conditions.     

Survival of step and ramp changes of temperature by the Queensland fruit fly, Dacus tryoni

ABSTRACT. Survival time at subzero temperatures is related to both long-term thermal history and the rate at which the insects are cooled. Insects cooled at one degree per hour survive for up to 3 times as long at a given test temperature than do insects cooled at 1C/min. Survival times are significantly shorter than times taken to freeze. Survival time at an extreme high temperature is related to long-term thermal history but the rate of heating makes no difference.     

Mark-recapture estimates of overwintering survival of the Queensland fruit fly,Dacus tryoni,in field cages

1. Four groups of 75 pairs each of marked adult Dacus tryoni that had emerged from field-collected guavas were liberated into cage-covered guava trees at the end of May, 1960 and sampled at weekly intervals to estimate the overwintering survivorship.
2. Estimates of numbers were made using three different methods as described byLeslie (1952) and one byJolly (1965). The Method A, approximate Method B andJolly models gave quite similar results although the Method A model was shown to be more sensitive to stochastic changes in the R/r ratios. The complete Method B model produced much smaller variances, due to its full utilization of the information in the data but it was very sensitive to small biases—even those caused by the restriction that recaptures must be whole numbers—and thus could not quite approach the exact expectations.
3. Although in some instances, statistical tests (Leslie , 1958) showed some significant deviations from randomness in recapture, these were not appreciable and did not affect the estimates.
4. When sufficient food was available, the flies in the field cages exhibited survival rates during the winter months of 1960 quite comparable to those of laboratory flies kept at room temperature with several exceptions: (a) there was a high mortality among the newly emerged flies during the initial week after release, (b) a high mortality during the last week of June and (c) a high mortality during the third week in July and the end of August which can be correlated with lack of rain, low humidity and warm daytime temperatures. Low temperatures apparently had little effect on survival.
5. There was no difference in the survival of males and females.
6. One replicate cage, whose flies had been anesthetized with carbon dioxide prior to release, exhibited significantly higher mortality rates than the other replicates.
7. Data from a fifth cage indicated that females were not inseminated during the winter months and did not produce mature eggs until the beginning of September.

     

The role of adult learning in the acceptance of host fruit for egglaying by the Queensland fruit fly, Dacus tryoni

Evidence presented indicates that exposure of wild Queensland fruit files, Dacus tryoni (Froggatt), to a given host fruit type (e.g. pear) for 3 days causes females to attempt oviposition to a greater degree in that fruit type than in other fruit types (e.g. tomato, grape). The effects of exposure to a particular fruit type proved reversible, suggesting that D. tryoni females were capable of learning. Females exposed to pear for 3 days appeared to retain the effect of such exposure on acceptance of tomato for up to 4 days but appeared to retain the effect on acceptance of grape for less than 2 days. The possible significance of prior experience of females with a particular fruit type on future ability to discriminate among varying-quality specimens of that type is discussed.

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Résumé Les éléments apportés montrent que l'exposition de D. tryoni Froggat à un type de fruit déterminé (par exemple, la poire) pendant 3 jours, conduit la femelle à tenter de pondre à une plus haute fréquence dans ce type de fruit que dans d'autres (par example; tomate ou raisin). Les effects de l'exposition à un type particulier de fruit étant réversibles, ceci suggère que les femelles de D. tryoni sont capables d'apprentissage. Des femelles exposées à des poires pendant 3 jours ont conservé l'effet d'une telle exposition pendant 4 jours en présence de tomates, mais moins de 2 jours en présence de raisin. La discussion porte sur la signification possible de la connaissance antérieure d'un type particulier de fruit sur l'aptitude ultérieure des femelles à distinguer parmi des spécimens de ce type, mais de qualités différentes.

     

Development and ultrastructure of the fat body cells and oenocytes of the Queensland fruit fly,Dacus tryoni (Frogg.)

Summary Half-way through the larval period in Dacus tryoni, the fat body cells begin to accumulate protein in the form of granules. Early in the pupal period, both the fat body cells and oenocytes become free in the body cavity. Meanwhile, an imaginal generation of hypodermal cells, while in the process of displacing the larval hypodermis, gives rise to an imaginal generation of oenocytes. Soon after, imaginal fat body cells also appear. A few days after emergence, the larval fat body cells and oenocytes disintegrate and their imaginal equivalents expand to fill the body cavity.This paper also describes the ultrastructure of the larval and imaginal fat body cells and of the imaginal oenocyte. In all three, tubular invaginations of the plasma membrane occupy the peripheral cytoplasm. At most stages, the fat body cells contain a considerable quantity of slightly distended, rough endoplasmic reticulum, which suggests that when these cells are not sequestering protein, they are secreting it into the blood. The imaginal oenocytes are packed with smooth endoplasmic reticulum, which supports other evidence that they participate in the synthesis of cuticular wax.For assistance with the electron microscopy, I thank Mr. Tony Webber and Miss Ann Miller of the Electron Microscopy Unit at Sydney University. For the loan of some sectioned material, I am grateful to Dr. D. T. Anderson.     

Genetic and molecular markers of the Queensland fruit fly, Bactrocera tryoni

Twenty-six microsatellite markers, along with two restriction fragment length polymorphism (RFLP) markers and three morphological markers, have been mapped to five linkage groups, corresponding to the five autosomes of the Queensland fruit fly, Bactrocera tryoni. All these molecular and genetic markers were genotyped in three-generation pedigrees. Eight molecular markers were also localized to the salivary gland polytene chromosomes by in situ hybridization. This provides a substantial starting point for an integrated genetic and physical map of B. tryoni.     

The integument of the Queensland fruit fly,Ddacus tryoni (Frogg.)

Summary During the pupal stage of Dacus tryoni, the hypodermis of the larva is replaced by an imaginal generation of smaller cells. The hypodermal cells of the tergal glands on the fifth abdominal segment of the adult were examined with the electron microscope; they contain slender, membrane-limited bundles of hollow wax filaments that traverse the cuticle in branched pore canals. Outside the glandular areas, the pore canals are narrower. The cuticle of the adult undergoes its greatest increase in thickness soon after emergence; it becomes sclerotized gradually. No epicuticle was detected with either the light or electron microscopes.Early in adult development, bristles are formed over the general surface of the terga. Most of these are innervated by single, bipolar nerve cells, and have more or less enlarged trichogen cells that appear to secrete wax through pore-plates in the cuticle. The bristles in different regions of the abdomen range in function from pure sensory receptors to pure secretors. The sensory bristles on the tergal glands were examined with the electron microscope.For assistance with the electron microscopy, I thank Mr. Tony Webber and Miss Ann Miller of the Electron Microscopy Unit at Sydney University. — Supported by a C.S.I.R.O. Junior Post-Graduate Studentship.     

Times for recovery from cold-torpor in the Queensland fruit fly Dacus tryoni: the relation to temperature during and after chilling

Recovery time after experience of a given minimum temperature below torpor threshold is related to the value of that minimum, the length of time spent at that minimum, and the temperature prevailing during the recovery period above torpor threshold. A model can predict recovery time for flies experiencing a given temperature fluctuation if the length of time spent at the minimum is expressed as a proportion of LE₅₀ at that minimum.The model has applications in defining the optimal protocol for chilling insects for use in the Sterile Insect Release Method. The model was confirmed by experiments showing that it is likely that flies will recover from non-lethal frosts before ant predators become active.

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Résumé Le temps de récupération après avoir subi une température minimal située au-dessous du seuil d'engourdissement dépend de la valeur de ce minimum, du temps passé à ce minimum, et de la température au-dessus du seuil d'engourdissement pendant la période de récupération. Un modèle mathématique permet d'estimer le temps de récupération après avoir subi une chute de température déterminée, en fonction du temps passé au minimum thermique exprimé comme une fraction du LE₅₀ (temps nécessaire pour tuer 50% des mouches) à ce minimum.Ce modèle s'est trouvé étayé par des observations montrant qu'il est probable que les mouches se remettent des gelées sublétales avant la reprise d'activité des fourmis prédatrices. Ce modèle peut être utilisé pour définir les conditions optimales de refroidissement des insectes utilisés lors de la libération d'individus stériles.

     

Upwind anemotaxis in response to cue-lure by the Queensland fruit fly, Bactrocera tryoni

Movements of mature male Bactrocera tryoni (Froggatt) (Diptera: Tephritidae) were observed individually in a wind tunnel under conditions of ‘cue-lure with wind’, ‘cue-lure with no wind’, ‘wind only’ and ‘no wind or cue-lure’. Further observations were made using a dense foliage array in the wind tunnel and a structured plume of cue-lure. Patterns of walking or flying were essentially the same in all of the first four treatments except that in the ‘cue-lure with wind’ treatment, over half of the flies moved in a consistent track upwind for at least 400 mm at some time during the first 5 min of observation. With clean wind, only 10% of the flies did this. The result was that mean net upwind displacement after 5 min in the ‘cue-lure with wind’ treatment significantly exceeded that in the other three treatments, the results of which did not differ significantly from each other. The upwind tracks were accomplished by either walking or flying (with or without stops) or by a combination of both. When the wind tunnel was filled with a dense foliage array, the results with cue-lure laden wind were similar to those obtained with the equivalent treatment without foliage, except that upwind tracks were predominantly in short stages. When flies were exposed to a structured plume of cue-lure odour (without foliage present), they did not apparently alter their behaviour on leaving or entering the plume, but some did make consistent upwind tracks while they were in the plume.     

Mass-rearing and sterilisation alter mating behaviour of male Queensland fruit fly, Bactrocera tryoni (Froggatt) (Diptera: Tephritidae)

Abstract  The effects of domestication and irradiation on the mating behaviour of males of Bactrocera tryoni (Froggatt) (Diptera: Tephritidae) were investigated by caging wild, laboratory-domesticated and sterile (laboratory-domesticated, gamma-irradiated) males with wild females. Mating behaviour of mass-reared males was different from that of wild males, although behaviour of wild and sterile males was similar. Mass-reared males engaged in mounting of other males much more frequently than wild and sterile males, and began calling significantly earlier before darkness. Unnatural selection pressures imposed in mass-rearing conditions may explain these changes in mass-reared male behaviour. Male calling did not appear to be associated with female choice of mating partners, although this does not exclude the possibility that calling is a cue used by females to discriminate among mating partners. Despite differences in behaviour, frequency of successful copulations and mating success were similar among wild, mass-reared and sterile males.     

The effects of thermal acclimation on immature mortality in the Queensland fruit fly Bactrocera tryoni and the light brown apple moth Epiphyas postvittana at a lethal temperature

Mortality data for non-acclimated and acclimated 3rd instar larvae and mid-term eggs of Bactrocera tryoni (Froggatt) (Diptera: Tephritidae) were obtained after immersing in hot water at 46 °C. Acclimation consisted of holding the larvae and eggs at 35 °C for 20 and 11 h respectively just prior to heat-treatment. The median lethal time (LT50) for acclimated larvae was found to be 6.9 min compared to 2.5 min for non-acclimated larvae. LT99.999 for acclimated larvae was 20.9 min compared to 8.7 min for non-acclimated larvae. LT50 for acclimated eggs was 5.0 min compared to 2.4 min for non-acclimated eggs. LT99.999 for acclimated eggs was 26.0 min compared to 6.6 min for non-acclimated eggs. For 3rd instar larvae, most acclimation effect on mortality had occurred by 8 h. A notable residual response was present 20 h after acclimation had occurred, reducing mortality at 46 °C for 4.5 min by roughly 25%. Mortality data at 46 °C were also obtained for non-acclimated and acclimated late instar larvae of Epiphyas postvittana (Walker) (Lepidoptera: Tortricidae). With this species, LT50 for acclimated larvae was 2.5 min compared to 1.1 min for non-acclimated larvae. LT99.999 for acclimated larvae was 9.5 min compared to 4.6 min for non-acclimated larvae.     

Factors affecting the dispersal of large‐scale releases of the Queensland fruit fly,Bactrocera tryoni

This study investigated two factors potentially affecting the spatial distribution of Queensland fruit fly Bactrocera tryoni (Froggatt) after release from a single point. First, the effect of age at release was investigated using a single cohort of 205 000 males from a mass‐rearing strain used for sterile insect releases. Approximately half were released as immature males and the remainder as sexually mature males (1 week later). Males were collected over 3 weeks from a grid of 135 traps, each containing a pheromone/insecticide bait, positioned between 4 and 500 m from the release point. Variation in the distribution of fly density around the release point was assessed by regressing trapped fly counts against distance. Unexpectedly, no significant differences were found in the spatial distribution of the flies. Second, the effect of inbreeding on spatial distribution was investigated using replicated simultaneous releases of two strains of B. tryoni. One strain was the existing (inbred) mass‐rearing strain that has been selected for high productivity in a mass‐rearing facility. The second strain was deliberately outbred but also selected for high productivity. Almost 100 000 males of each strain were released over the two experiments. Regression of trappings against distance differed significantly between strains in only one of five releases, but in all cases the outbred strain had a greater dispersal distance. As our trapping grid was not regular but contained gaps of up to 100 m, a small preliminary experiment investigated whether flies move faster along tree rows or across open fields. At distances up to 100 m, we found no detectable difference in fly distributions. These results are primarily relevant to the large‐scale point releases carried out as part of an existing B. tryoni sterile insect programme and are discussed in that context.     

Oviposition behaviour of two tephritid fruit flies,Dacus tryoni and Dacus jarvisi,as influenced by the presence of larvae in the host fruit

Summary When offered a choice, females of the fruit flies Dacus tryoni (Frogg.) and D. jarvisi (Tryon) strongly preferred to lay in fruits without larvae rather than fruits which already contained larvae. Fruits which contained even low densities of larvae, including newly hatched ones, received many fewer eggs than control fruits. This preference was not influenced by the species of larvae present in the fruits nor by the distance to uninfested fruits. Discrimination occurred when fruits with and without larvae were close together (10 cm apart) and also when they were separated by distances of about one metre. Laboratory assays suggested that the flies detect chemical changes in the fruit associated with the decomposition which accompanies larval feeding, but they do not seem to detect the larvae Perse. This behaviour may be significant when these two species utilise the same host since the species which is able to infest fruits first will reduce the availability of hosts for the other species. In contrast to many other Tephritids (e.g. Rhagoletis, Anastrepha and Ceratitis) female Dacus don't discriminate against fruits which contain eggs nor do they deposit a pheromone to deter oviposition by females that subsequently visit the fruit. An hypothesis is proposed to explain the absence of oviposition-deterring pheromones in Dacus, and their presence in many other species of Tephritidae, on the basis of differences in life history and population structure.     

Seasonal trends in acclimatization to cold in the Queensland fruit fly (Dacus tryoni,Diptera) and their prediction by means of a physiological model fed with climatological data

Summary This paper demonstrates the predictability and significance of changes in the temperature threshold for cold-torpor and in the threshold for survival at sub-zero temperatures in two contrasting types of climate. Both thresholds alter on a seasonal basis in both types of climate and it is observed that temperatures prevailing in autumn cause acclimatization for winter long before winter arrives.Torpor thresholds can be accurately predicted in laboratory-simulated climates by use of a model. When the model is used for field predictions, using shade temperature data from the field, it is only successful in predicting accurate threshold values for those parts of the year during which there is a downward trend in threshold.The relative importance of torpor-avoidance and frost-resistance is discussed with reference to records of the frequency and extent of various degrees of cold in the field. It is concluded that the marked lowering of torpor threshold prior to winter is of great significance in enabling overwintering flies to stay in their sheltered hibernacula. The relatively slight lowering of temperature threshold for frost-survival is only of significance in certain climates. It greatly improves chances of forst-survival in climates where yearly minima fall predominantly in the range-2.65°C to-4.05°C. Acclimation is not necessary for frost survival in climates where the yearly minima stay above-2.65°C whereas acclimation is ineffective in preventing frostinduced extinction when yearly minima fall below-4.5°C.     

Germ-line transformation of the Queensland fruit fly, Bactrocera tryoni, using a piggyBac vector in the presence of endogenous piggyBac elements

We report the heritable germ-line transformation of the Queensland fruit fly, Bactrocera tryoni, using a piggyBac vector marked with either the fluorescent protein DsRed or EGFP. A transformation frequency of 5–10% was obtained. Inheritance of the transgenes has remained stable over more than 15 generations despite the presence of endogenous piggyBac sequences in the B. tryoni genome. The sequence of insertion sites shows the usual canonical pattern of piggyBac integraton into TTAA target sites. An investigation of endogenous piggyBac elements in the B. tryoni genome reveals the presence of sequences almost identical to those reported recently for the B. dorsalis complex of fruit flies and two noctuid moths, suggesting a common origin of piggyBac sequences in these species. The availability of transformation protocols for B. tryoni has the potential to deliver improvements in the performance of the Sterile Insect Technique for this pest species.     

Additive and interactive effects of nutrient classes on longevity, reproduction, and diet consumption in the Queensland fruit fly (Bactrocera tryoni)

Insect lifespan is often closely linked to diet, and diet manipulations have been central to studies of ageing. Recent research has found that lifespan for some flies is maximised on a very low yeast diet, but once all yeast is removed, lifespan drops precipitously. Although effects of yeast availability on lifespan are commonly interpreted in terms of protein, yeast is a complex mix of nutrients and provides a rich source of vitamins, minerals and sterols. Elucidating which components of yeast are involved in this lifespan drop provides insights into more specific nutritional requirements and also provides a test for the commonplace interpretation of yeast in terms of protein. To this end, we fed Queensland fruit flies (Bactrocera tryoni) one of eight experimental diets that differed in the nutrient group(s) found in yeast that were added to sucrose: none, vitamins, minerals, amino acids, cholesterol, vitamin+mineral+cholesterol (VMC), vitamin+mineral+cholesterol+amino acids (VMCA), and yeast. We measured survival rates and egg production in single sex and mixed sex cages, as well as nutrient intake of individual flies. We found that the addition of minerals increased lifespan of both male and female flies housed in single sex cages by decreasing baseline mortality. The addition of just amino acids decreased lifespan in female flies; however, when combined with other nutrient groups found in yeast, amino acids increased lifespan by decreasing both baseline mortality and age-specific mortality. Flies on the yeast and VMCA diets were the only ones to show significant egg production. We conclude that the drop in lifespan observed when all yeast is removed is explained by missing micronutrients (vitamins, minerals and cholesterol) as well as the absence of protein in females, whereas minerals alone can explain the pattern for males. These results indicate a need for caution when interpreting effects of dietary yeast as effects of protein.     

Critical periods for developmental acclimation to cold in the Queensland fruit fly, Dacus tryoni

Rapid acclimation to cold can occur in Dacus tryoni during two short stages of its life history: the stage immediately prior to the “hopping larva” phase and the “pharate adult” stage within the puparium. Transfer from 25 to 15°C at either of these stages can produce full acclimation to cold within a few days. Acclimation is not detectable at other times in puparial life: during adult life it takes over 100 days.