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.

     

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.

     

Management and eradication options for Queensland fruit fly

Several tephritid fruit flies have explosive population growth and a wide host range, resulting in some of the largest impacts on horticultural crops, reducing marketable produce, and limiting market access. For these pests, early detection and eradication are routinely implemented in vulnerable areas. However, social and consumer concerns can limit the types of population management tools available for fruit fly incursion responses. Deterministic population models were used to compare eradication tools used at typical densities alone and in combination against the Queensland fruit fly (‘Qfly’), Bactrocera tryoni. The models suggested that tools that prevent egg laying are likely to be most effective at reducing populations. Tools that induced mortality once Qfly was sexually mature only slowed population growth, as successful mating still occurred. Release of sterile Qfly when using the sterile insect technique (SIT) interferes with the successful mating of wild flies, and of the tools investigated here, SIT caused the greatest reduction in the population at the prescribed release rate. Used in tandem with SIT, protein baits slightly improved the rate of population reduction, but the male annihilation technique (MAT) almost nullified control by SIT due to the mortality induced on sterile flies. The model suggested that the most rapid decrease in population size would be achieved by SIT plus protein baits. However, the model predicted both the SIT and protein baits when used alone would result in population reduction. The MAT can be used prior to SIT release to increase overflooding ratios.     

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.     

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.     

Domestication-related changes in sexual performance of Queensland fruit fly

In Sterile Insect Technique (SIT) programs, massive numbers of insects are reared, sterilized, and released in the field to impede reproduction of pest populations. The domestication and rearing processes used to produce insects for SIT programs may have significant evolutionary impacts on life history and reproductive biology. We assessed the effects of domestication on sexual performance of laboratory reared Queensland fruit fly, Bactrocera tryoni, by comparing an old (49 generations) and a young colony (5 generations). We evaluated mating propensity, mating latency, copula duration, sperm transfer, and ability to induce sexual inhibition in mates. Overall, both males and females from the old colony had greater mating propensity than those from the young colony. Copula duration was longer when females were from the old colony. There was no evidence of sexual isolation between the colonies as males and females from the two colonies had similar propensity to mate with flies from either colony. Males from the old colony transferred more sperm regardless of which colony their mate was from. Finally, males from both colonies were similarly able to induce sexual inhibition in their mates and were also similarly able to secure copulations with already-mated females. Positive effects of domestication on sperm transfer, coupled with maintained ability to induce sexual inhibition in mates and to secure copulations with previously mated females, highlights that domestication may have little effect, or even positive effects, on some aspects of sexual performance that may advantage mass-reared B. tryoni in SIT programs.     

Evaluation of yeasts in gel larval diet for Queensland fruit fly,Bactrocera tryoni

Queensland fruit fly, Bactrocera tryoni (“Q‐fly”), is Australia’s most economically important insect pest of horticultural and commercial crops especially in the eastern regions. The sterile insect technique (SIT) has been adopted as an environmentally benign and sustainable approach for management of Q‐fly outbreaks. High‐performance larval diets are required to produce the millions of flies needed each week for SIT. Yeast products contribute amino acids (protein) to fruit fly larval diets, as well as carbohydrate, fat and micronutrients, but there can be substantial variation in the nutritional composition and suitability of yeast products for use in larval diets. Gel larval diets have recently been developed for large‐scale rearing of Q‐fly for SIT, and composition of these diets requires optimization for both performance and cost, including choice of yeast products. We assessed performance of Q‐flies reared on gel larval diets that contained debittered brewer’s yeast (Lallemand LBI2240), hydrolysed yeast (Lallemand FNILS65), inactivated brewer’s yeast (Lallemand LBI2250) and inactivated torula yeast (Lallemand 2160‐50), including blends. Q‐flies performed poorly when reared on diets containing only or mostly hydrolysed yeast in terms of pupal number, pupal weight and percentage of fliers. Performance was also poor on diets containing high proportions of torula yeast. Overall, debittered brewer’s yeast is recommended as the best option for Q‐fly gel larval diet, as it is cheap, readily available, and produces flies with good performance in quality control assays. Inactivated brewer’s yeast produced flies of comparable quality with only a modest increase in cost and would also serve as an effective alternative.     

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.     

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.

     

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

Summary Testes of the Japanese freshwater turtle Clemmys japonica Temmnick et Schlegel were fixed in 3% potassium permanganate buffered to pH 7.2 with Veronal-acetate buffer, and thin sections of the tissue, embedded in epoxy Epon resin, were studied under the electron or light microscope.At the early stage of differentiation of the spermatid, the cytoplasm contains a few mitochondria provided with cristae which are oriented transversely or longitudinally. As the differentiation of spermatids proceeds, the mitochondrion has been modified into a cupshaped body with a wall consisting of several concentric layers. Such body has been referred to the mitochondrial lamellar body. The formation of such a body is mainly attributed to the mitochondrial cristae, and subsequently to the membrane system of the endoplasmic reticulum. In a more advanced stage of differentiation, the mitochondrial lamellar bodies appear wrapped around a bundle of tail filaments, and seem to present a very wide surface available for the localization of organized enzyme systems to facilitate the motion of spermatozoa.Prior to the formation of the mitochondrial lamellar bodies, the Golgi apparatus has been reorganized into a peculiar body with a floral appearance, consisting of numerous tubular elements, and revealing to be positive in PAS-reaction. The body has been designated as the tubular body which has never been demonstrated in any spermatogenic cells through animal kingdom.One to three tubules oval in cross section, approximately 430 × 700 Å in diameter, have been found in the nucleoplasm along the longitudinal axis of a greatly elongated, cone-shaped nucleus of the spermatid. The tubules open on the apex surface of the nucleus, but they are not encountered in the acrosome. A possible physiological significance of the tubules has been discussed in view of the function of the acrosomal tubules in the decapod and other species spermatozoa as well as on the basis of the metabolism of nucleus.This study was supported by Grant GM-8327 from the United States Public Health Service.We wish to express our gratitude to Dr. B. A. Afzelius, Wenner-Gren Institute, University of Stockholm, for his valuable suggestion to the present work.     

Fecundity,fertility and reproductive recovery of irradiated Queensland fruit fly Bactrocera tryoni

Pupae of the Queensland fruit fly or Q‐fly Bactrocera tryoni (Froggatt) are irradiated routinely to induce reproductive sterility in adults for use in sterile insect technique programmes. Previous studies suggest that adult sexual performance and survival under nutritional and crowding stress are compromised by the current target dose of radiation for sterilization (70–75 Gy), and that improved mating propensity and survival under stress by irradiated males may be achieved by reducing the target sterilization dose without reducing the level of induced sterility. This raises the question of the amount by which the irradiation dose can be reduced before residual fertility becomes unacceptable. The present study measures the levels of residual fertility in male and female irradiated Q‐flies at different irradiation doses (20, 30, 40, 50, 60 and 70 Gy), and investigates the possibility that fecundity and fertility increase between 10–15 and 30–35 days post emergence. Male flies require a higher dose than females to induce sterility, with no residual fertility found in females irradiated at doses of 50 Gy or above, and no residual fertility found in males irradiated at doses of 60 Gy or above. Irradiated females are more fecund at 30–35 days post emergence than at 10–15 days. However, fertility does not increase between 10 and 15 days post emergence and 30–35 days, even at doses below 50 Gy. The present study shows that there is scope to reduce the target sterilization dose for Q‐flies below that of the current dose range (70–75 Gy) at the same time as retaining an adequate safety margin above radiation doses at which residual fertility can be expected.     

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.     

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.     

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.     

Yeast hydrolysate supplement increases starvation vulnerability of Queensland fruit fly

Post‐teneral diets containing yeast hydrolysate are reported to increase longevity, reproductive development and sexual performance of Queensland fruit fly (‘Q‐fly’) Bactrocera tryoni Froggatt (Diptera: Tephritidae). Consequently, diets including yeast hydrolysate are recommended for sterile Q‐flies before release in sterile insect technique (SIT) programmes. However, in some tephritids, diets including yeast hydrolysate are associated with an increased vulnerability to starvation. In the present study, the effects of yeast hydrolysate supplementation before release are considered with respect to the longevity of released Q‐fly when food becomes scarce. Experiments are carried out in three settings of varying resemblance to field conditions: 5‐L laboratory cages, 107‐L outdoor cages and 14 140‐L field cages containing potted citrus trees. In all experimental settings, compared with flies that received only sucrose, male and female Q‐flies that are provided with yeast hydrolysate during the first 2 days of adult life have a significantly shorter survival when subsequently deprived of food. Yeast supplementation appears to commit Q‐flies to a developmental trajectory that renders them more vulnerable to starvation. The practical significance of these findings for SIT depends on how often the releases are carried out under conditions in which Q‐flies experience extreme food shortages in the field.     

Rapid acclimatization to low temperature in the Queensland fruit fly, Dacus tryoni

Dacus tryoni, which overwinters in conditions of wide diurnal fluctuations in temperature, is capable of rapid acclimatization with respect to thresholds for torpor and flight but ability for metabolic compensation is apparently lacking. The rate of reacclimatization on abrupt transfer to lower temperatures is quicker the less the temperature drops and extrapolation of this relationship suggests a time of the order of 1 min for a drop as small as 1°C. This explains why continuous reacclimatization appears to take place during gradual cooling, even when this is as fast as 1°C/min. The insect can rapidly produce its maximum response to extensive drops of temperature as long as the latter falls gradually at a rate no faster than 1°C/min. Acclimatization in nature is therefore ‘immediate’. Reacclimatization to higher temperatures (deacclimatization to cold) depends upon the temperature that the insects are raised to and thus is slower the smaller the rise and understandably is not accomplished more rapidly by a gradual transition.     

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.     

The influence of host fruit morphology on parasitization rates in the Caribbean fruit fly,Anastrepha suspensa

Among the host fruits of the Caribbean fruit fly there are a variety of sizes and shapes. These morphological differences may influence the vulnerability of the larvae to parasites. In the laboratory, Caribbean fruit fly larvae placed in the smaller of 2 different sizes of artificial ‘fruit’ (cloth spheres filled with a diet material) were parasitized at a higher rate by the braconid,Diachasmimorpha longicaudata (Ashmead) when spheres were presented separately. However, when parasites were simultaneously presented with 6 different sizes of ‘fruit’ there was no significant relationship between size and parasitization rate. This may be due to the parasites preference to search for larvae in larger ‘fruit’. In field collections of different species of host fruit, a significant inverse correlation exists between fruit radius and rate of parasitization. However, host fruit size accounts for only about 5% of the variance in yearly parasitization rates.      

Behavioural responses of female Queensland fruit fly, Bactrocera tryoni, to mineral oil deposits

Behavioural responses of Queensland fruit fly, Bactrocera tryoni (Froggatt) (Diptera: Tephritidae), females to fruit dipped in water and fruit dipped in 0.5% (vol/vol) aqueous emulsions of a mineral oil were determined and analysed. The mineral oil was an nC20–22 distillation fraction of the base oil used to produce an nC23 horticultural mineral oil. Females caged with oil‐treated fruit had significantly longer prelanding intervals than females caged with water‐dipped fruit. The latter was attacked immediately or shortly after being caged with flies whereas some oil‐dipped fruit was not attacked within 180 min. The percentage of landings that led to oviposition on water‐ and oil‐treated fruit were 58 and 13%, respectively, and the percentages ovipositing after probing were 74 and 25%, respectively. Likewise, average times spent probing were 7 vs. 31 s whereas average times spent ovipositing were 321 vs. 223 s. Females spent less than half as much time on oil‐treated fruit than on water‐treated fruit. Transition probabilities of rejection, when applied to the behaviour sequence indicated that oil‐treated fruits are about nine times less likely to be infested with B. tryoni.     

Cost of reproduction in the Queensland fruit fly: Y-model versus lethal protein hypothesis

The trade-off between lifespan and reproduction is commonly explained by differential allocation of limited resources. Recent research has shown that the ratio of protein to carbohydrate (P : C) of a fly''s diet mediates the lifespan–reproduction trade-off, with higher P : C diets increasing egg production but decreasing lifespan. To test whether this P : C effect is because of changing allocation strategies (Y-model hypothesis) or detrimental effects of protein ingestion on lifespan (lethal protein hypothesis), we measured lifespan and egg production in Queensland fruit flies varying in reproductive status (mated, virgin and sterilized females, virgin males) that were fed one of 18 diets varying in protein and carbohydrate amounts. The Y-model predicts that for sterilized females and for males, which require little protein for reproduction, there will be no effect of P : C ratio on lifespan; the lethal protein hypothesis predicts that the effect of P : C ratio should be similar in all groups. In support of the lethal protein hypothesis, and counter to the Y-model, the P : C ratio of the ingested diets had similar effects for all groups. We conclude that the trade-off between lifespan and reproduction is mediated by the detrimental side-effects of protein ingestion on lifespan.