A field test for host fruit odour discrimination and avoidance behaviour for Rhagoletis pomonella flies in the western United States

Prezygotic isolation due to habitat choice is important to many models of speciation-with-gene-flow. Habitat choice is usually thought to occur through positive preferences of organisms for particular environments. However, avoidance of non-natal environments may also play a role in choice and have repercussions for post-zygotic isolation that preference does not. The recent host shift of Rhagoletis pomonella (Diptera: Tephritidae) from downy hawthorn, Crataegus mollis, to introduced apple, Malus domestica, in the eastern United States is a model for speciation-with-gene-flow. However, the fly is also present in the western United States where it was likely introduced via infested apples ≤ 60 years ago. R. pomonella now attacks two additional hawthorns in the west, the native C. douglasii (black hawthorn) and the introduced C. monogyna (English ornamental hawthorn). Flight tunnel tests have shown that western apple-, C. douglasii- and C. monogyna-origin flies all positively orient to fruit volatile blends of their respective natal hosts in flight tunnel assays. Here, we show that these laboratory differences translate to nature through field-trapping studies of flies in the state of Washington. Moreover, western R. pomonella display both positive orientation to their respective natal fruit volatiles and avoidance behaviour (negative orientation) to non-natal volatiles. Our results are consistent with the existence of behaviourally differentiated host races of R. pomonella in the west. In addition, the rapid evolution of avoidance behaviour appears to be a general phenomenon for R. pomonella during host shifts, as the eastern apple and downy hawthorn host races also are antagonized by non-natal fruit volatiles.     

Odour movement, wind direction, and the problem of host-finding by tsetse flies

ABSTRACT. Movement of host odour was modelled in natural tsetse habitats with smoke and ultra-light 7-cm-long wind vanes; the speed and direction of the air movements were analysed from video recordings thereof. Wind of <1 ms^-1 did not move in straight lines, since large packets of air (>10 m across) often changed direction together. The rate of this change of direction (meander) correlated negatively with windspeed. In open woodland with a shrubby understorey (in which windspeed was reduced by a factor of >5 from that above the canopy, to ax 0.3 m s^-1), this wind meander fell by 2d? s^-1 change of direction for each 0.1 m s^-1 increase in windspeed (r²=0.96). Over open ground without shrub cover, the meander fell by 0.5d? s^-1 per 0.1 m s^-1 increase in windspeed (r²=0.85). In both situations, such meandering virtually ceased in winds of > 1 m s^-1. In woodland, the relationship between the direction of air movement near the surface of bare earth (one potential tsetse landing site) and that c. 0.5 m above ground level (flight height) was often weak (r²=0.2-0.4), but this problem would be reduced if the fly averaged the ground-level wind for at least 30 s. Odour (smoke) travelling from a source 15 m ‘upwind’ over open ground arrived at a notional tsetse fly for 80% of the time from a direction within 10d? of the true source direction. In typical tsetse woodland, however, the ‘odour’ arrived from all directions (including >90d? away from the source), with only a 30% bias towards the true source direction (±10d?). Evidently, tsetse must navigate up odour plumes by means that get round these difficulties-simple, moth-type upwind anemotaxis alone seems unlikely to be adequate.     

Odour learning and decision-making during food collection in the leaf-cutting antAcromyrmex lundi

Summary Recruited workers of the leaf-cutting antAcromyrmex lundi learn the odour of the food fragment initially carried by a successful scout, and use this cue as a decision criterion during food collection. Citral, a natural deterrent odour, can also be associated as a food signal, showing how powerful olfactory learning is in this species. The role of both food-odour learning and a previously reported leaf-marking pheromone during diet selection by leaf-cutting ants is discussed.     

Learning affects mate choice in female fruit flies

Learning in the context of mate choice can influence sexualselection and speciation. Relatively little work, however, hasbeen conducted on the role of learning in the context of matechoice, and this topic has been mostly ignored in insects eventhough insects have served as a prime model system in researchon sexual selection and incipient speciation. Extending recentwork indicating apparently adaptive learning in the contextof sexual behavior by male fruit flies (Drosophila melanogaster),I tested for the effect of learning on mate choice by femalefruit flies. Compared to young virgin females that experiencedcourtship by large males, young virgin females that experiencedcourtship by small males were more likely to mate with smalland large males in a test conducted a day after the experiencephase. These results, which are the first clear empirical demonstrationof learning in the context of mate choice by female insects,lay the foundation for research on the role of learning in insectsexual selection and speciation.     

Conditioning odor-shock associations in the black blowfly,Phormia regina

Discriminative classical conditioning of an olfactory avoidance response was demonstrated in the blowfly, Phormia regina.Learning indices were calculated as the fraction of flies avoiding the CS+ (conditioned stimulus paired with electric shock reinforcement) minus the fraction of flies avoiding the CS-(conditioned stimulus not paired with electric shock), averaged over two different groups of flies, in which reciprocal odors were used as the CS+. Avoidance responses to both odors presented simultaneously at a T-maze choice point yielded mean learning indices of zero for naive flies or pseudoconditioned (shock alone) or sensitized (odors alone) controls. In contrast, pairing an odor with electric shock produced a mean learning index significantly greater than zero. These results are similar to those reported for Drosophila melanogaster.Blow flies, however, showed much lower levels of associative learning than fruit flies.     

Male fruit flies learn to avoid interspecific courtship

Experimental data suggest, and theoretical models typicallyassume, that males of many fruit flies (Drosophila spp) areat least partially indiscriminate while searching for mates,and that it is mostly the females who exert selective mate choice,which can lead to incipient speciation. Evidence on learningby male D. melanogaster in the context of courtship, however,raises the possibility that the initially indiscriminate malesbecome more selective with experience. I tested this possibilityby comparing the courtship behavior of male D. melanogasterexperienced at courting females of the closely related species,D. simulans, and inexperienced males. I found that comparedwith the inexperienced males, the males experienced with courtingD. simulans females showed significantly lower courtship towardfemale D. simulans. Both male treatments, however, showed virtuallyidentical courtship durations with female D. melanogaster. Theseresults indicate that male fruit flies adaptively refine theircourtship behavior with experience and suggest that the malescontribute more to assortative mating and incipient speciationthan is commonly assumed.     

An effective trapping method for field studies of opiine braconid parasitoids of tephritid fruit flies

A simple but effective trap for conducting field studies with opiine braconid parasitoids of fruit fly larvae was designed from yellow plastic balls containing an internal odor source and coated on the exterior with Tanglefoot. Tests of the trap in large field cages showed that it was simpler to use, less expensive, and more effective than previously reported designs.      

实蝇共生菌研究进展

实蝇类昆虫(双翅目Diptera:实蝇科Tephritidae)是重要的农业害虫,为害农作物茎秆和水果,严重影响农业生产和果蔬贸易,被许多国家列为重要的检疫性有害生物。实蝇共生菌对宿主实蝇的取食、生殖、发育及环境适应能力具有重要作用。目前,已有21个属的共生细菌(Enterobacter、Klebsiella、Citrobacter、Pseudomonas、Providencia、Erwinia、Acetobacter、Serratia、Proteus、Hafnia、Cedecea、Arthrobacter、Lactobacillus、Micrococcus、Streptococcus、Staphylococcus、Vibrio、Hafnia、Deinococcacea、Bacillus、Wolba-chia)以及1个属的共生真菌(Candida)被鉴定。其中,肺炎杆菌Klebsiella pneumoniae、产酸克雷伯氏菌Klebsiella oxytoca、成团泛菌Pantoea agglomerans、费氏柠檬酸杆菌Citrobter freundii、阴沟肠杆菌Enterobacter cloaca...     

Remating inhibition in female Queensland fruit flies: effects and correlates of sperm storage

Reproductive success of male insects commonly hinges both on their ability to secure copulations with many mates and also on their ability to inseminate and inhibit subsequent sexual receptivity of their mates to rival males. We here present the first investigation of sperm storage in Queensland fruit flies (Tephritidae: Bactrocera tryoni; a.k.a. 'Q-flies') and address the question of whether remating inhibition in females is directly influenced by or correlated with number of sperm stored from their first mates. We used irradiation to disrupt spermatogenesis and thereby experimentally reduce the number of sperm stored by some male's mates while leaving other aspects of male sexual performance (mating probability, latency until copulating, copula duration) unaffected. Females that mated with irradiated rather than normal males were less likely to store any sperm at all (50% vs. 89%) and, if some sperm were stored, the number was greatly reduced (median 11 vs. 120). Despite the considerable differences in sperm storage, females mated by normal males and irradiated males were similarly likely to remate at the next opportunity, indicating (1) number of sperm stored does not directly drive female remating inhibition and (2) factors actually responsible for remating inhibition are similarly expressed in normal and irradiated males. While overall levels of remating were similar for mates of normal and irradiated males, factors responsible for female remating inhibition were positively associated with presence and number of sperm stored by mates of normal but not irradiated males. We suggest seminal fluids as the most likely factor responsible for remating inhibition in female Q-flies, as these are likely to be transported in proportion to number of sperm in normal males, be uninfluenced by irradiation, and be transported without systematic relation to sperm number in irradiated males.     

Associative learning of complex odours in parasitoid host location

In this paper we address the question how hymenopteran parasitoids deal with complex odour bouquets, using Microplitis croceipes (Cresson) (Hymenoptera: Braconidae) as a model. We examined the capacity of Microplitis croceipes to respond to individual compounds in flight chamber experiments after conditioning parasitoids with a mixture consisting of 2-octanone, methyl jasmonate and beta-caryophyllene. Parasitoids were given a choice between single compounds from the training mixture and beta-ocimene as an unrewarded alternative. When compared with control individuals lacking experience with the odour mixture, parasitoids trained to the odour blend showed an increased preference for 2-octanone and beta-caryophyllene, but not for methyl jasmonate. However, when trained with methyl jasmonate alone, parasitoids were able to respond to this compound. This demonstrates that parasitoids can learn to respond to individual compounds following experience with an odour mixture. However, for certain compounds of a mixture, learning can be blocked by other mixture components. Further experiments in which parasitoids were conditioned and challenged with two compound mixes confirmed that the olfactory background can affect recognition of individual compounds.     

Learning decreases heterospecific courtship and mating in fruit flies

Recent theory and data suggest that adaptive use of learning in the context of sexual behaviour could contribute to assortative mating. Experiments examining this issue indicated that male Drosophila persimilis that experienced courtship and rejection by heterospecific females exhibited significantly lower levels of heterospecific courtship and mating compared with those of inexperienced males. These results indicate that experience in the context of sexual behaviour in fruit flies could reduce gene flow between diverging populations, which may contribute to incipient speciation.     

Learning of apple fruit biotypes by apple maggot flies

Previously, we showed that after a female apple maggot fly, Rhagoletis pomonella,arrives on a host hawthorn or apple fruit, its propensity to accept (bore into) or reject that fruit prior to egg deposition can be modified by previous ovipositional experience with one or the other species and, hence, involves learning. Here, we present both field and laboratory evidence indicating that females also are able to learn characteristics of three different apple biotypes or cultivars: Early Macintosh, Red Delicious, and Golden Delicious. We suspect that females learn to discriminate among these three cultivars on the basis of differences in chemical stimuli among cultivars. The effect of fruit cultivar learning was not as strong as the effect of fruit species learning.     

Odour signals major histocompatibility complex genotype in an Old World monkey

The major histocompatibility complex (MHC) is an extraordinarily diverse cluster of genes that play a key role in the immune system. MHC gene products are also found in various body secretions, leading to the suggestion that MHC genotypes are linked to unique individual odourtypes that animals use to assess the suitability of other individuals as potential mates or social partners. We investigated the relationship between chemical odour profiles and genotype in a large, naturally reproducing population of mandrills, using gas chromatography–mass spectrometry and MHC genotyping. Odour profiles were not linked to the possession of particular MHC supertypes. Sex influenced some measures of odour diversity and dominance rank influenced some measures of odour diversity in males, but not in females. Odour similarity was strongly related to similarity at the MHC, and, in some cases, to pedigree relatedness. Our results suggest that odour provides both a cue of individual genetic quality and information against which the receiver can compare its own genotype to assess genetic similarity. These findings provide a potential mechanism underlying mate choice for genetic diversity and MHC similarity as well as kin selection.     

Exposure to stable flies reduces spatial learning in mice: involvement of endogenous opioid systems

Abstract. Biting flies influence both the physiology and behaviour of domestic and wild animals. This study demonstrates that relatively brief (60min) exposure to stable flies, Stomoxys calcitrans (L.), affects the spatial abilities of male mice. Stable fly exposure resulted in poorer subsequent performance in a water maze task in which individual mice had to learn the spatial location of a submerged hidden platform using extramaze visual cues. Determinations of spatial acquisition and retention were made with mice that had been previously exposed for 60min to either stable flies or house flies, Musca domestica (L.). Mice exposed to stable flies displayed over one day of testing (six blocks or sets of four trials) significantly poorer acquisition and retention of the water maze task than either mice that had been exposed to house flies or fly-naive mice. This attenuation of spatial learning occurred in the absence of any evident sensorimotor or motivational impairments. The reduction in spatial abilites involved endogenous opioid systems, as the decreased performance resulting from stable fly exposure was blocked by pre-treatment with the prototypic opiate antagonist, naltrexone. These results indicate that relatively brief exposure to biting flies can lead to a decrease in spatial abilities which is associated with enhanced endogenous opioid activity. These results support the involvement of endogenous opioid systems in the mediation of the behavioural and physiological effects of biting fly exposure. They further suggest that decreases in spatial abilities and performance may be part of the behavioural consequences of biting fly exposure in domestic and wild animals.     

Behavioural switch of female Mediterranean fruit fly, Ceratitis capitata: mating and oviposition activity in outdoor field cages in Hawaii

1 Laboratory-reared normal, and wild female Mediterranean fruit flies, Ceratitis capitata (Wiedemann), were assayed in outdoor field cages to assess the impact of a mating-induced behavioural switch on mating and subsequent oviposition activity. 2 Virgin females preferred interactions with males leading to mating over attraction to, and oviposition in, artificial yellow spheres containing guava odour or green apples hung in a guava tree. Laboratory-reared females previously mated with either laboratory-reared normal males or laboratory-reared irradiated (sterile) males showed little interest in remating with males and instead, were much more likely to be found arrested on artificial and real fruit and ovipositing. Oviposition on artificial fruit was five times greater by females that had mated with either normal or irradiated males than by virgin females. Wild females showed similar qualitative changes in the mating-induced behavioural switch; however, oviposition activity was significantly less than for laboratory-reared females. 3 These results confirm that mating has a profound effect on the behaviour of female Mediterranean fruit flies and that irradiated males are functionally equal with normal males (lab-reared or wild) in their ability to alter female behaviour. These results are discussed in the context of the sterile insect technique for control of Mediterranean fruit flies in the field.     

Orientation of tsetse flies to wind, within and outside host odour plumes in the field

Abstract Free-flying wild tsetse flies ( Glossina pallidipes Aust. and G. m. morsitans Westw.) were video recorded in Zimbabwe as they flew within an artificial host odour plume at 3, 7 or 15 m from the source, or in no odour, with and without a 0.75 m² vertical, black visual target present aligned with the wind. With no visual target present, flights in odour were strongly biased upwind, and in the absence of odour strongly biased downwind. With the target present, between 16% and 40% of the upwind approaching flies responded visually as they passed the target, by circling it, in proportion to the proximity of the source (taken to be proportional to the mean odour concentration). Crosswind approaching flies (for whom the target will have been visible for some metres away) circled more frequently (34–56%), but without obvious correlation with the odour concentration. Circling flies also responded orthokinetically, by slowing down as they passed the target. The departure directions relative to the wind of flies leaving the target were significantly affected by the odour concentration. At 3 m they left the target in all directions, except possibly avoiding due upwind. At 7 m they left with an obliquely upwind bias, but at 15 m and also in no odour, they left with a strong crosswind bias.