Sylvia S. Cooley

Associative learning in egglaying site selection by apple maggot flies.

Evidence is presented demonstrating that associative learning during oviposition in Crataegus or apple hosts can significantly influence the propensity of apple maggot flies to accept or reject these hosts in future encounters. The data suggest that within resource patches of a given host type there may be an enhancement of foraging efficiency.

Behavioral evidence for host races in Rhagoletis pomonella flies

SummaryOne of the most controversial putative cases of host race formation in insects is that of the apple maggot fly, Rhagoletis pomonella (Diptera: Tephritidae). A principal cause of the controversy is lack of relevant data. In laboratory and field enclosure experiments, we compared the host acceptance behavior of sympatric populations of flies originating from naturally infested hawthorn (the native host) and apple (an introduced host) in Amherst, Massachusetts or East Lansing, Michigan. In general, hawthorn fruit were accepted for ovipositional attempts nearly equally by apple and hawthorn origin females, whereas apples were accepted much more often by apple than hawthorn origin females. Similarly, males of apple and hawthorn origin exhibited about equal duration of residence on hawthorn fruits as sites at which to acquire potential mates, while males of apple origin tended to reside substantially longer than males of hawthorn origin on apples. Irrespective of fly origin, both sexes always responded more positively to hawthorn fruit than to apples. Because all flies assayed were naive (ruling out effects of prior host experience of adults) and because tests revealed no influence of pre-imaginal fruit exposure on pattern of host fruit acceptance by females, the combined evidence suggests the phenotypic differences we observed in host response pattern between hawthorn and apple origin flies may have an underlying genetic basis. Further tests showed that while larval progeny of flies of each origin survived better in naturally growing hawthorn fruit than in naturally growing apples, there was no differential effect of fly origin on larval survival ability in either host. We discuss our findings in relation to restriction in gene flow between sympatric populations of R. pomonella and in relation to current models of host shifts in insects.

On the nature of learning in oviposition site acceptance by apple maggot flies

Abstract In a previous study, we found that after a female apple maggot fly, Rhagoletis pomonella , had arrived on a host fruit, its propensity to accept (bore into) or reject that fruit prior to egg deposition was modifiable through experience, and hence involved learning. Here, we aimed to determine whether the true nature of the learned response was either (a) one in which a conditioned female, as a result of having become familiar with a particular fruit type, formed a greater propensity to accept that type in a future encounter than had a naive female, or (b) one in which a conditioned female formed no greater propensity to accept a particular fruit type but simply was less prone than a naive female to accept a novel fruit type. The results of two experiments supported the latter mechanism, while the results of a third experiment provided evidence for both mechanisms. The possible adaptive significance of learning to refuse a novel resource type is discussed.

Learning in oviposition site selection by Ceratitis capitata flies

Significant differences in propensity to attempt oviposition into various types of fruit were found among Mediterranean fruit flies, Ceratitis capitata, from the same wild population. Evidence presented suggests repeated exposure to one host fruit type conditions females to that host, and influences female response to other hosts in future encounters. Reversibility of conditioning was also demonstrated. Learning to refuse a novel host type is indicated as being more important than learning to accept a familiar host type in medfly conditioning to hosts.

Post-feeding bubbling behaviour in fluid-feeding Diptera: concentration of crop contents by oral evaporation of excess water

Abstract During long post‐feeding periods, adult tephritid flies (and apparently other non‐blood feeding Diptera and even Hymenoptera) engage in behaviour consisting of oral extrusion and aeration of liquid droplets of varying size (‘bubbling’), proboscis pumping, and occasional deposition of regurgitate on the substrate that is followed by subsequent re‐ingestion. Unlike evaporative cooling, which occurs in some insects when heat‐stressed, these regurgitation behaviours take place regularly at moderate temperatures. To our knowledge, the significance of these behaviours has not been studied. We suggest that through bubbling behaviour, fully gorged Rhagoletis pomonella (Walsh) (Diptera: Tephritidae) flies eliminate excess water by evaporation to concentrate nutrients suspended in dilute solution, and release crop volume to allow resumption of fluid feeding or minimize the water load while foraging for other resources. Fly weights were measured continuously during pre‐ and post‐feeding periods and in relation to occurrence of regurgitation behaviours. Fly weight losses during pre‐feeding were an order of magnitude lower than post‐feeding weight losses when flies exposed regurgitated liquid crop contents to air. During a bout of droplet extrusions, lasting on average 23 min, weight loss averaged 66% of the weight of liquid ingested by a fly in the preceding meal. Fly weight loss while bubbling was significantly correlated with duration of bubbling, temperature and relative humidity during post‐feeding, and initial fly weight (adj. R2 = 0.95). Fly age, volume of liquid ingested and rate of pre‐feeding weight loss did not significantly improve predicted weight loss through bubbling. Findings are discussed in relation to other behaviours adopted by fluid‐feeding insects to eliminate excess water.

Bioassay approaches to assessing behavioral responses of plum curculio adults (Coleoptera: Curculionidae) to host fruit odor.

We evaluated several approaches to developing a simple, sensitive, and reliable laboratory bioassay of responses of overwintered adult plum curculios (PCs),Conotrachelus nenuphar (Herbst), to host fruit odor or its attractive components. A high proportion of assayed PCs responded positively to odor of wild plums under no-choice, moving-air conditions in a wind tunnel and under dual-choice, still-air conditions in enclosed Petri dishes. Positive response to controls lacking host odor, however, was much greater in the wind tunnel, arguing in favor of bioassays under dual-choice conditions in still air to provide greater PC discrimination. Response to host odor (from wild plums or hexane extract of wild plums or Liberty apples) in Petri dish bioassay chambers proved greatest: (1) during the scotophase of PCs under total dark or dim red light conditions, (2) when Petri dishes were completely enclosed, (3) when PCs were starved for 24 or 48 hr, and (4) when PCs were tested within seven weeks after apple tree petal fall. Neither the sex of a PC nor the direction in which a PC was obliged to move to find the source of host odor (upward through a port in the Petri dish lid or downward through a port in the base) had a substantial effect on level of response to host odor or discrimination of host odor from a nonodorous control. We conclude that an enclosed Petri dish bioassay chamber of the type described here should be a valuable asset in the process of chemically identifying components of host fruit odor attractive to PCs.

How well can relative specialistRhagoletis flies learn to discriminate fruit for oviposition

We show that blueberry maggot females [Rhagoletis mendax(Curran)], apple maggot females [Rhagoletis pomonella(Walsh)], and walnut husk fly females [Rhagoletis suavis(Loew)], all relative specialists in range of fruit species attacked in nature, are able to learn to discriminate between types of fruit in which they oviposit. It appears, however, that these relative specialists express less capacity to learn fruit characters than relative generalist tephritids. This difference in expression of learning ability may be associated in part with differences in assignments species are asked to learn. Apparent differences in learning capability between relative specialist and relative generalist tephritids may therefore depend as much upon differences in the physical and chemical nature of host fruit as upon species differences in the adaptive value of learning.

Prior experience influences the fruit residence of male apple maggot flies,Rhagoletis pomonella

Male apple maggot flies spend considerable time residing on individual host fruit as territories on which they force-copulate arriving females in search of oviposition sites. Here, we present evidence from investigations in nature and the laboratory that shows the propensity of males to reside on a hawthorn or apple fruit as a territory is significantly modifiable through prior experience with fruit and, hence, involves learning. Previous studies revealed that after a female apple maggot fly, Rhagoletis pomonella, arrived on a host hawthorn or apple fruit, its propensity to accept or reject that fruit for egg-laying was similarly modifiable through prior fruit-exposure experience and also involved learning. We discuss how host fruit learning in males and females, in concert with genetic-based differences in host fruit residence and acceptance behavior between populations of flies originating from hawthorn and apple, could give rise to a reduction in gene flow between populations of flies on these two host types.