Host deprivation influence on response of Rhagoletis pomonella to its oviposition deterring pheromone

ABSTRACT. The response of host-depibved Rhagoletis pomonella Walsh (Diptera, Tephritidae) females to host fruit ( Crataegus viridis L.) marked with R.pomonella oviposition deterring pheromone was measured in the laboratory. A positive correlation was found between length of host deprivation (5, 10, 20, 40 or 80 min) and probability of host acceptance (i.e. oviposition) by flies. The results are discussed in relation to current theories on physiological control of insect feeding and oviposition behaviour.     

Oviposition deterring pheromone in Rhagoletis cerasi: Biological activity of a pure single compound verified in semi-field test

A pure single compound, a palmityl-glucopyranoside identified as component of the oviposition deterring pheromone (ODP) of the European cherry fruit fly was evaluated with respect to its biological activity by means of a semi-field bioassay in a large field-cage with host plants. The comparison of the observed behaviors of flies exposed to clean cherries, to cherries treated with a standard ODP solution and with the pure compound revealed that palmityl-glucopyranoside elicited the same response as did the crude ODP solution. Irritation indices are described that seem to be of use for the characterization of cherry fruit fly behavior in the presence and absence of ODP. The advantages of the field-cage test over the standard laboratory tests are discussed.

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Zusammenfassung Mit zunehmenden Reinigungsschritten am Rohextrakt des Markierungspheromones (ODP) der Kirschenfliege, Rhagoletis cerasi L., und zunehmenden Reinheitsgrad des aktiven Prinzipes nimmt die Menge an Substanz ab, welche sowohl für die chemische Analyse wie auch die Prüfung der biologischen Aktivität zur Verfügung steht. Beim Vorliegen einer nicht mehr wägbaren Menge eines reinen Stoffes im Sommer 1985, welcher als ein Palmityl-glucopyranosid identifiziert werden konnte (Hurter et al., 1987), musste deshalb auf den üblichen Verhaltenstest im Labor verzichtet und ein neuer Lösungsansatz gefunden werden. Nachdem ein erster elektrophysiologischer Test positive Resultate gezeight hatte, wurde die Fraktion Nr. 2634 in einem grossen Freilandkäfig mit Kirschenpflanzen und natürlichen Früchten in einem Verhaltenstest auf ihre biologische Wirksamkeit geprüft.Die in dieser Arbeit präsentierten Resultate zeigen nicht nur, dass mit dem Palmityl-glucopyranosid eine ähnliche Wirkung erzielt wurde wie mit natürlichen ODP, sondern zum erstenmal konnte demonstriert werden, dass in R. cerasi und möglicherweise in andern Fruchtfliegenarten eine einzelne chemische Substanz eine eiablagehemmende Wirkung ausüben kann.

     

The cost of reproduction in rosehip flies,Rhagoletis basiola: Eggs are time

Summary The hypothesis was tested that egg-laying by rosehip fruit flies,Rhagoletis basiola, would lead to reduced adult survivorship compared with oviposition-deprived flies, through experimental manipulation. Oviposition-deprived flies were assigned to two groups: mated and spinster. All three treatment groups were permitted to spend equal time and energy exploring and handling hosts but only the former group were allowed to deposit eggs (three/day). Results clearly showed that ovipositing flies suffered significantly higher mortality rates and that mated and spinster flies produced similar survivorship patterns. I consider these results both as a foraging problem (i.e. laying eggs in inferior hosts can cost both eggs and life) and as a more general life history problem.     

Oviposition deterring pheromone in Rhagoletis cerasi: Behavioral laboratory test to measure pheromone activity

Current investigations concerning the identification of the chemical nature of the oviposition deterring pheromone of the European cherry fruit fly, Rhagoletis cerasi L., are conducted in an interdisciplinary research program by combining chemistry with behavioral evaluation. The methods used to collect and evaluate the pheromone by an improved semi-quantitative behavior test are described.

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Zusammenfassung Nach einer einleitenden Übersicht über den heutigen Stand der Erkenntnisse auf dem Gebiet der eiablage-hemmenden Pheromone (ODP) bei Fruchtfliegen und insbesondere der Kirschenfliege wird der verbesserte Verhaltenstest beschrieben, welcher für den halb-quantitativen Nachweis von ODP-Aktivität entwickelt worden ist. Im laufenden interdisziplinären Forschungsprogramm an der Eidg. Forschungsanstalt Wädenswil wird versucht, durch enge Zusammenarbeit zwischen Biologe, Chemiker und Elektrophysiologe das ODP der Kirschenfliege möglichst mit geringem Anteil an Verunreinigung einzusammeln, zu reinigen und einer chemischen Identifikation der ODP Komponenten zugänglich zu machen.Die Möglichkeiten und Grenzen des Verhaltenstests werden diskutiert und die Empfindlichkeit des Verfahrens mittels einer Konzentrationsreihe von Pheromonextrakt aufgezeigt.

     

Prior experience affects the visual ability ofRhagoletis pomonella flies (Diptera: Tephritidae) to find host fruit

Females of the apple maggot fly,Rhagoletis pomonella, were allowed for 3 days to alight upon and oviposit in green or red 18- to 20-mm hawthorn host fruit (Crategus mollis) or green or red 45- to 55-mm apple host fruit (Malus pumila) hung from branches of potted host trees in field enclosures. Subsequently, when females were released individually on potted host trees harboring fruit of one of these types, their ability to find fruit of unfamiliar size proved unaffected by prior experience with fruit but their ability to find fruit of unfamiliar color was significantly affected. Specifically, females exposed to red hawthorns or red apples were less able to find green hawthorns or green apples than were females experienced with either of the latter fruit types. Fruit odor was found to have no effect on female ability to find familiar compared with unfamiliar green fruit. In contrast, a difference in size (or surface chemistry) between familiar and unfamiliar fruit but not a difference in fruit color had a significant negative influence on the propensity of alighting females to bore into unfamiliar fruit. Three bouts of experience with alighting upon and ovipositing into fruit over a period of about 1 h had no detectable effect on female ability to find unfamiliar fruit but did reduce propensity to bore into unfamiliar fruit. Our findings are discussed in relation to insect ability to learn visual and chemical stimuli of resources and insect propensity to form host races. We also discuss the potential impact of our findings on nonpesticidal, behavioral methods of managingR. pomonella in commercial apple orchards.     

Influence of protein hydrolysate baits on oviposition in the apple maggot, Rhagoletis pomonella

Ovipositional responses of apple maggot (AM), Rhagoletis pomonella (Walsh), females were studied in the laboratory on apples (var: Golden Delicious) treated with different rates of four protein hydrolysate baits in choice and no-choice tests. Protein hydrolysate baits at rates of 0.5 and 1% had no significant effect, but oviposition was greatly reduced at higher rates of 5 and 10%. Apple maggot females exposed to apples treated with protein hydrolysate baits at a rate of 10% made 41–71% fewer punctures and laid 41–73% fewer eggs than in untreated control. No oviposition activity was shown on apples treated with 25 and 100% Nulure®. In no-choice tests the AM females laid 75–96% fewer eggs in apples treated with 10 and 25% Nulure compared to controls and no oviposition occurred in apples treated with 100% Nulure. Apple maggot females arrived in similar numbers on apples treated with 10% Nulure and untreated apples, but only 5% of those arriving on Nulure-treated apples showed ovipositor boring with no egg deposition while 60% of females arriving on untreated apples showed ovipositor boring activity and laid an average of 2.5 eggs per apple. In another experiment, individual AM females displayed similar behavioral responses to 10% Nulure-treated apples; none of the 56 females tested on treated apples displayed ovipositor boring activity, but 59% of the females (N=56) tested on untreated apples displayed ovipositor boring within 5 min of their arrival. Ninetyeight percent of AM females stayed and fed on fruit surfaces for 5 min on Nulure-treated apples without ovipositor boring compared to only 2% on untreated apples. Of the females that arrived on untreated apples, 39% flew away within 5 min without ovipositor boring compared to only 2% of those that arrived on Nulure-treated apples. Results of these two behavioral experiments suggest that upon arrival on a protein bait-treated apple, an apparent change of behavior occurs in AM females and instead of attempting to oviposit, they attempt to feed on fruit surfaces resulting in reduced oviposition activity. These results indicate that the feeding and oviposition-related activities of AM females are probably mutually exclusive and that the feeding behavior preempts oviposition activities on host fruits treated with higher rates of protein hydrolysate baits.     

Apple maggot fly response to red sphere traps in relation to fly age and experience

Responses of the apple maggot fly,Rhagoletis pomonella (Walsh), to 8 cm red sticky sphere traps, baited with the synthetic fruit odor butyl hexanoate, were investigated in field-caged apple trees containing green or red Gravenstein apples. Trap capture rate and the probability of oviposition in apples before capture generally increased with female age and number of mature eggs in ovaries. Two days of pre-test exposure of mature females to red hawthorns, green Red Delicious apples or green Gravenstein apples had no significant effect on the likelihood of a fly finding a red sphere. However, before capture on a sphere or departing a tree, hawthorn-exposed females found significantly fewer apples and laid significantly fewer eggs than females exposed to Gravenstein apples. Variation in duration of pre-test exposure (1–4 days) of flies to Gravenstein apples had no detectable influence on female response to apples or to a red sphere in a test tree. The relevance of these findings to effectiveness of sphere traps, forR. pomonella control in commercial orchards is discussed.     

Realized heritability of behavioral responsiveness to an oviposition deterring pheromone in the azuki bean weevil Callosobruchus chinensis

Artificial selection was conducted to reduce the behavioral responsiveness of female bruchid beetles, Callosobruchus chinensis, to the oviposition deterring pheromone excreted by conspecific females. Significant responses to selection were observed after two generations of selection. Realized heritability was estimated as 0.052 ± 0.017 from cumulative responses regressed on cumulative selection differentials. These results indicate that this pheromone communication system has significant additive genetic variance needed for its evolution.     

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.     

Host-marking pheromone and use of previously established oviposition sites by the mediterranean fruit fly (Diptera: Tephritidae)

Under controlled conditions, the Mediterranean fruit fly, Ceratitis capitata(Wiedemann) preferred to initiate oviposition into preexisting, naturally formed oviposition punctures in a host fruit, kumquat (Fortunella japonica),over establishing new sites on the fruit. This preference was expressed despite the presence of naturally deposited host-marking pheromone (HMP)shown previously to deter oviposition. An almost-identical preference for existing punctures was expressed when females were presented with fruit bearing artificially made punctures on which HMP had been naturally deposited. Using artificial punctures and HMP extracts, the occurrence of punctures was manipulated independently of the presence of HMP.Under field-cage conditions, we found that (1) punctures stimulated egg-laying on kumquats, regardless of HMP treatment; (2) HMP extract inhibited egg-laying, regardless of the occurrence of punctures; and (3) the extent to which HMP inhibited egg-laying was greater on fruit free of punctures than on fruit bearing them. The physiological, evolutionary, and pest management implications of these results are discussed.     

On the evolutionary ecology of marking pheromones

Summary Many parasitic insects mark hosts with a pheromone after oviposition. The evolutionary ecology of such marking pheromones was studied to determine (i) under what ecological and behavioral conditions such pheromones could evolve and (ii) why so many of these marking pheromones are water-soluble and thus short-lived. We used a number of different techniques. First, the fitness values of individual normal (nonmarking) and mutant (marking) insects foraging for hosts were computed using dynamic state-variable models. Second, population level models were used to study when a population of non-marking individuals can be invaded by marking individuals. Third, behavior-rich simulations (developed originally for apple maggot,Rhagoletis pomonella) were used to test experimentally some of the hypotheses generated using the individual and population-level models. Finally, we developed a model for the benefit over time to an individual by marking. This model shows that when benefit is measured in terms of larval survival, nearly all of the benefit to a mother is obtained from short-lived marks. Genetical theories of pheromone evolution and the connection between our results and existing theories of altruistic behavior are discussed.     

Effects of pyriproxyfen on ovarian development and fecundity ofRhagoletis pomonella flies

Three-day old female apple maggot flies,Rhagoletis pomonella (Walsh), were topically exposed to different doses (0.1, 1, 10, and 100 μg per fly) of a juvenile hormone mimic, pyriproxyfen, in the laboratory. Pyriproxyfen had little lethal effect on females except at the extremely high dose of 100 μg. It also had no significant effect on egg viability of treated females. A non-lethal dose of 1 μg per fly did, however, enhance significantly the fecundity (egg production) as well as the ovarian development (number of eggs in ovaries and length of egg folicles) of treated flies. We conclude that pyriproxyfen could be a useful aid in exploring endocrine regulation of feeding and reproductive physiology behavior inR. pomonella, about which current knowledge is scant.     

Host marking behavior in phytophagous insects and parasitoids

Oviposition behavior in phytophagous insects and entomophagous parasitoids is often modified by the presence of conspecific brood (eggs and larvae). Often, females avoid laying eggs on or in hosts bearing brood, a behavior that acts to reduce the level of competition suffered by their offspring. Avoidance of occupied hosts is typically mediated by cues and/or signals associated with brood. In this article, we review the role of Marking Pheromones (MPs) as signals of brood presence in both phytophagous and entomophagous insects. We place information about the function and evolution of MPs in the context of recent theory in the field of animal communication. We highlight the dynamics of host-marking systems and discuss how effects of MPs vary according to factors such as female experience and egg load. We also examine variation in the form and function of MP communication across a variety of insect taxa. While studies of MP communication in phytophagous insects have focused on the underlying behavioral mechanisms and chemistry of MP communication, studies in entomophagous insects have focused on the functional aspects of MPs and their role in decision-making in insects. We argue that an approach that incorporates the important contributions of both of these somewhat independent, but complementary areas of research will lead to a more complete understanding of MPs in insects. Finally, we suggest that MP systems are model systems for the study of animal signaling and its evolution.     

Recognition and avoidance of previously laid egg masses by the oblique-banded leafroller (Lepidoptera: Tortricidae)

Investigations were conducted on oviposition by laboratory-reared oblique-banded leafrollers, Choristoneura rosaceana Harris. In paperbag oviposition chambers, females deposited their egg masses in a significantly nonrandom, uniform manner in 16 of 20 replicates. Inpetri dish oviposition chambers, gravid females were exposed to egg masses recently deposited by other females. As determined by the magnitude of the mean direction vector of the second egg mass in relation to the first, females detected and avoided egg masses laid by other females. In addition, they avoided waxed paper from which a recently oviposited egg mass had been removed. This phenomenon is most likely due to a pheromone laid down with each egg mass.     

Multiple Mating and Reproductive Success of Male and Female Apple Maggot Flies, Rhagoletis pomonella (Diptera: Tephritidae)

Mature male and female apple maggot flies mated frequently on a field-caged host tree during a 14-day study. Each sex averaged one mating per day (mean of 1.0 ± 0.1), but some females mated up to eight times per day and some males up to six times per day. Reproductive success was estimated based both on observed numbers of matings in the field cage and on previous work relating fecundity and fertility to female mating status. Male and female flies did not differ in mean or variance in reproductive success, indicating that this is a polygamous mating system consisting of both polygyny and polyandry. We discuss the significance of this with regard to the framework of insect mating control and frequency proposed by others. We also discuss behaviors (such as movements, agonistic encounters, occurrences on fruit, and ovipositions) that do and do not show correlations with mating success in apple maggot flies.