Learning to find fruit in Ceratitis capitata flies

Wild Mediterranean fruit fly females, Ceratitis capitata (Wiedemann), from an essentially monophagous population on the island of Hawaii were exposed to natural mock orange (Murraya paniculata) or sweet orange (Citrus sinensis) host fruit hung from branches of potted trees for 3-day periods in field enclosures. Subsequently, when flies were released individually onto potted trees harboring one or the other (or a mixture) of these fruit types, a higher proportion visited the type of fruit with which they were familiar (and visitors found familiar fruit faster) compared with the fruit type with which they were unfamiliar. Moreover, fruit-finding flies of this monophagous population attempted oviposition exclusively in the familiar fruit type, and thus appeared to be just as capable of learning to accept fruit for oviposition as wild flies from a previously-tested polyphagous population on the island of Maui. Additional tests were conducted in which flies were exposed to natural or colored-wax-covered mock oranges or sweet oranges and tested for response to colored-wax-covered natural or artificial fruit. Results suggested that fruit size was the principal character learned and used in finding mock orange or sweet orange fruit, while fruit color and odor appeared to be of little or no importance in this regard.

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Résumé Des femelles sauvages d'une population essentiellement monophage de C. capitata Wiedemann, provenant de l'île de Hawaï, ont été mises en présence pendant des périodes de 3 jours dans des enceintes dans la nature à des fruits de Murraya paniculata et de Citrus sinensis suspendus à des branches d'arbres empotés. Quand les mouches ont été libérées individuellement sur les arbres empotés portant l'un ou l'autre de ces fruits (ou leur mélange), une plus forte proportion a visité le fruit avec lequel elles étaient familiarisées (et l'ont trouvé plus vite) que le fruit avec lequel elles ne n'étaient pas. De plus, les femelles découvrant des fruits de cette population monophage ne tentèrent de pondre que dans le type de fruit avec lequel elle étaient familiarisées. Elles se montrèrent aussi capables que les mouches d'une population polyphage de l'île de Maui d'apprendre à accepter de nouveaux fruits pour pondre. Des expériences complémentaires ont été réalisées dans lesquelles les mouches étaient mises en présence de fruits de M. paniculata ou de C. sinensis naturels ou couverts de cire colorée ou encore de fruits artificiels. Les résultats suggèrent que la taille du fruit est le principal critère d'apprentissage utilisé pour trouver M. paniculata ou C. sinensis, la couleur et l'odeur du fruit étant apparus comme de moindre importance ou sans effet.

     

Life history response of female Mediterranean fruit flies, Ceratitis capitata, to periods of host deprivation

Female medflies were subjected to four different patterns of host deprivation at each of three levels for a 24-day period. Treatments ranged from host absence 8 days post-eclosion to host absence 2 out of 3 days. Survival was recorded daily and egg production was recorded on the days in which hosts were present.The direct effect of host deprivation is to deny females the opportunity to oviposit. This disrupts their reproductive cycle and reduces their overall reproductive effort. Reduced reproductive effort at young ages increases survival (up to 2-fold) and daily reproduction (up to 8-fold) at older ages. Therefore, the long term effect of host deprivation is to postpone senescence.The implications of these findings regarding medfly sensescence are discussed in relation to: (i) reduction in population extinction rates during periods of host scarcity and (ii) egging strategies in medfly mass-rearing.

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Résumé Les femelles de C. capitata ont été privées d'hôtes suivant 4 modalités pendant 3 durées (33, 50 et 67% de leur vie) par période de 24 heures. Les modalités se répartissaient de la privation pendant 8 jours après l'éclosion jusqu'à 1 jour sur 3. La survie a été notée chaque jour et la ponte pour tous les jours où il y avait des hôtes.L'effect direct de l'absence d'hôte est d'empêcher la femelle de pondre. Ceci interrompt leur cycle de ponte et réduit leur activité reproductrice totale. Un effort reproductif réduit des femelles jeunes augmente la survie (jusqu'à 2 fois) et la ponte quotidienne (jusqu'à 8 fois) chez les femelles plus âgées. Ainsi, l'effet à long terme de la privation d'hôtes est d'ajourner la sénescence.Les implications de ces observations sur la sénescence de C. capitata sont discutées quant à: (1) la réduction des taux d'extinction des populations pendant les périodes de rareté d'hôtes; et: (2) les stratégies d'obtention de pontes dan les élevages de masse de C. capitata.

     

Function of multiple sperm storage organs in female Mediterranean fruit flies (Ceratitis capitata, Diptera: Tephritidae)

Sperm storage organs allow females to temporally separate insemination from fertilization, manipulate ejaculates and control fertilization. In the reproductive tract of female fruit flies (Diptera: Tephritidae), sperm are found in two different organs--a pair or triplet of spermathecae, and a "fertilization chamber". In order to understand the specific function of each of these organs, we tested the following hypotheses: (1) Sperm are distributed equally amongst the various sperm storage organs; (2) Both organ types maintain sperm viability; and (3) Sperm used in fertilization come from the fertilization chamber. We counted sperm in spermathecae and fertilization chamber of Mediterranean fruit flies (Ceratitis capitata) every 3 days for 18 days following insemination, and used a live/dead staining technique to determine the viability of sperm in these organs. Finally, by extirpating spermathecae from inseminated females and allowing them to oviposit, we were able to identify the fertilization chamber as the source of fertilizing sperm. Numbers of sperm in the spermathecae declined from an average of 3575 on the day of copulation to 649, 18 days later. Conversely, the fertilization chamber maintained a fairly constant level of sperms, ranging between an average of 207 cells on day 3 to 115 sperms on day 18. Throughout the period we monitored, we found high levels of sperm viability in both organs (> 80%). Sperm viability was similarly high in the fertilization chambers of females without spermathecae. However, fertility of eggs laid by these females declined rapidly, as did the number of sperm in the fertilization chamber. We conclude that both the spermathecae and the fertilization chamber are active sperm storage organs, with separate functions: the spermathecae for long-term storage and the fertilization chamber, periodically filled by the spermathecae, a staging point for fertilizing sperm. We suggest that the use of both organs by females results in sperm economy, which adaptively prolongs the intervals between copulations.     

Molecular sexing in the Mediterranean fruit fly, Ceratitis capitata

Molecular methods have been devised for sexing Mediterranean fruit fly (medfly) individuals using minimal amounts of material from any stage of the life cycle. Molecular sexing methods are particularly valuable when material is obtained from pre-adult stages and sex identification based on morphological characters is not possible. These methods may also be useful for adult stage material in situations where only limited amounts or poorly preserved specimens are available. The sexing methods described here use the polymerase chain reaction (PCR) to amplify sequences known to originate from the sex chromosomes of this species. One method co-amplifies homologous regions of the ITS1 ribosomal DNA from both the X and Y chromosomes. Males and females are distinguished based on the restriction fragment pattern produced after digestion of the PCR products with the restriction enzyme ApoI. A second method identifies males based on the positive amplification of a repetitive DNA sequence originating from the Y chromosome. Both methods are shown to be capable of establishing the sex identity of individuals using only minimal amounts of material from any stage of the life cycle.     

Host-specific demographic studies of the Mediterranean fruit fly Ceratitis capitata

Abstract. 1. This study was conducted in Greece using a wild strain of medflies ( Ceratitis capitata [Weidemann]) to assess age-specific vital rates in adults, and development and survival in pre-adults when reared on different host fruit. The results were used to construct life tables.
2. The demographic analyses suggested that there are basically four aspects of the medfly's life history which are of major importance: (i) multiple, highlyoverlapping generations; (ii) high net reproduction while young; (iii) high larval fitness in certain hosts; (iv) lack of diapause.
3. Reasons why these characteristics are felt to be important are briefly discussed.     

Effects of male sterility on female remating in the mediterranean fruitfly, Ceratitis capitata

Mating-induced reductions in female receptivity are common in insects. These responses are of interest because of their utility in insect pest control. In addition, the control of receptivity is likely to be the subject of sexual conflict over remating frequency. We investigated the specific effect of male sterility on female receptivity in an important pest species, the Mediterranean fruitfly (medfly), in which sterile males are often used for population suppression. Sterile males performed less courtship, obtained significantly fewer first and second matings than fertile males, and reduced female receptivity significantly less effectively than did fertile males. We modelled the likelihood of fertile matings and show that the low mating success of sterile males represents a significant problem for medfly sterile insect technique (SIT) programmes.     

Adhesive powder uptake and transfer by Mediterranean fruit flies, Ceratitis capitata (Dipt., Tephritidae)

Abstract:  Entostat^TM is an electrostatically charged wax powder that is used as a carrier particle in novel delivery systems for contaminating target insect pests with insecticides, biologicals or pheromones. Here, the adhesion of two forms of Entostat to the Mediterranean fruit fly (medfly) Ceratitis capitata (Wiedemann) was examined, and the adhesion of Entostat to live and dead medflies was compared. From controlled contaminations of medflies, it was shown that live medflies acquired larger quantities of Entostat than dead medflies, which could be due to the electrostatic charge shown to be carried by live insects. Air-milled Entostat (7.59  μ m mean diameter) adhered in larger quantities to medflies than pestle and mortar-ground Entostat (9.17  μ m mean diameter). Exposing medflies to different quantities of Entostat affected the initially adhering quantity but did not alter the proportion of powder retained over time. Medfly males contaminated with air-milled Entostat were shown to transfer small quantities to females during mating. This documentation of secondary powder transfer underscores the potential for using slow-acting killing agents on the basis of this delivery system.     

Responses of male Mediterranean fruit flies, Ceratitis capitata, to trimediure in a wind tunnel of novel design

ABSTRACT. The responses of male Ceratitis capitata (Wiedemann) (Diptera; Tephritidae) to trimediure were investigated in a wind tunnel of novel design with disposable plastic walls. Trimediure was presented to the flies either in the form of a discrete 'plume' or uniformly dispersed in the air passing through the tunnel. The flies showed an increase in flight activity in response to the two forms of stimulation and performed a zig-zag anemotactic flight in response to the plume, reminiscent of the behaviour of moths responding to a sex pheromone. They performed a straight, upwind, anemotactic response to the uniformly dispersed odour. The design of wind tunnel used in these tests was found to have several advantages: air flow and turbulence can be easily varied and the plastic walls can be changed to eliminate contamination problems.     

Two-way selection for mating activity in the Mediterranean fruit fly, Ceratitis capitata

No significant variation in mating activity was observed among eight laboratory strains of the Mediterranean fruit fly (medfly), Ceratitis capitata (Wiedemann) (Diptera: Tephritidae). Two methods were used to select strains showing high and low mating activity: a single pair technique (SP) assessing time to mating and a mass technique (M) based on a mating index. Reciprocal pairings between fast and slow selected lines showed that the difference between the SP lines was female-determined whereas the difference between the M lines depended on the behaviour of both sexes. M selection, irrespective of its direction, affected two courtship parameters, vibration distance and vibration duration both of which tended to be shorter. M selection was also associated with a reduction in startle activity in females.

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Résumé Aucune variation significative n'a été observée dans les activités sexuelles de huit souches de laboratoire de la mouche méditerranéenne du fruit, Ceratitis capitata (Wiedemann) (Diptera Tephritidae).Deux méthodes ont été utilisées pour sélectionner des souches manifestant une activité sexuelle basse ou élevée: une technique utilisant un seul couple (SP), basée sur la durée de la période avant copulation, et une technique de masse (M), basée sur un index d'accouplement.Des appariements réciproques entre les lignées rapides et lentes ont montré que la différence entre les lignées SP était déterminée par les femelles, tandis que la différence entre les lignées M résultait du comportement des deux sexes. La sélection M, quelle que soit sa direction, affects deux paramètres, la distance entre partenaires lors de la vibration et la durée de la vibration qui tous deux tendent à se raccourcir. La sélection M est aussi associée à une réduction de la réaction des femelles à un brusque stimulus lumineux (Startle activity).

     

地中海实蝇地理种群遗传分化研究

以mtDNA的CO Ⅰ基因的部分序列作为分子标记,研究了来自安哥拉、黎巴嫩、刚果共和国、尼日利亚、贝宁、多哥、加纳、科特迪瓦、南非、埃及、法国、约旦、秘鲁、智利、美国佛罗里达和美国夏威夷等16个国家和地区共75个个体的地中海实蝇Ceratitis capitaza地理种群序列特征和单倍型特征,并分析了各种群间的遗传分化水平和进化关系,结果表明,在供试的样品中发现单倍型的数目是29,其中有5种为共享倍型.地中海实蝇各地理种群的遗传多态性水平差异较大,总体遗传距离D等于0.007.非洲南撒哈拉区(安哥拉、刚果、尼日利亚、贝宁、多哥、加纳、科特迪瓦)的种群的单倍型数、单倍型百分比和群内遗传距离等遗传多态性参数值都高于其它供试地区,多样性明显较其他地区丰富.     

Effects of egg load on finding and acceptance of host fruit in Ceratitis capitata flies

Abstract. In studies conducted on potted host trees in field cages and in the laboratory, we examined the influence of egg load on the finding and acceptance of high-ranking (kumquat) and lower-ranking (grapefruit) hosts for oviposition by wild-origin Mediterranean fruit fly females, Ceratitis capitata (Wiedemann). By prescribing the periods during which females had access to protein prior to testing, we generated four classes of females having progressively increasing egg loads but not differing in population origin, age, degree of protein hunger at testing, or amount of prior experience with host fruit (none). Egg load had no discernible effect on behaviour associated with finding either type of fruit but did have a significant effect on several behaviours associated with oviposition after alighting on fruit. Increasing egg load led to increasing propensity to engage in ovipositional-type behaviour on both kumquats and grapefruits. There was no evidence, however, to support a hypothesis that medflies would become less discriminating against grapefruits relative to kumquats as egg load increased. Relative to kumquats, grapefruits were accepted for oviposition by intermediate and high egg load females to a substantially greater degree in laboratory cages than on trees, suggesting that results of laboratory cage experiments on host discrimination by tephritid flies may poorly reflect differences in behavioural responses expressed under less constrained conditions.     

Cytogenetic analysis of chromosome 5 from the Mediterranean fruit fly,Ceratitis capitata

A cytogenetic map of chromosome 5 from the Mediterranean fruit fly, Ceratitis capitata, was constructed. Six mutations were located by translocation, transposition or deletion mapping. This knowledge allowed alignment and orientation of the existing linkage map with the polytene chromosomes. In addition, mapping of mutations used as selectable markers in genetic sex-separation strains is an essential prerequisite for the improvement of genetic stability during mass-rearing.     

Viability and behaviour of translocations that suppress female recombination in the Mediterranean fruit fly,Ceratitis capitata (Wied.)

Summary The viability of a series of recombination suppressor (RS) strains in Ceratitis capitata, all previously found to contain a reciprocal autosomal translocation, was assessed for egg hatchability and adult emergence in both the homozygous and heterozygous state. Except in T 30C, which contains a Y-autosome translocation in addition to the A-A translocation, egg hatch was significantly reduced in all heterozygous translocation strains, and ranged from 42.4% to 58.5% in seeded eggs compared to a control value of 82.8%. Adult emergence from hatched eggs was affected to a lesser extent, but with a range of 59.5% to 84.2%, compared to the control value of 83.1%, remained significantly reduced in 4 of the 6 translocation strains, as well as in the male line of T 30C. In the homozygous configuration all strains, except T 19 and T 109, showed a significant reduction in egg hatchability, whereas adult emergence was not adversely affected. A significant reduction in the egg hatchability of the translocation heterozygotes compared to that of the homozygotes was observed in 5 of the 7 strains, the observed reduction in T 55/109 being non-significant while that of T 30C was significantly increased. The behaviour of translocations as recombination suppressors and their suitability for inclusion in breeding schemes for the isolation of induced recessive mutations is discussed.This work forms part of a Joint FAO/IAEA research programme on the development of genetic sexing mechanisms for the Mediterranean fruit fly, Ceratitis capitata (Wied.)     

Male diet and age influence to inhibit female remating in Ceratitis capitata (Diptera: Tephritidae)

This study evaluated the influence of age and adult nutritional status of Ceratitis capitata males on their ability to inhibit female remating. Their roles and that of copula duration on the amount of sperm transferred to female spermathecae were also analysed. After emergence, adults were kept in separate groups according to their diets (either high protein – 6.5 g of brewer’s yeast, or low protein – 3.5 g of brewer’s yeast) and their age at the time of use in experiments (4, 8, 12, 16 and 20 days old). The results demonstrated that: (i) male age was not a factor that influenced remating when females mated with well‐nourished males; however, the youngest males (4 days old) in the low‐protein group were less efficient in preventing female remating; (ii) 12‐ and 16‐day‐old males fed on a high‐protein diet transferred and produced more sperm than males of other groups; (iii) there was no correlation between copula duration and the amount of sperm transferred to the female; the longest copula durations were observed among low‐protein‐diet/20‐day‐old males. These results suggest that age and nutrition influence the ability of the males to inhibit female remating. The sterile insect technique is most effective when females do not remate after coupling with sterile males, and therefore, the capacity to inhibit female remating is an important characteristic of males released in the field.     

Susceptibility of the Mediterranean fruit fly (Ceratitis capitata) and the Natal fruit fly (Ceratitis rosa) to entomopathogenic nematodes

The potential of entomopathogenic nematodes, Heterorhabditis bacteriophora, Heterorhabditis zealandica and Steinernema khoisanae, to infect pupariating larvae, pupae and adults of Ceratitis capitata and Ceratitis rosa was investigated in laboratory bioassays. Pupariating larvae and adult flies were susceptible to nematode infection, with no infection recorded for the pupae. Pupariating larvae of C. capitata were generally more susceptible to infection than those of C. rosa. Significantly more larvae of C. capitata were infected by H. bacteriophora. For C. rosa, highest infectivity of larvae was obtained with H. zealandica. In contrast, adults of both species were highly infected by S. khoisanae.     

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.     

Inter-population variation among Ceratitis capitata flies in host acceptance pattern

Significant inter-populational differences in propensity to attempt boring into (accept) various types of fruit for oviposition were found among Ceratitis capitata females from two wild sources and one laboratory source. Evidence suggests that (a) fruit size had a strong influence whereas fruit taxonomic status had little influence on the acceptance pattern of each population, and (b) at least a portion of the inter-populational variation had a genetic basis.

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Zusammenfassung Bei zwei Wildherkünften und einem Laboratoriumsstamm von Ceratitis capitata wurden Weibchen auf ihre Neigung zur Probebohrung in Früchte vor der Eiablage untersucht; dabei ergaben sich gesicherte Unterschiede zwischen den Populationen. Die Resultate deuten drauf hin, (a) dass die Fruchtgrösse einen grossen, und die taxonomische Stellung der Früchte einen kleinen Einfluss auf das Annahmemuster der Populationen ausübt, und (b) dass mindestens ein Teil der Unterschiede zwischen den Populationen genetisch bedingt ist.

     

Performance of controlled-release formulations of trimedlure to attract the Mediterranean fruit fly, Ceratitis capitata

Mediterranean fruit flies (Ceratitis capitata Wied.) are detected by traps baited with the attractant, trimedlure (TML), deposited on the standard cotton dental roll dispenser. Since this dispenser is effective for only 2–4 weeks, controlled-release dispensers may extend the field life of TML-baited traps. A polymeric plug containing 70% by weight TML was highly effective for 8–12 weeks. Other formulations were less effective in the tests with released flies. A release rate of ca. 17 mg of standard TML/day or higher gave insect captures which were generally lower but statistically equivalent to those with the reference dispenser which released 61–63 mg/day. Normalized insect catches increased approximately as the square root of the release rate; the curvilinear regression equation suggested that TML release rates of ca. 52 and 15 mg/day give insect captures equal to ca. 100% and 50%, respectively, of the reference catches. The most attractive TML isomer (TML-C) was a highly effective attractant at a release rate of ca. 4 mg/day.

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Résumé C. capitata était attiré par des pièges alimentés avec de la trimedlure (TML), attractif déposé sur un distributeur commercial de coton dentaire en rouleau. Puisque ce distributeur ne fonctionne que 2 à 4 semaines, des distributeurs à libération contrôlée peuvent augmenter le temps de fonctionnement de pièges fournis en TML. Un bouchon en polymère contenant en poids 70% de TML a été très efficace pendant 8 à 12 semaines. D'autres formules ont été moins efficaces avec des mouches lâchées. Un taux de libération, de 17 mg ou plus de TML moyen par jour, capture moins, mais sans différence statistique significative, qu'un distributeur témoin qui libérait 61 à 63 mg/j. Les captures d'insectes ont augmenté à peu près comme le carré du taux de libération; l'équation de régression curviligne a conduit à penser que la libération de TML à des taux de 52 et 15 mg/j, provoque des captures égales à 100% et 50% des captures de référence. L'isomère le plus attractif (TML-C) a été un attractif très efficace au taux de libération de 4 mg/j.