Orange-derived stimuli regulating oviposition in the Mediterranean fruit fly

Abstract:  Some visual and olfactory host stimuli influencing oviposition of the Mediterranean fruit fly ( Ceratitis capitata Wiedemann) in orange fruit were investigated in a laboratory-reared strain of this species. Mated females of C. capitata were found to be attracted to the same extent by fragrant orange fruits and odourless sham oranges, while unmated females were notably less attracted than mated females by the above objects. Mated females laid significantly more eggs in orange-coloured than in whitish (optically neutral) spherical wax dummies (diameter, 9.0 cm) as well as in both orange-coloured and whitish wax dummies supplemented with high internal humidity (80–100%) compared with respective wax dummies supplied with low internal humidity (30–50%). The orange-like colour together with high internal humidity provides basic stimuli supporting adequate oviposition of C. capitata in oranges. The oviposition rate of female C. capitata was not significantly changed when graded dosages ranging from 0.004 to 19.6  μ l of orange peel oil per cm² were added to orange-coloured wax dummies, while oviposition was considerably subdued by addition of 3.9  μ l orange peel oil and completely disrupted by addition of 9.8 or 19.6  μ l of orange peel oil per cm² of whitish wax dummies. Oviposition of C. capitata in ripe orange fruit may thus be interpreted by the predilection of this tephritid species for an orange-coloured, glossy pericarp, being capable of counteracting the deterrent effect of the essential oil found in orange flavedo.     

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.     

Enhancement of citrus resistance to the Mediterranean fruit fly

Precolorbreak treatment of Marsh grapefruit and Shamouti and Valencia oranges with 20 or 50 ppm gibberellic acid (GA) reduced fruit susceptibility to laboratory infestation by Ceratitis capitata (Wiedemann). Treatment effects were dose-dependent and the benefits were most pronounced with the orange cultivars. The implications of these findings are discussed in relation to use of GA treatment for improved fruit fly control in citrus.

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Zusammenfassung In den hier beschriebenen Versuchen wurde untersucht, wie weit durch die Anwendung von Gibberilinsäure (GA) im Freiland die Resistenz von Pampelmuse (Marsh), sowie von Shamoutiund Valencia-Orangen gegenüber Befall durch die Mittelmeerfruchtfliege, Ceratitis capitata (Wiedemann) gesteigert werden kann.In den Jahren 1982/83 spritzten wir in Israel Bäume dieser drei Zitrusarten mit GA in den Konzentrationen von 5 ppm und 50 ppm zu zwei verschiedenen Zeitpunkten vor dem Farbumschlag der Früchte (September, Oktober). Die so behandelten Früchte wurden anschliessend während der Reifeperiode in verschiedenen Zeitpunkten adulten Mittelmeerfruchtfliegen für die Eiablage dargeboten. Diese Exposition wurde sowohl im Freiland mit Früchten am Baum sowie im Laboratorium mit gepflückten Früchten durchgeführt. Untersucht wurde einerseits die Anfälligkeit resp. Resistenz der Früchte bezüglich der Eiablage, andererseits die Entwicklung der Larven in den Früchten.Wir stellten fest, dass die Behandlung mit GA bei Pampelmuse in Abhängigkeit vom Zeitpunkt und Konzentration der Behandlung eine gewisse erhöhte Befallsresistenz bewirken konnte. Bezüglich der beiden geprüften Orangensorten konnten wegen ungünstigen Witterungsbedingungen während der Reifezeit keine schlüssigen Befunde erarbeitet werden.Im Winterhalbjahr 1983/84 wurden Bäume aller drei Zitrussorten vor dem Farbwechsel der Früchte mit 20 ppm und 50 ppm GA behandelt. Um den negativen Einfluss der Witterung in den Expositionsversuchen auszuschalten, wurden die Versuchsfrüchte den Fruchtfliegen während der Reifeperiode ausschliesslich im Laboratorium dargeboten. In dieser Versuchsperiode stellten wir auch bei Orangen eine erhöhte Widerstandsfähigkeit gegen Fruchtfliegenbefall fest, und zwar in erhöhtem Ausmass bei Valencia-Orangen. Gleichzeitig beobachteten wir eine stark verzögerte Entwicklung des Farbwechsels von grün auf orange als Folge der hohen GA-Konzentration.

     

Male biased sex ratio in the Mediterranean fruit fly Ceratitis capitata, an example of Y-chromosome meiotic drive

A case of Y-chromosome meiotic drive is reported in the Mediterranean fruit fly Ceratitis capitata. It arose in an irradiated male and results in excess of males. Male excess is inherited strictly from father to son. A Y-linked factor MP (male producer) is proposed. Higher drive can be selected, but distortion declines rapidly in the absence of selection. Hybrid males from crosses between driving males and nondriving females also show drive but to a reduced extent, suggesting the action of suppressors. Sex ratio distortion is independent of postzygotic mortality, and is not associated with an obvious chromosome arrangement. Spermiogenesis in driving males is characterised by abnormalities in sperm tails and reduced numbers in some sperm cysts, whereas neighbouring cysts of the same MP testis are essentially wild type. The average number of missing sperms plus deformed sperms approximates to the average depression in female recovery among the progenies of siblings, suggesting that most of the missing or abnormal sperms would have given rise to females, that is, they would have been X-bearing. To explain the heterogeneity between neighbouring cysts, a theory is proposed that links it to variation in X-chromosome sensitivity to MP, arising by random suppression of the genetic basis of sensitivity during the six mitotic divisions in the origin of the cyst from its stem cell before meiosis.     

Male rank and optimal lek size

Widemo and Owens presented a model that calculates the expectedcopulation rates of males on leks of a range of sizes. Theyclaim that a negative relationship between lek size and malemating skew will result in low-ranking males having greateroptimal lek sizes than higher ranking rivals. Widemo and Owensoffered no proof of their claim, and their model assumes thatthe rank of a male does not change as lek size increases, whereasin reality, rank may change as more males arrive. We presenta general model that allows rank to change as lek size increases.We show that the crucial determinant of whether optimal leksize increases with male rank is whether relative competitive differencesincrease with lek size. Contrary to the claim of Widemo andOwens, the relationship between skew and lek size has no directbearing on the optimal levels of aggregation of males of differentrank. We show that a negative relationship between skew andlek size can exist even when high-ranking males have the greatestoptimal lek sizes.     

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.     

Assessing the invasive potential of the Mediterranean fruit fly in California and Italy

The polyphagous tropical Mediterranean fruit fly (Ceratitis capitata Weid. (medfly)) was detected in California in 1975, and a large-scale detection/eradication campaign was begun in the absence of sound knowledge of the fly’s potential invasiveness and geographic distribution. Persistent measurable populations of the fly have not been found in California, but a scientific explanation for this has not developed. A physiologically based demographic system model (CASAS) was developed to examine the effects of temperature on medfly’s potential distribution across the ecological zones of Arizona–California (AZ–CA), and in Italy where the fly is established. The system model simulates the daily age-mass structured dynamics of a tree host composed of sub-unit populations of leaves, stem, roots and fruit, as well as the age-structured dynamics of medfly life stages. Total pupae tree⁻¹ year⁻¹ was used as the metric of favorability for medfly at 151 locations in AZ–CA during 1995–2006, and at 84 locations in Italy during 1999–2005. The results were mapped using GRASS GIS. AZ and the southern desert areas of CA are unfavorable for medfly because of high summer temperatures, while much of CA, including many frost-free areas, is too cold. Only the area of south coastal CA (San Diego, Orange and Los Angeles Counties) is potentially favorable for medfly, but in the absence of measurable populations, we cannot say whether it is established there. The majority of medfly discoveries over the past 35 years have occurred in south coastal CA, but discoveries also occurred in Santa Clara County in northern CA, mostly during 1975 and 1980–1981. Santa Clara County, just south of San Francisco Bay, is generally marginal for medfly, but favorability increased approximately 10% during the period 1979–1981. Medfly has been established in Italy for decades, and our model predicts its wide distribution in the southern and western regions of the country. The fly is restricted in northern areas and at higher elevations of Italy by winter temperatures. Temperature is expected to increase in CA and the Mediterranean Basin. We used two scenarios consisting of increasing observed daily temperatures +2 and +3°C to examine the effects on the potential distribution of the fly in CA and Italy. Increasing temperatures expand the favorable range for medfly northward along the coast of CA, but decrease it in the southern reaches of current favorability. A similar but greater increase in geographic range is predicted for Italy. We examine critically some ongoing eradication programs in CA, and question the scientific basis for them. We also review some climate matching approaches used to assess the potential geographic distribution of invasive species.     

Nutritional reserves regulate male participation in Mediterranean fruit fly leks

1. The mating system of Mediterranean fruit flies ( Ceratitis capitata , Diptera: Tephritidae), is based on male leks that form on the foliage of trees. Following observations that not all males participate in leks, the hypotheses that (a) small males are absent from leks and (b) males with inadequate nutrient reserves are unable to participate in leks were examined.
2. The size and weight of lekking males ( n = 183) and resting males captured at the same time in the vicinity of leks ( n = 148) were established. In addition, using biochemical techniques, the amounts of sugar, glycogen, lipid and protein in each individual were established quantitatively.
3. There was no significant size difference between lekking and resting males. However, lekking males were significantly heavier and contained significantly more sugars and protein than resting males.
4. In conclusion, leks are exclusive, and only males with adequate nutritional reserves may join. Thus, in this species, reproductive success is closely linked to foraging success.     

A new long-life trimedlure dispenser for Mediterranean fruit fly

New agricultural techniques are attempting to reduce the application of synthesized pesticides and replace them with new environmentally friendly methods such as mass trapping, mating disruption, or chemosterilization techniques. All these methods are based on the release of a lure for insect attraction or confusion. The success of the chosen method depends on the quality of the attractant emission from the dispenser. Currently, used dispensers with a polymeric matrix and new dispensers with mesoporous inorganic materials were evaluated to obtain more efficient emission kinetics. In this study, the selected pest was the Mediterranean fruit fly, Ceratitis capitata (Wiedemann) and the lure used was trimedlure (TML). The dispensers were validated by means of a field study comparing insect catches with attractant release values. As a result, we have demonstrated that mesoporous dispensers have a clearly longer lifetime than the polymeric plug. Furthermore, the attractant release rate is less dependent on temperature in mesoporous than in polymeric dispensers.     

Susceptibility of low-chill blueberry cultivars to Mediterranean fruit fly, oriental fruit fly, and melon fly (Diptera: Tephritidae)

No-choice tests were conducted to determine whether fruit of southern highbush blueberry, Vaccinium corymbosum L., hybrids are hosts for three invasive tephritid fruit flies in Hawaii. Fruit of various blueberry cultivars was exposed to gravid female flies of Bactrocera dorsalis Hendel (oriental fruit fly), Ceratitis capitata (Wiedemann) (Mediterranean fruit fly), or Bactrocera cucurbitae Coquillet (melon fly) in screen cages outdoors for 6 h and then held on sand in the laboratory for 2 wk for pupal development and adult emergence. Each of the 15 blueberry cultivars tested were infested by oriental fruit fly and Mediterranean fruit fly, confirming that these fruit flies will oviposit on blueberry fruit and that blueberry is a suitable host for fly development. However, there was significant cultivar variation in susceptibility to fruit fly infestation. For oriental fruit fly, 'Sapphire' fruit produced an average of 1.42 puparia per g, twice as high as that of the next most susceptible cultivar 'Emerald' (0.70 puparia per g). 'Legacy', 'Biloxi', and 'Spring High' were least susceptible to infestation, producing only 0.20-0.25 oriental fruit fly puparia per g of fruit. For Mediterranean fruit fly, 'Blue Crisp' produced 0.50 puparia per g of fruit, whereas 'Sharpblue' produced only 0.03 puparia per g of fruit. Blueberry was a marginal host for melon fly. This information will aid in development of pest management recommendations for blueberry cultivars as planting of low-chill cultivars expands to areas with subtropical and tropical fruit flies. Planting of fruit fly resistant cultivars may result in lower infestation levels and less crop loss.     

Effect of marking pheromone on clutch size in the Mediterranean fruit fly

Abstract Using acridine orange to selectively stain eggs, we showed that wild-collected female Mediterranean fruit flies (Ceratitis capitata Wiedemann) laid fewer eggs per clutch in fruit previously infested with eggs than in uninfested fruit. This effect is apparently attributable to marking pheromone deposited by females after oviposition: clutch size on fruit infested with eggs but free of marking pheromone was not statistically different from that on uninfested fruit. Clutch size on uninfested fruit on which marking pheromone was artificially transferred was significantly lower than that on uninfested and untreated fruit. Marking pheromone had a comparable though not statistically significant effect on the clutch size of females originating from a strain maintained in the laboratory for several hundred generations.     

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.     

Thermotolerance and HSP70 expression in the Mediterranean fruit fly Ceratitis capitata

The relationship between Hsp70 expression and thermotolerance has been well documented in Drosophila melanogaster. However, there is limited information on this relationship in other insect species. In this report we describe the Hsp70-thermotolerance relationship in one of the major fruit fly pests, Ceratitis capitata (medfly). Hsp70 expression and thermotolerance were assayed at a range of temperatures in several stages of medfly development. The most thermotolerant stage was found to be the late larval stage (100% survival at 41 °C) followed by adult flies and late embryos (100% survival at 39 °C). These three stages showed a positive relationship between Hsp70 expression and thermotolerance. Mid-larval and mid-embryonic stages were found less thermotolerant and the Hsp70-thermotolerance relationship was not evident. Early embryos did not express Hsp70 at any temperature and exhibited the lowest thermotolerance. The relationship between Hsp70 and inducible thermotolerance was also studied in late larvae. A pretreatment at 37-39 °C increased thermotolerance at higher temperatures by approximately 1 °C. In parallel, the pretreatment increased Hsp70 expression suggesting a close link between Hsp70 expression and inducible thermotolerance. The increased Hsp70 levels after pretreatment were found to be due to the increased levels of the hsp70 RNA.     

Mediterranean fruit fly as a potential vector of bacterial pathogens

The Mediterranean fruit fly (Ceratitis capitata) is a cosmopolitan pest of hundreds of species of commercial and wild fruits. It is considered a major economic pest of commercial fruits in the world. Adult Mediterranean fruit flies feed on all sorts of protein sources, including animal excreta, in order to develop eggs. After reaching sexual maturity and copulating, female flies lay eggs in fruit by puncturing the skin with their ovipositors and injecting batches of eggs into the wounds. In view of the increase in food-borne illnesses associated with consumption of fresh produce and unpasteurized fruit juices, we investigated the potential of Mediterranean fruit fly to serve as a vector for transmission of human pathogens to fruits. Addition of green fluorescent protein (GFP)-tagged Escherichia coli to a Mediterranean fruit fly feeding solution resulted in a dose-dependent increase in the fly's bacterial load. Flies exposed to fecal material enriched with GFP-tagged E. coli were similarly contaminated and were capable of transmitting E. coli to intact apples in a cage model system. Washing contaminated apples with tap water did not eliminate the E. coli. Flies inoculated with E. coli harbored the bacteria for up to 7 days following contamination. Fluorescence microscopy demonstrated that the majority of fluorescent bacteria were confined along the pseudotrachea in the labelum edge of the fly proboscis. Wild flies captured at various geographic locations were found to carry coliforms, and in some cases presumptive identification of E. coli was made. These findings support the hypothesis that the common Mediterranean fruit fly is a potential vector of human pathogens to fruits.     

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.     

Invasion genetics of the Mediterranean fruit fly: variation in multiple nuclear introns

Biological invasions generally start from low initial population sizes, leading to reduced genetic variation in nuclear and especially mitochondrial DNA. Consequently, genetic approaches for the study of invasion history and population structure are difficult. An extreme example is the Mediterranean fruit fly, Ceratitis capitata (Medfly), for which successive invasions during this century have resulted in a loss of 60% of ancestral genetic variation in isozymes and 75% of variation in mitochondrial DNA. Using Medflies as an example, we present a new approach to invasion genetics that measures DNA sequence variation within introns from multiple nuclear loci. These loci are so variable that even relatively recently founded Medfly populations within California and Hawaii retain ample genetic diversity. Invading populations have only lost 35% of the ancestral genetic variation. Intron variation will allow high-resolution genetic characterization of invading populations in both natural and managed systems, although non-equilibrium methods of analysis may be necessary if the genetic diversity represents sorting ancestral polymorphism.     

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).