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Ceratitis Capitata) (Tephritidae) Bisexual and Genetic Sexing Strains: Development, Evaluation and Economics

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Ceratitis Capitata) (Tephritidae) Bisexual and Genetic Sexing Strains: Development, Evaluation and Economics Proceedings of 6th International Fruit Fly Symposium 6–10 May 2002, Stellenbosch, South Africa pp. 367–381 Comparison of Mediterranean fruit fly (Ceratitis capitata) (Tephritidae) bisexual and genetic sexing strains: development, evaluation and economics C. Caceres1, J.P. Cayol2, W. Enkerlin2*, G. Franz1, J. Hendrichs2 & A.S. Robinson1 1Entomology Unit, Agriculture and Biotechnology Laboratory, Agency’s Laboratories, A-2444 Seibersdorf, Austria 2Joint FAO/IAEA Division of Nuclear Techniques in Food and Agriculture, IAEA, Wagramerstrasse 5, A-1400, Vienna, Austria In Medfly, Ceratitis capitata, sterile insect technique (SIT) programmes, the use of genetic sexing strains (GSS) is now routine. The use of these strains in mass-rearing facilities enables them to produce only males for irradiation and release.The advantages of using these strains are described together with genetic approaches used for their construction. The early use of the strains in mass- rearingfacilitieshighlightedimportantlimitationsthatwerenotapparentduringtheirconstruction and small-scale evaluation. Using a combination of new genetic and rearing approaches the problems have essentially been solved. The potential use of one GSS in many different geographic locations raised concerns about mating compatibility. A very detailed field-cage study using populations from very diverse areas showed that these strains had no unexpected impact on mating compatibility. The economics of the use of GSS in SIT programmes is described in detail, covering both their mass production and use in the field. The analysis reveals a much improved cost/benefit ratio when GSS are used. Future improvements that will further enhance the use of GSS are also discussed. BACKGROUND Laboratories at Seibersdorf in Austria, and the The sterile insect technique (SIT) is an estab- funding of three Coordinated Research Projects lished technology for the suppression and/or (CRPs) (IAEA 1990, 1997; Genetica 2002). With eradication of selected key insect pests of man, his the completion of the final CRP, almost all of livestock and crops (Tan 2000). The technology the Medfly rearing facilities worldwide are using involves the mass production and release, on an these genetic sexing strains (GSS) for their SIT area-wide basis,of large numbers of sexually sterile programmes (Robinson et al. 1999; Table 1). insects into a field population. The released males Prior to the start of the first CRP very little was mate with the females in the field resulting in no known about the genetics of Medfly, and the task production of offspring. Following repeated of this CRP was to develop many of the essential releases of the sterilized insects, the field popula- genetic tools, e.g. mutations, polytene chromo- tion is suppressed and in certain circumstances some analysis and male-linked translocations, eradication can be achieved (Hendrichs 2000). which are essential for the construction of a GSS. To be effective,SIT does not require the release of Table 1. Medfly mass-rearing facilities using GSS with sterile females as they do not contribute to the the potential capacity in millions of males/week. transfer of sterility to the wild population. Thus, mass production of sterile females is unnecessary Country Strain Capacity and the technique requires that only sterile male insects are released (Knipling 1955; Robinson et al. Argentina SEIB 6 (VIENNA 7/Tol) 80 1999; McInnis et al. 1994; Franz & McInnis 1995; Australia VIENNA 7/Mix 7 Hendrichs et al. 1995). In some cases the release Chile SEIB 6 (VIENNA 7) 40 of females can have a negative impact. Greece (Crete) SEIB 7/Mix 3 As a result of this situation, the Joint FAO/IAEA Guatemala VIENNA 7/Tol 1600 Division of Nuclear Techniques in Food and Hawaii (CDFA) VIENNA 7/Tol 100 Agriculture initiated activities in 1981 to develop Peru VIENNA 7/Mix 100 special strains for the Medfly, Ceratitis capitata, Portugal (Madeira) VIENNA 7/Mix 50 that could be used to produce only males for area- Seibersdorf VIENNA 7-D53/Mix 25 wide SIT programmes. These activities included South Africa VIENNA 7-D53/Mix 7 Hawaii VIENNA 7/Tol 200 research and development at the FAO/IAEA (USDA/APHIS) *To whom correspondence should be addressed. Total 2212 E-mail: [email protected] 368 Proceedings of the 6th International Fruit Fly Symposium At the conclusion of the second CRP, the first sterile females) and thus released at a more transfer of a GSS to an operational SIT programme mature age, thereby reducing losses before in Guatemala took place. males reach full sexual capacity; Studies carried out during the second CRP e. simplified and more precise monitoring demonstrated the efficacy of an all-male release activities when using female attractants, in the field (McInnis et al. 1994) and included infor- as the recapture of sterile males is largely mation on a temperature-sensitive mutation, tsl, reduced, thus also significantly reducing the in Medfly (Franz et al. 1996). The use of this risk of mis-identification (plus the value of mutation has been central to the practical applica- not removing many valuable sterile males); tion of GSS in Medfly. Some of the most serious f. no damage in certain types of fruit due to problems to be confronted in the use of GSS in absence of oviposition stings by sterile females, operational programmes concerned the stability and reduced transfer of pathogenic fungi and of the strains under mass-rearing conditions. bacteria to such fruit; These have now been essentially solved by a g. increased applicability of SIT for Medfly combination of new rearing methods and the suppression in fruit growing regions, because use of more appropriate translocations. the reduced cost and absence of oviposition Initially it was assumed that the sex determina- damage to the fruit enables the routine use tion mechanism of Medfly would be very similar to of sterile males as a biological ‘insecticide’ to that of Drosophila melanogaster. However, it is replace chemical bait-sprays during fruiting now known that in most tephritid fruit flies there is seasons; probably a single gene that initiates male sex h. increased bio-safety,as an accidental release of determination (Zapater & Robinson 1986; Will- non-irradiated flies would only include males, hoeft & Franz 1996) in contrast to D. melanogaster, and escaping females from mass-rearing where a chromosomal balance system operates. would have reduced fitness. This is particu- The use of polytene chromosome analysis larly important for mass-rearing facilities (Kerremans et al. 1990, 1992; Zacharopoulou located in fruit fly-free areas or areas where et al. 1991) has been essential for the develop- Medfly eradication is the objective. ment of stable GSS, by enabling both mutations In conclusion, the most important benefit is the and translocation breakpoints to be accurately increased efficiency of SIT, as it has been shown mapped (Franz & Kerremans 1993; Franz et al. in many Medfly studies that male-only releases 1994; Kerremans & Franz 1994, 1995). introduce 3–4 times more sterility into the target population than do bi-sexual releases (McInnis DEVELOPMENT,IMPROVEMENTS AND et al. 1986; Robinson et al. 1986; McInnis et al. REARING OF MEDFLY GENETIC SEXING 1994; Rendon et al. 2000). STRAINS FOR USE IN SIT Choice of sexing system Benefits of genetic sexing strains (GSS) The practicality and economics of GSS use The benefits of using GSS for Medfly SIT have depends on the choice of the appropriate been articulated many times and were summa- selectable marker in the sexing system. Initially rized by Hendrichs et al. (1995) as follows: the white pupae (wp) mutation (Rössler 1979) was a. economic savings in rearing, irradiation, used in combination with seed sorters to separate packaging, transport and release; white (female) and brown (male) pupae. However, b. increased male quality as male pupae can be it became clear that this type of selectable marker irradiated 24 h before emergence, instead of has two significant disadvantages. First of all, the 48 h when also irradiating females; sex separation can be achieved only after the costly c. several-fold increase in field effectiveness as mass-rearing step, i.e. expensive diet has to be the sterile sperm is not wasted in matings with wasted on rearing females and, in addition, the sterile females, and sterile males compete females had to be killed and disposed of in a safe better for wild females. Some studies also way. Second, the use of seed sorters is not easy. showed that sterile males disperse more in Such machines are very expensive, especially the absence of sterile females; considering the separation capacity needed in d. in the absence of sterile females in the fly most mass-rearing facilities; they are relatively emergence containers, sterile males can be complicated and are not very accurate (c. 5% held longer (without mating taking place with contamination with females in the ‘male-only’ Caceres et al.: Medfly bisexual and genetic sexing strains: development, evaluation and economics 369 product and a significant loss of male pupae). translocation breakpoint on that chromosome Furthermore, the sorting process can damage made it possible to significantly reduce the the pupae, resulting in flies with reduced flight impact of a particular genetic destabilizing ability. event. Acknowledging these severe disadvantages led There is now a clear understanding of the ge- to the development of an alternative sexing netic factors that are involved in the stability of system. This is
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