Neoclassical development of genetic sexing strains for insect pest and disease vector control

Giovanni Petrucci , Maria-Eleni Gregoriou , Philippos Aris Papathanos , Marc F. Schetelig , Zhijian Tu , Kostas Bourtzis

Insect Science ›› 2026, Vol. 33 ›› Issue (2) : 618 -639.

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Insect Science ›› 2026, Vol. 33 ›› Issue (2) :618 -639. DOI: 10.1111/1744-7917.70192
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Neoclassical development of genetic sexing strains for insect pest and disease vector control
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Abstract

The sterile insect technique, which consists of the mass production and release of sterile insects to control populations of pests and disease vectors, has been effectively used for decades. An important component of sterile insect technique field applications is the availability of sex separation systems that reliably and economically eliminate females from mass-reared sterile insect populations destined for field release. Genetic sexing strains are important for the effectiveness and cost-efficiency of insect population control programs, including sterile insect technique. Classical approaches to generate genetic sexing strains, such as irradiation-induced chromosomal translocations, have yielded stable strains for species like the Mediterranean fruit fly, Ceratitis capitata. However, significant efforts are needed to establish genetic sexing strains using classical genetic methods, as large-scale random mutagenesis and screening are needed. We introduce here a neoclassical genetic approach, leveraging CRISPR-based gene-editing to target known genes to develop selectable genetic markers, followed by genetic rescue in a male-specific manner to speed up the development of genetic sexing strains and enhance their precision, stability, and adaptability. The integration of molecular tools, genetic markers like the white pupae and temperature-sensitive lethal, and strategies for maintaining genetic stability are discussed. We also review the challenges and opportunities in applying classical, transgenic, and neoclassical genetic approaches to improve genetic sexing strains for pest management.

Keywords

cardinal / CRISPR / gene editing / sterile insect technique / temperature-sensitive lethal / white pupae

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Giovanni Petrucci, Maria-Eleni Gregoriou, Philippos Aris Papathanos, Marc F. Schetelig, Zhijian Tu, Kostas Bourtzis. Neoclassical development of genetic sexing strains for insect pest and disease vector control. Insect Science, 2026, 33 (2) : 618-639 DOI:10.1111/1744-7917.70192

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References

[1]

Ahmed, H.M.M., Hildebrand, L. and Wimmer, E.A. (2019) Improvement and use of CRISPR/Cas9 to engineer a sperm-marking strain for the invasive fruit pest Drosophila suzukii. BMC Biotechnology, 19, 85.

[2]

Alphey, L. (2002) Re-engineering the sterile insect technique. Insect Biochemistry and Molecular Biology, 32, 1243–1247.

[3]

Andrade, P.P., da Silva Ferreira, M.A., Muniz, M.S. and de Casto Lira-Neto, A. (2018) GM insect pests under the Brazilian regulatory framework: development and perspectives. BMC Proceedings, 12, 16.

[4]

Ant, T., Koukidou, M., Rempoulakis, P., Gong, H.F., Economopoulos, A., Vontas, J. et al. (2012) Control of the olive fruit fly using genetics-enhanced sterile insect technique. BMC Biology, 10, 51.

[5]

Asad, M., Chang, Y., Liao, J. and Yang, G. (2025) CRISPR/Cas9 genome editing in the Diamondback moth: current progress, challenges, and prospects. International Journal of Molecular Sciences, 26, 1515.

[6]

Augustinos, A.A., Targovska, A., Cancio-Martinez, E., Schorn, E., Franz, G., Cáceres, C. et al. (2017) Ceratitis capitata genetic sexing strains: laboratory evaluation of strains from mass-rearing facilities worldwide. Entomologia Experimentalis et Applicata, 164, 305–317.

[7]

Augustinos, A.A., Misbah-ul-Haq, M., Carvalho, D.O., de la Fuente, L.D., Koskinioti, P. and Bourtzis, K. (2020) Irradiation induced inversions suppress recombination between the M locus and morphological markers in Aedes aegypti. BMC Genetics, 21, 142.

[8]

Augustinos, A.A., Nikolouli, K., Duran de la Fuente, L., Misbah-ul-Haq, M., Carvalho, D.O. and Bourtzis, K. (2022) Introgression of the Aedes aegypti red-eye genetic sexing strains into different genomic backgrounds for sterile insect technique applications. Frontiers in Bioengineering and Biotechnology, 10, 821428.

[9]

Aumann, R.A., Gouvi, G., Gregoriou, M.E., Rehling, T., Sollazzo, G., Bourtzis, K. et al. (2025) Decoding and engineering temperature-sensitive lethality in Ceratitis capitata for pest control. Proceedings of the National Academy of Sciences USA, 122, e2503604122.

[10]

Aumann, R.A., Häcker, I. and Schetelig, M.F. (2020) Female-to-male sex conversion in Ceratitis capitata by CRISPR/Cas9 HDR-induced point mutations in the sex determination gene transformer-2. Scientific Reports, 10, 18611.

[11]

Aumann, R.A., Schetelig, M.F. and Häcker, I. (2018) Highly efficient genome editing by homology-directed repair using Cas9 protein in Ceratitis capitata. Insect Biochemistry and Molecular Biology, 101, 85–93.

[12]

Aviles, E.I., Rotenberry, R.D., Collins, C.M., Dotson, E.M. and Benedict, M.Q. (2020) Fluorescent markers rhodamine B and uranine for Anopheles gambiae adults and matings. Malaria Journal, 19, 236.

[13]

Bassett, A.R. and Liu, J.L. (2014) CRISPR/Cas9 and genome editing in Drosophila. Journal of Genetics and Genomics, 41, 7–19.

[14]

Baumhover, A.H., Graham, A.J., Bitter, B.A., Hopkins, D.E., New, W.D., Dudley, F.H. et al. (1955) Screw-worm control through release of sterilized flies. Journal of Economic Entomology, 48, 462–466.

[15]

Bedo, D.G. (1986) Polytene and mitotic chromosome analysis in Ceratitis capitata (Diptera; Tephritidae). Canadian Journal of Genetics and Cytology, 28, 180–188.

[16]

Bello, B., Resendez-Perez, D. and Gehring, W.J. (1998) Spatial and temporal targeting of gene expression in Drosophila by means of a tetracycline-dependent transactivator system. Development (Cambridge, England), 125, 2193–2202.

[17]

Bernardini, F., Haghighat-Khah, R.E., Galizi, R., Hammond, A.M., Nolan, T. and Crisanti, A. (2018) Molecular tools and genetic markers for the generation of transgenic sexing strains in Anopheline mosquitoes. Parasites & Vectors, 11, 660.

[18]

Bhalla, S.C. and Craig, G.B. (1970) Linkage analysis of chromosome i of Aedes aegypti. Canadian Journal of Genetics and Cytology, 12, 425–435.

[19]

Bi, H.L., Xu, J., He, L., Zhang, Y., Li, K. and Huang, Y.P. (2019) CRISPR/Cas9-mediated ebony knockout results in puparium melanism in Spodoptera litura. Insect Science, 26, 1011–1019.

[20]

Bi, H.L., Xu, J., Tan, A.J. and Huang, Y.P. (2016) CRISPR/Cas9-mediated targeted gene mutagenesis in Spodoptera litura. Insect Science, 23, 469–477.

[21]

Biedler, J.K., Aryan, A., Qi, Y., Wang, A., Martinson, E.O., Hartman, D.A. et al. (2024) On the origin and evolution of the mosquito male-determining factor nix. Molecular Biology and Evolution, 41, msad276.

[22]

Bohle, F., Schneider, R., Mundorf, J., Zühl, L., Simon, S. and Engelhard, M. (2024) Where does the EU-path on new genomic techniques lead us? Frontiers in Genome Editing, 6, 1377117.

[23]

Bouyer, J. and Lefrançois, T. (2014) Boosting the sterile insect technique to control mosquitoes. Trends in Parasitology, 30, 271–273.

[24]

Brand, A.H. and Perrimon, N. (1993) Targeted gene expression as a means of altering cell fates and generating dominant phenotypes. Development (Cambridge, England), 118, 401–415.

[25]

Buchman, A. and Akbari, O.S. (2019) Site-specific transgenesis of the Drosophila melanogaster Y-chromosome using CRISPR/Cas9. Insect Molecular Biology, 28, 65–73.

[26]

Caceres, C. (2002) Mass rearing of temperature sensitive genetic sexing strains in the mediterranean fruit fly (Ceratitis Capitata). Genetica, 116, 107–116.

[27]

Cáceres, C., Bourtzis, K., Gouvi, G., Vreysen, M.J.B., Bimbilé Somda, N.S., Hejníčková, M. et al. (2023) Development of a novel genetic sexing strain of Ceratitis capitata based on an X-autosome translocation. Scientific Reports, 13, 16167.

[28]

Cáceres, C., Cayol, J.P., Enkerlin, W., Franz, G., Hendrichs, J. and Robinson, A.S. (2004) Comparison of Mediterranean fruit fly (Ceratitis capitata) (Tephritidae) bisexual and genetic sexing strains: development, evaluation and economics. Proceedings of 6th International Fruit Fly Symposium, 1, 367–381.

[29]

Carballar-Lejarazú, R., Ogaugwu, C., Tushar, T., Kelsey, A., Pham, T.B., Murphy, J. et al. (2020) Next-generation gene drive for population modification of the malaria vector mosquito, Anopheles gambiae. Proceedings of the National Academy of Sciences USA, 117, 22805–22814.

[30]

Catteruccia, F., Benton, J.P. and Crisanti, A. (2005) An Anopheles transgenic sexing strain for vector control. Nature Biotechnology, 23, 1414–1417.

[31]

Cayol, J.P. (1999) Changes in sexual behavior and life history traits of tephritid species caused by mass-rearing processes. In Fruit Flies (Tephritidae) (eds. M. Aluja & A. Norrbom), pp. 861–878. CRC Press, Taylor and Francis Group, LLC, Florida, USA.

[32]

Chen, C., Compton, A., Nikolouli, K., Wang, A., Aryan, A., Sharma, A. et al. (2022) Marker-assisted mapping enables forward genetic analysis in Aedes aegypti, an arboviral vector with vast recombination deserts. Genetics, 222, iyac140.

[33]

Choo, A., Fung, E., Chen, I.Y., Saint, R., Crisp, P. and Baxter, S.W. (2020) Precise single base substitution in the shibire gene by CRISPR/Cas9-mediated homology directed repair in Bactrocera tryoni. BMC Genetics, 21, 127.

[34]

Clymans, R., Van Kerckvoorde, V., Beliën, T., Bylemans, D. and De Clercq, P. (2020) Marking Drosophila suzukii (Diptera: Drosophilidae) with fluorescent dusts. Insects, 11, 152.

[35]

Cock, M.J.W., Murphy, S.T., Kairo, M.T.K., Thompson, E., Murphy, R.J. and Francis, A.W. (2016) Trends in the classical biological control of insect pests by insects: an update of the BIOCAT database. BioControl, 61, 349–363.

[36]

Concha, C., Palavesam, A., Guerrero, F.D., Sagel, A., Li, F., Osborne, J.A. et al. (2016) A transgenic male-only strain of the new world screwworm for an improved control program using the sterile insect technique. BMC Biology, 14, 72.

[37]

Concha, C., Yan, Y., Arp, A., Quilarque, E., Sagel, A., de León, A.P. et al. (2020) An early female lethal system of the New World screwworm, Cochliomyia hominivorax, for biotechnology-enhanced SIT. BMC Genetics, 21, 143.

[38]

Criscione, F., Qi, Y., Saunders, R., Hall, B. and Tu, Z. (2013) A unique Y gene in the Asian malaria mosquito Anopheles stephensi encodes a small lysine-rich protein and is transcribed at the onset of embryonic development. Insect Molecular Biology, 22, 433–441.

[39]

Criscione, F., Qi, Y. and Tu, Z. (2016) GUY1 confers complete female lethality and is a strong candidate for a male-determining factor in Anopheles stephensi. eLife, 5, e19281.

[40]

Curtis, C.F., Akiyama, J. and Davidson, G. (1976) A genetic sexing system in Anopheles gambiae species A. Mosquito News, 36, 492–498.

[41]

Davydova, S., Liu, J., Kandul, N.P., Braswell, W.E., Akbari, O.S. and Meccariello, A. (2023) Next-generation genetic sexing strain establishment in the agricultural pest Ceratitis capitata. Scientific Reports, 13, 19866.

[42]

Doudna, J.A. and Charpentier, E. (2014) The new frontier of genome engineering with CRISPR-Cas9. Science, 346, 1258096.

[43]

Dyck, V.A., Hendrichs, J. and Robinson, A.S. (2021) Sterile Insect Technique: Principles and Practice in Area-Wide Integrated Pest Management. CRC Press, Taylor and Francis Group, LLC, Florida, USA.

[44]

EASAC (2024) EASAC welcomes reform of EU legislation on new genome techniques. https://easac.eu/news/details/easac-welcomes-reform-of-eu-legislation-new-genome-techniques.

[45]

EPA (2020) Human health and environmental risk assessment for the new product OX5034 containing the tetracycline-repressible transactivator protein variant (tTAV-OX5034; new active ingredient) protein, a DsRed2 protein variant (DsRed2-OX5034; new inert ingredient), and the genetic material (Vector pOX5034) necessary for their production in OX5034 Aedes aegypti. https://www.regulations.gov/document/EPA-HQ-OPP-2019-0274-0359.

[46]

EPA (2022) Following review of available data and public comments, EPA expands and extends testing of genetically engineered mosquitoes to reduce mosquito populations. https://www.epa.gov/pesticides/following-review-available-data-and-public-comments-epa-expands-and-extends-testing#:~:text=EPA%20has%20approved%20an%20experimental%20use%20permit%20%28EUP%29,Aedes%20aegypti%20%28OX5034%29%20mosquitoes%20to%20reduce%20mosquito%20populations.

[47]

Epsky, N.D., Hendrichs, J., Katsoyannos, B.I., Vásquez, L.A., Ros, J.P., Zümreoglu, A. et al. (1999) Field evaluation of female-targeted trapping systems for Ceratitis capitata (Diptera: Tephritidae) in seven countries. Journal of Economic Entomology, 92, 156–164.

[48]

European Commission. (2023) Proposal for a regulation of the European Parliament and of the Council on Plants obtained by certain new genomic techniques and their food and feed, and amending regulation (EU) 2017/625. https://eur-lex.europa.eu/legal-content/EN/TXT/HTML/?uri=CELEX:52023PC0411.

[49]

European Commission. (2021) Study on the status of new genomic techniques under union law and in light of the court of justice ruling in case C-528/16. https://food.ec.europa.eu/system/files/2021-04/gmo_mod-bio_ngt_eu-study.pdf.

[50]

European Court of Justice. (2018) Judgment in Case C-528/16, Confédération paysanne and Others v Premier Ministre and Ministre de l'Agriculture, de l'Agroalimentaire et de la Forêt. https://curia.europa.eu/juris/document/document.jsf;jsessionid=DFF4D1E9E0550FE99E436A47B4CDFDE5?text=&docid=204387&pageIndex=0&doclang=EN&mode=req&dir=&occ=first&part=1&cid=2996154.

[51]

FAO/IAEA, 1st RCM Report. (2019) First Research Coordination Meeting Joint FAO/IAEA Division of Nuclear Techniques in Food and Agriculture. Research Coordination Meeting on Generic Approach for the Development of Genetic Sexing Strains for SIT Applications. Vienna International Centre, Vienna, Austria. https://www.iaea.org/sites/default/files/20/11/d44003-rcm1report_20200304_0.pdf.

[52]

FAO/IAEA, 2nd RCM Report (2021) Second Research Coordination Meeting Joint FAO/IAEA Division of Nuclear Techniques in Food and Agriculture. Research Coordination Meeting on Generic Approach for the Development of Genetic Sexing Strains for SIT Applications. Vienna International Centre, Vienna, Austria. https://www.iaea.org/sites/default/files/21/11/d44003-crp_rcm2-report.pdf.

[53]

FAO/IAEA, 3rd RCM Report (2023) Third Research Coordination Meeting Joint FAO/IAEA Division of Nuclear Techniques in Food and Agriculture. Research Coordination Meeting on Generic Approach for the Development of Genetic Sexing Strains for SIT Applications. Vienna International Centre, Vienna, Austria. https://www.iaea.org/sites/default/files/23/08/d44003_crp_rcm_3_report_for_website_final_20230713.pdf.

[54]

FAO/IAEA/USDA (2019) Product Quality Control for Sterile Mass-Reared and Released Tephritid Fruit Flies, Version 7.0. IAEA, Vienna, Austria.

[55]

Feng, X., Kambic, L., Nishimoto, J.H.K., Reed, F.A., Denton, J.A., Sutton, J.T. et al. (2021) Evaluation of gene knockouts by CRISPR as potential targets for the genetic engineering of the mosquito Culex quinquefasciatus. The CRISPR Journal, 4, 595–608.

[56]

Fisher, K. and Caceres, C. (2000) A filter rearing system for mass reared genetic sexing strains of Mediterranean fruit fly (Diptera: Tephritidae). Area-Wide Control of Fruit Flies and Other Insect Pests. Joint Proceedings of the International Conference on Area-Wide Control of Insect Pests. 28 May-2 June, 1998 and the Fifth International Symposium on Fruit Flies of Economic Importance, Penang, Malaysia, 1−5 June, 1998, pp. 543–550. Penerbit Universiti Sains Malaysia, Pulau Pinang, Malaysia.

[57]

Flores, S., Montoya, P., Toledo, J., Enkerlin, W. and Liedo, P. (2014) Estimation of populations and sterility induction in Anastrepha ludens (Diptera: Tephritidae) fruit flies. Journal of Economic Entomology, 107, 1502–1507.

[58]

Franz, G. (2002) Recombination between homologous autosomes in medfly (Ceratitis capitata) males: type-1 recombination and the implications for the stability of genetic sexing strains. Genetica, 116, 73–84.

[59]

Franz, G., Bourtzis, K. and Cáceres, C. (2021) Practical and operational genetic sexing systems based on classical genetic approaches in fruit flies, an example for other species amenable to large-scale rearing for the sterile insect technique. Sterile Insect Technique (eds. V.A Dyck, J. Hendrichs & A.S. Robinson), pp. 575–604. CRC Press, Taylor and Francis Group, LLC, Florida, USA.

[60]

Franz, G., Willhoeft, U., Kerremans, P., Hendrichs, J. and Rendon, P. (1997) Development and application of genetic sexing systems for the Mediterranean fruit fly based on a temperature sensitive lethal mutation. Final Research Co-Ordination Meeting on Evaluation of Genetically Altered Medflies for Use in Sterile Insect Technique Programmes. Clearwater, FL (United States), 11–13 Jun 1994. pp. 85–95. IAEA, Austria

[61]

Fu, G., Condon, K.C., Epton, M.J., Gong, P., Jin, L., Condon, G.C. et al. (2007) Female-specific insect lethality engineered using alternative splicing. Nature Biotechnology, 25, 353–357.

[62]

Fu, G., Lees, R.S., Nimmo, D., Aw, D., Jin, L., Gray, P. et al. (2010) Female-specific flightless phenotype for mosquito control. Proceedings of the National Academy of Sciences USA, 107, 4550–4554.

[63]

Gamez, S., Chaverra-Rodriguez, D., Buchman, A., Kandul, N.P., Mendez-Sanchez, S.C., Bennett, J.B. et al. (2021) Exploiting a Y chromosome-linked Cas9 for sex selection and gene drive. Nature Communications, 12, 7202.

[64]

Gilles, J.R.L., Schetelig, M.F., Scolari, F., Marec, F., Capurro, M.L., Franz, G. et al. (2014) Towards mosquito sterile insect technique programmes: exploring genetic, molecular, mechanical and behavioural methods of sex separation in mosquitoes. Acta Tropica, 132, S178–S187.

[65]

Gong, P., Epton, M.J., Fu, G., Scaife, S., Hiscox, A., Condon, K.C. et al. (2005) A dominant lethal genetic system for autocidal control of the Mediterranean fruitfly. Nature Biotechnology, 23, 453–456.

[66]

Gossen, M. and Bujard, H. (1992) Tight control of gene expression in mammalian cells by tetracycline-responsive promoters. Proceedings of the National Academy of Sciences USA, 89, 5547–5551.

[67]

Gossen, M., Freundlieb, S., Bender, G., Müller, G., Hillen, W. and Bujard, H. (1995) Transcriptional activation by tetracyclines in mammalian cells. Science, 268, 1766–1769.

[68]

Häcker, I., Bourtzis, K. and Schetelig, M.F. (2021) Applying modern molecular technologies in support of the sterile insect technique. Sterile Insect Technique (eds. V.A. Dyck, J. Hendrichs & A.S. Robinson), pp. 657–702. CRC Press, Taylor and Francis Group, LLC, Florida, USA.

[69]

Hall, A.B., Basu, S., Jiang, X., Qi, Y., Timoshevskiy, V.A., Biedler, J.K. et al. (2015) A male-determining factor in the mosquito Aedes aegypti. Science, 348, 1268–1270.

[70]

Hammond, A., Galizi, R., Kyrou, K., Simoni, A., Siniscalchi, C., Katsanos, D. et al. (2016) A CRISPR-Cas9 gene drive system targeting female reproduction in the malaria mosquito vector Anopheles gambiae. Nature Biotechnology, 34, 78–83.

[71]

Harris, A.F., McKemey, A.R., Nimmo, D., Curtis, Z., Black, I., Morgan, S.A. et al. (2012) Successful suppression of a field mosquito population by sustained release of engineered male mosquitoes. Nature Biotechnology, 30, 828–830.

[72]

Harris, A.F., Nimmo, D., McKemey, A.R., Kelly, N., Scaife, S., Donnelly, C.A. et al. (2011) Field performance of engineered male mosquitoes. Nature Biotechnology, 29, 1034–1037.

[73]

Hawkins, N.J., Bass, C., Dixon, A. and Neve, P. (2019) The evolutionary origins of pesticide resistance. Biological Reviews, 94, 135–155.

[74]

Heinrich, J.C. and Scott, M.J. (2000) A repressible female-specific lethal genetic system for making transgenic insect strains suitable for a sterile-release program. Proceedings of the National Academy of Sciences USA, 97, 8229–8232.

[75]

Hendrichs, J., Franz, G. and Rendon, P. (1995) Increased effectiveness and applicability of the sterile insect technique through male-only releases for control of Mediterranean fruit flies during fruiting seasons. Journal of Applied Entomology, 119, 371–377.

[76]

Hendrichs, J., Pereira, R. and Vreysen, M.J.B. (2021) Area-wide Integrated Pest Management: Development and Field Application. CRC Press.

[77]

Herbillon, F., Diouf, E.G., Brévault, T., Haramboure, M., Fellous, S. and Piou, C. (2024) Life history traits of the target pest and transmission routes of the biocide are critical for the success of the boosted sterile insect technique. Current Research in Insect Science, 6, 100101.

[78]

Hibino, Y. and Iwahashi, O. (1991) Appearance of wild females unreceptive to sterilized males on Okinawa Is. in the eradication program of the melon fly, Dacus cucurbitae Coquillett (Diptera:Tephritidae). Applied Entomology and Zoology, 26, 265–270.

[79]

Hsu, P.D., Lander, E.S. and Zhang, F. (2014) Development and applications of CRISPR-Cas9 for genome engineering. Cell, 157, 1262–1278.

[80]

Ioannidou, C., Gregoriou, M.E., Schetelig, M.F., Drosopoulou, E., Mathiopoulos, K.D. and Bourtzis, K. (2025) CRISPR/Cas9-based white pupae mutant lines in Bactrocera spp. for sterile insect technique applications. Insect Science, https://doi.org/10.1111/1744-7917.70190.

[81]

Johnson, B.J., Mitchell, S.N., Paton, C.J., Stevenson, J., Staunton, K.M., Snoad, N. et al. (2017) Use of rhodamine B to mark the body and seminal fluid of male Aedes aegypti for mark-release-recapture experiments and estimating efficacy of sterile male releases. PLoS Neglected Tropical Diseases, 11, e0005902.

[82]

Kaiser, P.E., Seawright, J.A., Dame, D.A. and Joslyn, D.J. (1978) Development of a genetic sexing system for Anopheles albimanus. Journal of Economic Entomology, 71, 766–771.

[83]

Kalajdzic, P. and Schetelig, M.F. (2017) CRISPR/Cas-mediated gene editing using purified protein in Drosophila suzukii. Entomologia Experimentalis et Applicata, 164, 350–362.

[84]

Kandul, N.P., Liu, J., Hsu, A.D., Hay, B.A. and Akbari, O.S. (2020) A drug-inducible sex-separation technique for insects. Nature Communications, 11, 2106.

[85]

Kerremans, P. and Franz, G. (1995) Isolation and cytogenetic analyses of genetic sexing strains for the medfly, Ceratitis capitata. Theoretical and Applied Genetics, 91, 255–261.

[86]

Kerremans, P., Bourtzis, K. and Zacharopoulou, A. (1990) Cytogenetic analysis of three genetic sexing strains of Ceratitits capitata. Theoretical and Applied Genetics, 80, 177–182.

[87]

Kioy, D., Jannin, J. and Mattock, N. (2004) Focus: human African trypanosomiasis. Nature Reviews Microbiology, 2, 186.

[88]

Kistler, K.E., Vosshall, L.B. and Matthews, B.J. (2015) Genome engineering with CRISPR-Cas9 in the mosquito Aedes aegypti. Cell Reports, 11, 51–60.

[89]

Klassen, W., Curtis, C.F. and Hendrichs, J. (2021) History of the sterile insect technique. Sterile Insect Technique. Principles and Practice in Area-Wide Integrated Pest Management. 2nd edn. (eds. V.A. Dyck, J. Hendrichs & A.S. Robinson), pp. 1–44. CRC Press, Taylor and Francis Group, LLC, Florida, USA.

[90]

Knipling, E.F. (1955) Possibilities of insect control or eradication through the use of sexually sterile males. Journal of Economic Entomology, 48, 459–462.

[91]

Koskinioti, P., Augustinos, A.A., Carvalho, D.O., Misbah-ul-Haq, M., Pillwax, G., de la Fuente, L.D. et al. (2021) Genetic sexing strains for the population suppression of the mosquito vector Aedes aegypti. Philosophical Transactions of the Royal Society B, 376, 20190808.

[92]

Koutroumpa, F.A., Monsempes, C., François, M.C., de Cian, A., Royer, C., Concordet, J.P. et al. (2016) Heritable genome editing with CRISPR/Cas9 induces anosmia in a crop pest moth. Scientific Reports, 6, 29620.

[93]

Krzywinska, E., Dennison, N.J., Lycett, G.J. and Krzywinski, J. (2016) A maleness gene in the malaria mosquito Anopheles gambiae. Science, 353, 67–69.

[94]

Kuba, H., Kohama, T., Kakinohana, H., Yamagishi, M., Kinjo, K., Sokei, Y. et al. (2020) The successful eradication programs of the melon fly in Okinawa. Fruit Fly Pests (eds. B.A. McPheron & G.J. Steck), pp. 543–550. CRC Press, Taylor and Francis Group, LLC, Florida, USA.

[95]

Labbé, G.M.C., Scaife, S., Morgan, S.A., Curtis, Z.H. and Alphey, L. (2012) Female-specific flightless (fsRIDL) phenotype for control of Aedes albopictus. PLoS Neglected Tropical Diseases, 6, e1724.

[96]

Leftwich, P.T., Spurgin, L.G., Harvey-Samuel, T., Thomas, C.J.E., Paladino, L.C., Edgington, M.P. et al. (2020) Genetic pest management and the background genetics of release strains. Philosophical Transactions of the Royal Society B: Biological Sciences, 376, 20190805.

[97]

Li, J. and Handler, A.M. (2019) CRISPR/Cas9-mediated gene editing in an exogenous transgene and an endogenous sex determination gene in the Caribbean fruit fly, Anastrepha suspensa. Gene, 691, 160–166.

[98]

Li, J. and Handler, A.M. (2017) Temperature-dependent sex-reversal by a transformer-2 gene-edited mutation in the spotted wing drosophila, Drosophila suzukii. Scientific Reports, 7, 12363.

[99]

Lim, J.T., Bansal, S., Chong, C.S., Dickens, B., Ng, Y., Deng, L. et al. (2024) Efficacy of Wolbachia-mediated sterility to reduce the incidence of dengue: a synthetic control study in Singapore. The Lancet Microbe, 5, e422–e432.

[100]

Lindquist, D.A., Abusowa, M. and Hall, M.J.R. (1992) The New World screwworm fly in Libya: a review of its introduction and eradication. Medical and Veterinary Entomology, 6, 2–8.

[101]

Lines, J.D. and Curtis, C.F. (1985) Genetic sexing systems in Anopheles arabiensis patton (Diptera: Culicidae). Journal of Economic Entomology, 78, 848–851.

[102]

Lino, C.A., Harper, J.C., Carney, J.P. and Timlin, J.A. (2018) Delivering CRISPR: a review of the challenges and approaches. Drug Delivery, 25, 1234–1257.

[103]

Liu, P., Jin, B., Li, X., Zhao, Y., Gu, J., Biedler, J.K. et al. (2020) Nix is a male-determining factor in the Asian tiger mosquito Aedes albopictus. Insect Biochemistry and Molecular Biology, 118, 103311.

[104]

Lutrat, C., Burckbuchler, M., Olmo, R.P., Beugnon, R., Fontaine, A., Akbari, O.S. et al. (2023) Combining two genetic sexing strains allows sorting of non-transgenic males for Aedes genetic control. Communications Biology, 6, 646.

[105]

Lutrat, C., Giesbrecht, D., Marois, E., Whyard, S., Baldet, T. and Bouyer, J. (2019) Sex sorting for pest control: it's raining men! Trends in Parasitology, 35, 649–662.

[106]

Lutrat, C., Olmo, R.P., Baldet, T., Bouyer, J. and Marois, E. (2022) Transgenic expression of Nix converts genetic females into males and allows automated sex sorting in Aedes albopictus. Communications Biology, 5, 210.

[107]

Ma, S., Chang, J., Wang, X., Liu, Y., Zhang, J., Lu, W. et al. (2014) CRISPR/Cas9 mediated multiplex genome editing and heritable mutagenesis of BmKu70 in Bombyx mori. Scientific Reports, 4, 4489.

[108]

Malcolm, C.A. and Mali, P. (1986) Genetic sexing of Anopheles stephensi with the larval morphological mutant Bl. Genetica, 70, 37–42.

[109]

Mallapaty, S. (2019) Australian gene-editing rules adopt ‘middle ground’. Nature, https://doi.org/10.1038/d41586-019-01282-8.

[110]

Markert, M.J., Zhang, Y., Enuameh, M.S., Reppert, S.M., Wolfe, S.A. and Merlin, C. (2016) Genomic access to monarch migration using TALEN and CRISPR/Cas9-mediated targeted mutagenesis. G3 Genes|Genomes|Genetics, 6, 905–915.

[111]

Martín-Park, A., Che-Mendoza, A., Contreras-Perera, Y., Pérez-Carrillo, S., Puerta-Guardo, H., Villegas-Chim, J. et al. (2022) Pilot trial using mass field-releases of sterile males produced with the incompatible and sterile insect techniques as part of integrated Aedes aegypti control in Mexico. PLoS Neglected Tropical Diseases, 16, e0010324.

[112]

Mavragani-Tsipidou, P., Zacharopoulou, A., Drosopoulou, E., Augustinos, A.A., Bourtzis, K. and Marec, F. (2014) Protocols for cytogenetic mapping of arthropod genomes: Tephritid fruit flies of economic importance. Protocols for Cytogenetic Mapping of Arthropod Genomes (ed. I. Sakharov), pp. 1–62. CRC Press, Taylor and Francis Group, LLC, Florida, USA.

[113]

McClelland, G.A.H. (1966) Sex-linkage at two loci affecting eye pigment in the mosquito Aedes aegypti (diptera: Culicidae). Canadian Journal of Genetics and Cytology, 8, 192–198.

[114]

McCombs, S.D. and Saul, S.H. (1995) Translocation-based genetic sexing system for the oriental fruit fly (Diptera: Tephritidae) based on pupal color dimorphism. Annals of the Entomological Society of America, 88, 695–698.

[115]

McDonald, P.T. and Asman, S.M. (1982) A genetic-sexing strain based on malathion resistance for Culex tarsalis. Mosquito News, 42, 531–536.

[116]

McFarlane, G.R., Whitelaw, C.B.A. and Lillico, S.G. (2018) CRISPR-based gene drives for pest control. Trends in Biotechnology, 36, 130–133.

[117]

McInnis, D.O., Lance, D.R. and Jackson, C.G. (1996) Behavioral resistance to the sterile insect technique by mediterranean fruit fly (Diptera: Tephritidae) in Hawaii. Annals of the Entomological Society of America, 89, 739–744.

[118]

McInnis, D.O., Tam, S., Grace, C. and Miyashita, D. (1994) Population suppression and sterility rates induced by variable sex ratio, sterile insect releases of Ceratitis capitata (Diptera: Tephritidae) in Hawaii. Annals of the Entomological Society of America, 87, 231–240.

[119]

McInnis, D.O., Tam, S., Lim, R., Komatsu, J., Kurashima, R. and Albrecht, C. (2004) Development of a pupal color-based genetic sexing strain of the melon fly, Bactrocera cucurbitae (Coquillett) (Diptera: Tephritidae). Annals of the Entomological Society of America, 97, 1026–1033.

[120]

Meccariello, A., Monti, S.M., Romanelli, A., Colonna, R., Primo, P., Inghilterra, M.G. et al. (2017) Highly efficient DNA-free gene disruption in the agricultural pest Ceratitis capitata by CRISPR-Cas9 ribonucleoprotein complexes. Scientific Reports, 7, 10061.

[121]

Meccariello, A., Salvemini, M., Primo, P., Hall, B., Koskinioti, P., Dalíková, M. et al. (2019) Maleness-on-the-Y (MoY) orchestrates male sex determination in major agricultural fruit fly pests. Science, 365, 1457–1460.

[122]

Meza, J.S., Bourtzis, K., Zacharopoulou, A., Gariou-Papalexiou, A. and Cáceres, C. (2020) Development and characterization of a pupal-colour based genetic sexing strain of Anastrepha fraterculus sp. 1 (Diptera: Tephritidae). BMC Genetics, 21, 134.

[123]

Meza, J.S., Schetelig, M.F., Zepeda-Cisneros, C.S. and Handler, A.M. (2014) Male-specific Y-linked transgene markers to enhance biologically-based control of the Mexican fruit fly, Anastrepha ludens (Diptera: Tephritidae). BMC Genomic Data, 15, S4.

[124]

Misbah-ul-Haq, M., Augustinos, A.A., Carvalho, D.O., Duran de la Fuente, L. and Bourtzis, K. (2022a) The effect of an irradiation-induced recombination suppressing inversion on the genetic stability and biological quality of a white eye-based Aedes aegypti genetic sexing strain. Insects, 13, 946.

[125]

Misbah-ul-Haq, M., Carvalho, D.O., Duran De La Fuente, L., Augustinos, A.A. and Bourtzis, K. (2022b) Genetic stability and fitness of Aedes Aegypti red-eye genetic sexing strains with pakistani genomic background for sterile insect technique applications. Frontiers in Bioengineering and Biotechnology, 10, 871703.

[126]

Mojica, F.J.M. and Rodriguez-Valera, F. (2016) The discovery of CRISPR in archaea and bacteria. The FEBS Journal, 283, 3162–3169.

[127]

Munstermann, L.E. and Craig, G.B. (1979) Genetics of Aedes aegypti: updating the linkage map. Journal of Heredity, 70, 291–296.

[128]

Mysore, K., Sun, L., Hapairai, L.K., Wang, C.W., Roethele, J.B., Igiede, J. et al. (2021) A broad-based mosquito yeast interfering RNA pesticide targeting Rbfox1 represses Notch signaling and kills both larvae and adult mosquitoes. Pathogens, 10, 1251.

[129]

Nguyen, T.N.M., Choo, A. and Baxter, S.W. (2021) Lessons from Drosophila: engineering genetic sexing strains with temperature-sensitive lethality for sterile insect technique applications. Insects, 12, 243.

[130]

Nidhi, S., Anand, U., Oleksak, P., Tripathi, P., Lal, J.A., Thomas, G. et al. (2021) Novel CRISPR–Cas systems: an updated review of the current achievements, applications, and future research perspectives. International Journal of Molecular Sciences, 22, 3327.

[131]

Nikolouli, K., Compton, A., Tu, Z.J. and Bourtzis, K. (2025) Evaluation of ebony as a potential selectable marker for genetic sexing in Aedes aegypti. Parasites & Vectors, 18, 76.

[132]

Niyazi, N., Caceres, C., Delprat, A., Wornoayporn, V., Santos, E.R., Franz, G. et al. (2005) Genetics and mating competitiveness of Ceratitis capitata (Diptera: Tephritidae) strains carrying the marker sergeant, Sr2. Annals of the Entomological Society of America, 98, 119–125.

[133]

Ntoyi, N.L., Mashatola, T., Bouyer, J., Kraupa, C., Maiga, H., Mamai, W. et al. (2022) Life-history traits of a fluorescent Anopheles arabiensis genetic sexing strain introgressed into South African genomic background. Malaria Journal, 21, 254.

[134]

Ogaugwu, C.E., Schetelig, M.F. and Wimmer, E.A. (2013) Transgenic sexing system for Ceratitis capitata (Diptera: Tephritidae) based on female-specific embryonic lethality. Insect Biochemistry and Molecular Biology, 43, 1–8.

[135]

Oye, K.A., Esvelt, K., Appleton, E., Catteruccia, F., Church, G., Kuiken, T. et al. (2014) Regulating gene drives. Science, 345, 626–628.

[136]

Papathanos, P.A., Bossin, H.C., Benedict, M.Q., Catteruccia, F., Malcolm, C.A., Alphey, L. et al. (2009) Sex separation strategies: past experience and new approaches. Malaria Journal, 8, S5.

[137]

Papathanos, P.A., Bourtzis, K., Tripet, F., Bossin, H., Virginio, J.F., Capurro, M.L. et al. (2018) A perspective on the need and current status of efficient sex separation methods for mosquito genetic control. Parasites & Vectors, 11, 654.

[138]

Paulo, D.F., Nguyen, T.N.M., Ward, C.M., Corpuz, R.L., Kauwe, A.N., Rendon, P. et al. (2025) Functional genomics implicates ebony in the black pupae phenotype of tephritid fruit flies. Communications Biology, 8, 60.

[139]

Phuc, H.K., Andreasen, M.H., Burton, R.S., Vass, C., Epton, M.J., Pape, G. et al. (2007) Late-acting dominant lethal genetic systems and mosquito control. BMC Biology, 5, 11.

[140]

Potter, C.J., Tasic, B., Russler, E.V., Liang, L. and Luo, L. (2010) The Q system: a repressible binary system for transgene expression, lineage tracing, and mosaic analysis. Cell, 141, 536–548.

[141]

Prates, L.H.F., Aumann, R.A., Sievers, I., Rehling, T. and Schetelig, M.F. (2025) Functional validation of a white pupae minimal gene construct in Ceratitis capitata (Diptera: Tephritidae). Insect Science, https://doi.org/10.1111/1744-7917.70058.

[142]

Prates, L.H.F., Fiebig, J., Schlosser, H., Liapi, E., Rehling, T., Lutrat, C. et al. (2024) Challenges of robust RNAi-mediated gene silencing in Aedes Mosquitoes. International Journal of Molecular Sciences, 25, 5218.

[143]

Ramírez-Santos, E., Rendon, P., Gouvi, G., Zacharopoulou, A., Bourtzis, K., Cáceres, C. et al. (2021) A novel genetic sexing strain of Anastrepha ludens for cost-effective sterile insect technique applications: improved genetic stability and rearing efficiency. Insects, 12, 499.

[144]

Ren, X., Sun, J., Housden, B.E., Hu, Y., Roesel, C., Lin, S. et al. (2013) Optimized gene editing technology for Drosophila melanogaster using germ line-specific Cas9. Proceedings of the National Academy of Sciences USA, 110, 19012–19017.

[145]

Robinson, A.S. (2002) Genetic sexing strains in medfly, Ceratitis Capitata, sterile insect technique programmes. Genetica, 116, 5–13.

[146]

Roger, C.R. (2023) NGT: The European Commission plays a “simultaneously” approach. European Scientist. http://www.europeanscientist.com/en/features/ngt-the-european-commission-plays-a-simultaneously-approach/.

[147]

Roseman, R.R., Pirrotta, V. and Geyer, P.K. (1993) The su(Hw) protein insulates expression of the Drosophila melanogaster white gene from chromosomal position-effects. The EMBO Journal, 12, 435–442.

[148]

Rössler, Y. (1979) The genetics of the Mediterranean fruit fly: a “white pupae” mutant. Annals of the Entomological Society of America, 72, 583–585.

[149]

Schetelig, M.F., Caceres, C., Zacharopoulou, A., Franz, G. and Wimmer, E.A. (2009) Conditional embryonic lethality to improve the sterile insect technique in Ceratitis capitata (Diptera: Tephritidae). BMC Biology, 7, 4.

[150]

Schetelig, M.F. and Handler, A.M. (2012) Strategy for enhanced transgenic strain development for embryonic conditional lethality in Anastrepha suspensa. Proceedings of the National Academy of Sciences USA, 109, 9348–9353.

[151]

Schetelig, M.F., Schwirz, J. and Yan, Y. (2021) A transgenic female killing system for the genetic control of Drosophila suzukii. Scientific Reports, 11, 12938.

[152]

Schetelig, M.F., Targovska, A., Meza, J.S., Bourtzis, K. and Handler, A.M. (2016) Tetracycline-suppressible female lethality and sterility in the Mexican fruit fly, Anastrepha ludens. Insect Molecular Biology, 25, 500–508.

[153]

Schneeberger, K. (2014) Using next-generation sequencing to isolate mutant genes from forward genetic screens. Nature Reviews Genetics, 15, 662–676.

[154]

Sharma, A., Heinze, S.D., Wu, Y., Kohlbrenner, T., Morilla, I., Brunner, C. et al. (2017a) Male sex in houseflies is determined by Mdmd, a paralog of the generic splice factor gene CWC22. Science, 356, 642–645.

[155]

Sharma, A., Kumar, V., Shahzad, B., Tanveer, M., Sidhu, G.P.S., Handa, N. et al. (2019) Worldwide pesticide usage and its impacts on ecosystem. SN Applied Sciences, 1, 1446.

[156]

Sharma, S., Kooner, R. and Arora, R. (2017b) Insect pests and crop losses. Breeding Insect Resistant Crops for Sustainable Agriculture (Eds. R. Arora & S. Sandhu), pp. 45–66. Springer, Singapore.

[157]

Shen, X., Song, S., Li, C. and Zhang, J. (2022) Synonymous mutations in representative yeast genes are mostly strongly non-neutral. Nature, 606, 725–731.

[158]

Shetty, N.J. (1987) Genetic sexing system for the preferential elimination of females in Culex quinquefasciatus. Journal of the American Mosquito Control Association, 3, 84–86.

[159]

Silicheva, M., Golovnin, A., Pomerantseva, E., Parshikov, A., Georgiev, P. and Maksimenko, O. (2010) Drosophila mini-white model system: new insights into positive position effects and the role of transcriptional terminators and gypsy insulator in transgene shielding. Nucleic Acids Research, 38, 39–47.

[160]

Sim, S.B., Kauwe, A.N., Ruano, R.E.Y., Rendon, P. and Geib, S.M. (2019) The ABCs of CRISPR in Tephritidae: developing methods for inducing heritable mutations in the genera Anastrepha, Bactrocera and Ceratitis. Insect Molecular Biology, 28, 277–289.

[161]

Singh, S., Rahangdale, S., Pandita, S., Saxena, G., Upadhyay, S.K., Mishra, G. et al. (2022) CRISPR/Cas9 for insect pests management: a comprehensive review of advances and applications. Agriculture, 12, 1896.

[162]

Sollazzo, G., Gouvi, G., Nikolouli, K., Aumann, R.A., Djambazian, H., Whitehead, M.A. et al. (2023) Genomic and cytogenetic analysis of the Ceratitis capitata temperature-sensitive lethal region. G3 Genes|Genomes|Genetics, 13, jkad074.

[163]

Sollazzo, G., Nikolouli, K., Gouvi, G., Aumann, R.A., Schetelig, M.F. and Bourtzis, K. (2024) Deep orange gene editing triggers temperature-sensitive lethal phenotypes in Ceratitis capitata. BMC Biotechnology, 24, 7.

[164]

Spinner, S.A.M., Barnes, Z.H., Puinean, A.M., Gray, P., Dafa'alla, T., Phillips, C.E. et al. (2022) New self-sexing Aedes aegypti strain eliminates barriers to scalable and sustainable vector control for governments and communities in dengue-prone environments. Frontiers in Bioengineering and Biotechnology, 10, 975786.

[165]

Sproule, A., Broughton, S., De Lima, F., Hardie, D., Monzu, N. and Woods, B. (2001) The Fight Against Fruit Flies in Western Australia. Department of Primary Industries and Regional Development, Western Australia, Perth. Bulletin, 4504.

[166]

Szendrei, Z. and Rodriguez-Saona, C. (2010) A meta-analysis of insect pest behavioral manipulation with plant volatiles. Entomologia Experimentalis et Applicata, 134, 201–210.

[167]

Teng, F., Guo, F., Feng, J., Lu, Y. and Qi, Y. (2024) Distribution analysis of TRH in Bactrocera dorsalis using a CRISPR/Cas9-mediated reporter knock-in strain. Insect Molecular Biology, 33, 283–292.

[168]

Thomas, D.D., Donnelly, C.A., Wood, R.J. and Alphey, L.S. (2000) Insect population control using a dominant, repressible, lethal genetic system. Science, 287, 2474–2476.

[169]

Tomaszkiewicz, M., Medvedev, P. and Makova, K.D. (2017) Y and W chromosome assemblies: approaches and discoveries. Trends in Genetics, 33, 266–282.

[170]

Tudi, M., Daniel Ruan, H., Wang, L., Lyu, J., Sadler, R., Connell, D. et al. (2021) Agriculture development, pesticide application and its impact on the environment. International Journal of Environmental Research and Public Health, 18, 1112.

[171]

Van den Bossche, P., de La Rocque, S., Hendrickx, G. and Bouyer, J. (2010) A changing environment and the epidemiology of tsetse-transmitted livestock trypanosomiasis. Trends in Parasitology, 26, 236–243.

[172]

Verhulst, N.O., Loonen, J.A. and Takken, W. (2013) Advances in methods for colour marking of mosquitoes. Parasites & Vectors, 6, 200.

[173]

Vogel, E., Santos, D., Mingels, L., Verdonckt, T.W. and Broeck, J.V. (2019) RNA interference in insects: protecting beneficials and controlling pests. Frontiers in Physiology, 9, 1912.

[174]

Vreysen, M.J., Hendrichs, J. and Enkerlin, W.R. (2006a) The sterile insect technique as a component of sustainable area-wide integrated pest management of selected horticultural insect pests. Journal of Fruit and Ornamental Plant Research, 14, 107.

[175]

Vreysen, M.J.B., Barclay, H.J. and Hendrichs, J. (2006b) Modeling of preferential mating in areawide control programs that integrate the release of strains of sterile males only or both sexes. Annals of the Entomological Society of America, 99, 607–616.

[176]

Vreysen, M.J.B., Saleh, K.M., Ali, M.Y., Abdulla, A.M., Zhu, Z.R., Juma, K.G. et al. (2000) Glossina austeni (Diptera: Glossinidae) eradicated on the island of unguja, zanzibar, using the sterile insect technique. Journal of Economic Entomology, 93, 123–135.

[177]

Wang, J.Y. and Doudna, J.A. (2023) CRISPR technology: a decade of genome editing is only the beginning. Science, 379, eadd8643.

[178]

Wappner, P., Kramer, K.J., Hopkins, T.L., Merritt, M., Schaefer, J. and Quesada-AlluÉ, L.A. (1995) White pupa: a Ceratitis capitata mutant lacking catecholamines for tanning the puparium. Insect Biochemistry and Molecular Biology, 25, 365–373.

[179]

Ward, C.M., Aumann, R.A., Whitehead, M.A., Nikolouli, K., Leveque, G., Gouvi, G. et al. (2021) White pupae phenotype of tephritids is caused by parallel mutations of a MFS transporter. Nature Communications, 12, 491.

[180]

WHO (2024) Vector-borne diseases. https://www.who.int/news-room/fact-sheets/detail/vector-borne-diseases#:~:text=Vector-borne%20diseases%20are%20human%20illnesses%20caused%20by%20parasites%2C,viruses%20and%20bacteria%20that%20are%20transmitted%20by%20vectors.

[181]

Whyard, S., Erdelyan, C.N., Partridge, A.L., Singh, A.D., Beebe, N.W. and Capina, R. (2015) Silencing the buzz: a new approach to population suppression of mosquitoes by feeding larvae double-stranded RNAs. Parasites & Vectors, 8, 96.

[182]

Willhoeft, U. and Franz, G. (1996) Identification of the sex-determining region of the Ceratitis capitata Y chromosome by deletion mapping. Genetics, 144, 737–745.

[183]

Wise de Valdez, M.R., Nimmo, D., Betz, J., Gong, H.F., James, A.A., Alphey, L. et al. (2011) Genetic elimination of dengue vector mosquitoes. Proceedings of the National Academy of Sciences USA, 108, 4772–4775.

[184]

Wray-Cahen, D., Hallerman, E. and Tizard, M. (2024) Global regulatory policies for animal biotechnology: overview, opportunities and challenges. Frontiers in Genome Editing, 6, 1467080.

[185]

Xu, X., Harvey-Samuel, T., Yang, J., You, M. and Alphey, L. (2021) CRISPR/Cas9-based functional characterization of the pigmentation gene ebony in Plutella xylostella. Insect Molecular Biology, 30, 615–623.

[186]

Yamada, H., Benedict, M.Q., Malcolm, C.A., Oliva, C.F., Soliban, S.M. and Gilles, J.R. (2012) Genetic sex separation of the malaria vector, Anopheles arabiensis, by exposing eggs to dieldrin. Malaria Journal, 11, 208.

[187]

Yamada, H., Vreysen, M.J.B., Bourtzis, K., Tschirk, W., Chadee, D.D. and Gilles, J.R.L. (2015) The Anopheles arabiensis genetic sexing strain ANO IPCL1 and its application potential for the sterile insect technique in integrated vector management programmes. Acta Tropica, 142, 138–144.

[188]

Yan, Y., Aumann, R.A., Häcker, I. and Schetelig, M.F. (2023) CRISPR-based genetic control strategies for insect pests. Journal of Integrative Agriculture, 22, 651–668.

[189]

Yan, Y. and Scott, M.J. (2015) A transgenic embryonic sexing system for the Australian sheep blow fly Lucilia cuprina. Scientific Reports, 5, 16090.

[190]

Yan, Y. and Scott, M.J. (2020) Building a transgenic sexingstrain for genetic control of the Australian sheep blow fly Lucilia cuprina using two lethaleffectors. BMC Genetics, 21, 141.

[191]

Yu, Z., Chen, H., Liu, J., Zhang, H., Yan, Y., Zhu, N. et al. (2014) Various applications of TALEN- and CRISPR/Cas9-mediated homologous recombination to modify the Drosophila genome. Biology Open, 3, 271–280.

[192]

Zacharopoulou, A. (1987) Cytogenetic analysis of mitotic and salivary gland chromosomes in the Medfly Ceratitis capitata. Genome, 29, 67–71.

[193]

Zacharopoulou, A., Augustinos, A.A., Drosopoulou, E., Tsoumani, K.T., Gariou-Papalexiou, A., Franz, G. et al. (2017) A review of more than 30 years of cytogenetic studies of Tephritidae in support of sterile insect technique and global trade. Entomologia Experimentalis et Applicata, 164, 204–225.

[194]

Zacharopoulou, A., Bourtzis, K. and Kerremans, P. (1991) A comparison of polytene chromosomes in salivary glands and orbital bristle trichogen cells in Ceratitis capitata. Genome, 34, 215–219.

[195]

Zepeda-Cisneros, C.S., Meza Hernández, J.S., García-Martínez, V., Ibañez-Palacios, J., Zacharopoulou, A. and Franz, G. (2014) Development, genetic and cytogenetic analyses of genetic sexing strains of the Mexican fruit fly, Anastrepha ludens Loew (Diptera: Tephritidae). BMC Genomic Data, 15, S1.

[196]

Zhao, S., Xing, Z., Liu, Z., Liu, Y., Liu, X., Chen, Z. et al. (2019) Efficient somatic and germline genome engineering of Bactrocera dorsalis by the CRISPR/Cas9 system. Pest Management Science, 75, 1921–1932.

[197]

Zheng, X., Zhang, D., Li, Y., Yang, C., Wu, Y., Liang, X. et al. (2019) Incompatible and sterile insect techniques combined eliminate mosquitoes. Nature, 572, 56–61.

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2025 International Atomic Energy Agency and The Author(s). Insect Science published by John Wiley & Sons Australia, Ltd on behalf of Institute of Zoology, Chinese Academy of Sciences.

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