Ecological geneticsEcological geneticsЭкологическая генетика1811-09322411-9202Eco-Vector67734110.17816/ecogen677341AZONEWGenetic toxicologyГенетическая токсикологияReview ArticleMolecular genetic patterns of pyrethroid resistance in diamondback moth (Plutella xylostella; Linnaeus, 1758)Молекулярно-генетическая природа резистентности капустной моли Plutella xylostella (Linnaeus, 1758) к пиретроидамhttps://orcid.org/0000-0002-0308-6108EmelyanovDmitrii A.ЕмельяновДмитрий Александрович
Russian Federation
dimitriy.nord@yandex.ruhttps://orcid.org/0009-0005-4885-3904BogomazFeodor D.БогомазФедор Денисович
Russian Federation
pickayut2006@gmail.comhttps://orcid.org/0009-0009-3296-3868ShabanovaKsenia A.ШабановаКсения Алексеевна
Russian Federation
kotukkc@gmail.comhttps://orcid.org/0000-0001-8569-66653877-6598MatveevaTatiana V.МатвееваТатьяна Валерьевна
Russian Federation

Dr. Sci. (Biology), Professor

д-р биол. наук, профессор

radishlet@gmail.comAll-Russian Research Institute of Plant ProtectionВсероссийский научно-исследовательский институт защиты растенийNorth-Western State Medical University named after I.I. MechnikovСеверо-Западный государственный медицинский университет им. И.И. МечниковаPeter the Great Saint Petersburg Polytechnic UniversityСанкт-Петербургский политехнический университет Петра Великого05112025090220262343613741903202505112025Copyright ©; 2025, Eco-VectorCopyright ©; 2025, Эко-Вектор2025Eco-VectorЭко-Векторhttps://eco-vector.com/for_authors.php#07https://journals.eco-vector.com/ecolgenet/article/view/677341

The diamondback moth (Plutella xylostella; Linnaeus, 1758) is a globally significant pest of cruciferous crops, causing substantial economic losses. Resistance to pyrethroid insecticides, which are widely used for its control, has become a major issue. This review explores the molecular and genetic mechanisms underlying pyrethroid resistance in P. xylostella, focusing on mutations in the voltage-gated sodium channel gene (Pxpara), which is the primary target of pyrethroids. The review involved an analysis of P. xylostella populations from various regions, particularly in Asia and Australia, where resistance to pyrethroids is prevalent. Molecular techniques, including KASP assays and PCR analysis followed by sequencing, were employed to identify and characterize resistance-associated mutations in the Pxpara gene. Several key mutations in the Pxpara gene were identified, including T929I, M918I, L1014F, and F1020S, which are associated with pyrethroid resistance. These mutations were found to be widespread in Asian populations, with a high prevalence observed in China. An analysis of publications on resistance mechanisms in other insect species revealed resistance mutations at the same sites in a wide range of species, indicating shared mechanisms. The identified mutations in the Pxpara gene provide valuable markers for resistance detection. The development of diagnostic tools based on these findings is crucial for effective resistance management and sustainable pest control. The review also emphasizes the need for integrated pest management approaches to mitigate the spread of resistance and reduce reliance on chemical insecticides.

Капустная моль [Plutella xylostella (Linnaeus, 1758)] — глобально значимый вредитель крестоцветных культур, вызывающий значительные экономические потери. Устойчивость к пиретроидным инсектицидам, широко используемым для ее контроля, в настоящее время представляет серьезную проблему. В приведенном обзоре обобщены молекулярные и генетические механизмы, лежащие в основе устойчивости к пиретроидам у P. xylostella, с упором на мутации в гене потенциал-зависимого натриевого канала (Pxpara), который является основной мишенью пиретроидов. В данной работе представлен анализ популяций P. xylostella из различных регионов, особенно из Азии и Австралии, где была детально изучена молекулярная природа устойчивости к пиретроидам. Молекулярные методы, включая КАСП-анализ (KASP) и генотипирование на основе классической ПЦР с последующим секвенированием ее продуктов, были использованы для выявления и характеристики частот встречаемости мутаций в гене Pxpara, продукт которого является мишенью действия пиретроидов. Среди них можно выявить несколько ключевых SNP, приводящих к заменам аминокислот: T929I, M918I, L1014F и F1020S. Было обнаружено, что они широко распространены в азиатских популяциях, с высокой частотой встречаются в Китае. Анализ литературных данных о механизмах устойчивости у других видов насекомых позволил выявить мутации резистентности в тех же сайтах у большого количества видов, что указывает на ее общие механизмы — данные аминокислотные замены снижают чувствительность натриевого канала к пиретроидам, что приводит к устойчивости насекомых к данным веществам. Разработка ДНК-диагностикумов на основе этих данных имеет решающее значение для эффективного управления устойчивостью и успешной борьбы с вредителями.

diamondback mothpyrethroidsresistancesodium channelsmutationsmolecular diagnosticsresistance managementintegrated pest managementкапустная мольпиретроидырезистентностьнатриевые каналымутациимолекулярная диагностикауправление устойчивостьюинтегрированная защита растенийМинистерство науки и высшего образования Российской ФедерацииMinistry of Science and Higher Education of the Russian Federation075-15-2025-651The work was supported by the Ministry of Education and Science of the Russian Federation as part of the project “Chemical and Biological Control of Major Sucking Pests on Vegetable Crops” (Agreement No. 075-15-2025-651 of August 20, 2025)Работа выполнена при финансовой поддержке Минобрнауки России по проекту «Химический и биологический контроль основных сосущих вредителей на овощных культурах» (соглашение № 075-15-2025-651 от 20.08.2025)Talekar NS, Shelton AM. Biology, ecology, and management of the diamondback moth. Annu Rev Entomol. 1993;38(1):275–301. doi: 10.1146/annurev.en.38.010193.001423Mason P. Plutella xylostella (diamondback moth). CABI Compendium. 2022. doi: 10.1079/cabicompendium.42318Coulson SJ, Hodkinson ID, Webb NR, et al. Aerial colonization of high Arctic islands by invertebrates: the diamondback moth Plutella xylostella (Lepidoptera: Yponomeutidae) as a potential indicator species. Divers Distrib. 2002;8(6):327–334. doi: 10.1046/j.1472-4642.2002.00157.x EDN: BEUZOFShelton AM. Management of the diamondback moth: déjà vu all over again? The management of diamondback moth and other crucifer pests: Proceedings of the Fourth International Workshop 26–29 November 2001; Melbourne, Victoria, Australia. Editors N.M. Endersby, P.M. Ridland, The Regional Institute Gosford, N.S.W. 2004.Zalucki MP, Shabbir A, Silva R, et al. Estimating the economic cost of one of the world’s major insect pests, Plutella xylostella (Lepidoptera: Plutellidae): just how long is a piece of string? J Econ Entomol. 2012;105(4): 1115–1129. doi: 10.1603/EC12107Sparks TC, Crossthwaite AJ, Nauen R, et al. Insecticides, biologics, and nematicides: updates to IRAC’s mode of action classification-a tool for resistance management. Pestic Biochem Physiol. 2020;167:104587. doi: 10.1016/j.pestbp.2020.104587 EDN: ZTIAJJFurlong MJ, Shi ZH, Liu YQ, et al. Experimental analysis of the influence of pest management practice on the efficacy of an endemic arthropod natural enemy complex of the diamondback moth. J Econ Entomol. 2004;97(6):1814–1827. doi: 10.1093/jee/97.6.1814Badenes-Pérez FR. Trap crops and insectary plants in the order Brassicales. Ann Entomol Soc Am. 2018;112(4):318–329. doi: 10.1093/jee/100.1.215da Silva Ponce F, Trento DA, Toledo C, et al. Low tunnels with shading meshes: an alternative for the management of insect pests in kale cultivation. Scientia Horticulturae. 2021;288:110284. doi: 10.1016/j.scienta.2021.110284 EDN: CONNLLZhao JZ, Li YX, Collins H, et al. Monitoring and characterization of diamondback moth (Lepidoptera: Plutellidae) resistance to spinosad. J Econ Entomol. 2002;95(2):430–436. doi: 10.1603/0022-0493-95.2.430Ankersmit GW. DDT resistance in Plutella maculipennis (Curt.) (Lep.) in Java. Bulletin Entomological Research. 1953;44(3):421–425. doi: 10.1017/S0007485300025530Johnson DR. Plutella maculipennis resistance to DDT in Java. J Econ Entomol. 1953;46(1):176.Iqbal M, Verkerk RHJ, Furlong MJ, et al. Evidence for resistance to Bacillus thuringiensis (Bt) subsp. kurstaki HD-1, Bt subsp. aizawai and abamectin in field populations of Plutella xylostella from Malaysia. Pesticide Science. 1996;48(1): 89–97. doi: 10.1002/(SICI)1096-9063(199609)48:1<89:: AID-PS450>3.0.CO;2-BAttique MNR, Khaliq A, Sayyed AH. Could resistance to insecticides in Plutella xylostella (Lep., Plutellidae) be overcome by insecticide mixtures? J Appl Entomol. 2006;130(2):122–127. doi: 10.1111/j.1439-0418.2006.01035.xRibeiro LMS, Siqueira HAA, Wanderley-Teixeira V, et al. Field resistance of Brazilian Plutella xylostella to diamides is not metabolism-mediated. Crop Prot. 2017;93:82–88. doi: 10.1016/j.cropro.2016.11.027Palmquist K, Salatas J, Fairbrother A. Pyrethroid insecticides: use, environmental fate, and ecotoxicology. In: Insecticides-advances in integrated pest management. 2012. P. 251–278.Alesho NA, Kostina MN, Kaira AN. Modern methods and means of destroying harmful insects and mites—vectors of human disease pathogens. Moscow: RMAPO; 2015. 68 p. ISBN: 978-5-7249-2502-0 (In Russ.)Williamson MS, Martinez-Torres D, Hick CA, et al. Identification of mutations in the housefly para-type sodium channel gene associated with knockdown resistance (KDR) to pyrethroid insecticides. Mol Gen Genet. 1996;252(1):51–60. doi: 10.1007/BF02173204 EDN: UVZCELDong K, Du Y, Rinkevich F, et al. Molecular biology of insect sodium channels and pyrethroid resistance. Insect Biochem Mol Biol. 2014;50(1):1–17. doi: 10.1016/j.ibmb.2014.03.012 EDN: SKRSTHBrackenbury WJ, Isom LL. Na+ channel β subunits: overachievers of the ion channel family. Front Pharmacol. 2011;2:53. doi: 10.3389/fphar.2011.00053Catterall WA. From ionic currents to molecular mechanisms: the structure and function of voltage-gated sodium channels. Neuron. 2000;26(1):13–25. doi: 10.1016/S0896-6273(00)81133-2Dong K. Insect sodium channels and insecticide resistance. Invert Neurosci. 2007;7(1):17–30. doi: 10.1007/s10158-006-0036-9 EDN: TLJOLWLoughney K, Kreber R, Ganetzky B. Molecular analysis of the para locus, a sodium channel gene in Drosophila. Cell. 1989;58(6):1143–1154. doi: 10.1016/0092-8674(89)90512-6Soderlund DM, Bloomquist JR. Neurotoxic actions of pyrethroid insecticides. Annu Rev Entomol. 1989;34:77–96. doi: 10.1146/annurev.en.34.010189.000453Feng G, Deak P, Chopra M, et al. Cloning and functional analysis of TipE, a novel membrane protein that enhances Drosophila para sodium channel function. Cell. 1995;82(6):1001–1011.Warmke JW, Reenan RAG, Wang P, et al. Functional expression of Drosophila para sodium channels: modulation by the membrane protein TipE and toxin pharmacology. J Gen Physiol. 1997;110(2):119–133. doi: 10.1085/jgp.110.2.119Bourdin CM, Moignot B, Wang L, et al. Intron retention in mRNA encoding ancillary subunit of insect voltage-gated sodium channel modulates channel expression, gating regulation, and drug sensitivity. PLoS One. 2013;8(8):e67290. doi: 10.1371/journal.pone.0067290Du Y, Nomura Y, Satar G, et al. Molecular evidence for dual pyrethroid-receptor sites on a mosquito sodium channel. Proc Natl Acad Sci USA. 2013;110(29):11785–11790. doi: 10.1073/pnas.1305118110 EDN: RFIMZRLee SH, Smith T, Knipple DC, et al. Mutations in the house fly Vssc1 sodium channel gene associated with super-kdr resistance abolish the pyrethroid sensitivity of Vssc1/tipE sodium channels expressed in Xenopus oocytes. Insect Biochem Mol Biol. 1999;29(2):185–194. doi: 10.1016/S0965-1748(98)00122-2Derst C, Walther C, Veh RW, et al. Four novel sequences in Drosophila melanogaster homologous to the auxiliary Para sodium channel subunit TipE. Biochem Biophys Res Commun. 2006;339(3):939–948. doi: 10.1016/j.bbrc.2005.11.096Narahashi T. Neuronal ion channels as the target sites of insecticides. Pharmacol Toxicol. 1996;79(1):1–14. doi: 10.1111/j.1600-0773.1996.tb00234.x EDN: YAOKDVSilver KS, Du Y, Nomura Y, et al. Voltage-gated sodium channels as insecticide targets. Adv Insect Physiol. 2014;46:389–433. doi: 10.1016/B978-0-12-417010-0.00005-7 EDN: SKQIVVSoderlund DM. Molecular mechanisms of pyrethroid insecticide neurotoxicity: recent advances. Arch Toxicol. 2012;86(2):165–181. doi: 10.1007/s00204-011-0726-x EDN: FKRDKBShen XJ, Cao LJ, Chen JC, et al. A comprehensive assessment of insecticide resistance mutations in source and immigrant populations of the diamondback moth Plutella xylostella (L.). Pest Manag Sci. 2023;79(2):569–583. doi: 10.1002/ps.7223 EDN: YCOCHGLiu Z, Ma H, Li K, et al. Frequencies of insecticide resistance mutations detected by the amplicon sequencing in Plutella xylostella (Lepidoptera: Plutellidae) and Spodoptera exigua (Lepidoptera: Noctuidae) from China. J Econ Entomol. 2024;117(4):1648–1654. doi: 10.1093/jee/toae109 EDN: FXOHKCKwon DH, Choi BR, Park HM, et al. Knockdown resistance allele frequency in field populations of Plutella xylostella in Korea. Pestic Biochem Physiol. 2004;80(1):21–30. doi: 10.1016/j.pestbp.2004.06.001Endersby NM, Viduka K, Baxter SW, et al. Widespread pyrethroid resistance in Australian diamondback moth, Plutella xylostella (L.), is related to multiple mutations in the para sodium channel gene. Bull Entomol Res. 2011;101(4):393–405. doi: 10.1017/S0007485310000684Sonoda S. Molecular analysis of pyrethroid resistance conferred by target insensitivity and increased metabolic detoxification in Plutella xylostella. Pest Manag Sci. 2010;66(5):572–575. doi: 10.1002/ps.1918Busvine JR. Mechanism of resistance to insecticide in houseflies. Nature. 1951;168(4266):193–195. doi: 10.1038/168193a0Soderlund DM, Knipple DC. The molecular biology of knockdown resistance to pyrethroid insecticides. Insect Biochem Mol Biol. 2003;33(6): 563–577. doi: 10.1016/S0965-1748(03)00023-7 EDN: BDHQVSSchuler TH, Martinez-Torres D, Thompson AJ, et al. Toxicological, electrophysiological, and molecular characterisation of knockdown resistance to pyrethroid insecticides in the diamondback moth, Plutella xylostella (L.). Pestic Biochem Physiol. 1998;59(3):169–182. doi: 10.1006/pest.1998.2320Vais H, Williamson MS, Devonshire AL, et al. The molecular interactions of pyrethroid insecticides with insect and mammalian sodium channels. Pest Manag Sci. 2001;57(10):877–888. doi: 10.1002/ps.392Soderlund DM. Pyrethroids, knockdown resistance and sodium channels. Pest Manag Sci. 2008,64(6):610–616. doi: 10.1002/ps.1574.Usherwood PNR, Davies TGE, Mellor IR, et al. Mutations in DIIS5 and the DIIS4-S5 linker of Drosophila melanogaster sodium channel define binding domains for pyrethroids and DDT. FEBS Lett. 2007;581(28):5485–5492. doi: 10.1016/j.febslet.2007.10.057Chang CC, Dai SM, Chen CY, et al. Insecticide resistance and characteristics of mutations related to target site insensitivity of diamondback moths in Taiwan. Pestic Biochem Physiol. 2024;203:106001. doi: 10.1016/j.pestbp.2024.106001 EDN: VOUOUYChen M, Du Y, Nomura Y, et al. Chronology of sodium channel mutations associated with pyrethroid resistance in Aedes aegypti. Arch Insect Biochem Physiol. 2020;104(2):e21686. doi: 10.1002/arch.21686 EDN: WZDFIEKawada H, Higa Y, Kasai S. Reconsideration of importance of the point mutation L982W in the voltage-sensitive sodium channel of the pyrethroid resistant Aedes aegypti (L.) (Diptera: Culicidae) in Vietnam. PLoS One. 2023;18(5):e0285883. doi: 10.1371/journal.pone.0285883 EDN: GEGJNFNazemi A, Khajehali J, Van Leeuwen T. Incidence and characterization of resistance to pyrethroid and organophosphorus insecticides in Thrips tabaci (Thysanoptera: Thripidae) in onion fields in Isfahan, Iran. Pestic Biochem Physiol. 2016;129:28–35. doi: 10.1016/j.pestbp.2015.10.013Jouraku A, Kuwazaki S, Iida H, et al. T929I and K1774N mutation pair and M918L single mutation identified in the voltage-gated sodium channel gene of pyrethroid-resistant Thrips tabaci (Thysanoptera: Thripidae) in Japan. Pestic Biochem Physiol. 2019;158:77–87. doi: 10.1016/j.pestbp.2019.04.012Wu M, Gotoh H, Waters T, et al. Identification of an alternative knockdown resistance (kdr)-like mutation, M918L, and a novel mutation, V1010A, in the Thrips tabaci voltage-gated sodium channel gene. Pest Manag Sci. 2014;70(6):977–981. doi: 10.1002/ps.3638Toda S, Morishita M. Identification of three point mutations on the sodium channel gene in pyrethroid-resistant Thrips tabaci (Thysanoptera: Thripidae). J Econ Entomol. 2009;102(6):2296–2300. doi: 10.1603/029.102.0635Boné E, Acevedo GR, Sterkel M, et al. Characterization of the pyrethroid resistance mechanism in a Blattella germanica (Dictyoptera: Blattellidae) strain from Buenos Aires (Argentina). Bull Entomol Res. 2022;112(1):21–28. doi: 10.1017/S000748532100050X EDN: VRWRJLPridgeon JW, Appel AG, Moar W, et al. Variability of resistance mechanisms in pyrethroid resistant German cockroaches (Dictyoptera: Blattellidae). Pestic Biochem Physiol. 2002;73(3):149–156. doi: 10.1016/S0048-3575(02)00103-7 EDN: BCWQGHDang K, Doggett SL, Leong XY, et al. Multiple mechanisms conferring broad-spectrum insecticide resistance in the tropical bed bug (Hemiptera: Cimicidae). J Econ Entomol. 2021;114(6):2473–2484. doi: 10.1093/jee/toab205 EDN: AEGMENSoh LS, Veera Singham G. Cuticle thickening associated with fenitrothion and imidacloprid resistance and influence of voltage-gated sodium channel mutations on pyrethroid resistance in the tropical bed bug, Cimex hemipterus. Pest Manag Sci. 2021;77(11):5202–5212. doi: 10.1002/ps.6561 EDN: BBQUVIValmorbida I, Hohenstein JD, Coates BS, et al. Association of voltage-gated sodium channel mutations with field-evolved pyrethroid resistant phenotypes in soybean aphid and genetic markers for their detection. Sci Rep. 2022;12(1):12020. doi: 10.1038/s41598-022-16366-1 EDN: SZMFDMKoç N, İnak E, Nalbantoğlu S, et al. Biochemical and molecular mechanisms of acaricide resistance in Dermanyssus gallinae populations from Turkey. Pestic Biochem Physiol. 2022;180:104985. doi: 10.1016/j.pestbp.2021.104985 EDN: RQDHOHNyoni BN, Gorman K, Mzilahowa T, et al. Pyrethroid resistance in the tomato red spider mite, Tetranychus evansi, is associated with mutation of the para-type sodium channel. Pest Manag Sci. 2011;67(8):891–897. doi: 10.1002/ps.2145Marshall KL, Moran C, Chen Y, et al. Detection of KDR pyrethroid resistance in the cotton aphid, Aphis gossypii (Hemiptera: Aphididae), using a PCR-RFLP assay. J Pestic Sci. 2012;37(2):169–172. doi: 10.1584/jpestics.D11-017Eleftherianos I, Foster SP, Williamson MS, et al. Characterization of the M918T sodium channel gene mutation associated with strong resistance to pyrethroid insecticides in the peach-potato aphid, Myzus persicae (Sulzer). Bull Entomol Res. 2008;98(2):183–191. doi: 10.1017/S0007485307005524Sun H, Nomura Y, Du Y, et al. Characterization of two mutations at predicted pyrethroid receptor site 2 in the sodium channels of Aedes aegypti and Nilaparvata lugens. Insect Biochem Mol Biol. 2022;148:103814. doi: 10.1016/j.ibmb.2022.103814 EDN: BRYSCGGuerrero FD, Jamroz RC, Kammlah D, et al. Toxicological and molecular characterization of pyrethroid-resistant horn flies, Haematobia irritans: identification of kdr and super-kdr point mutations. Insect Biochem Mol Biol. 1997;27(8–9):745–755. doi: 10.1016/S0965-1748(97)00057-X EDN: AJGPLXHaddi K, Berger M, Bielza P, et al. Identification of mutations associated with pyrethroid resistance in the voltage-gated sodium channel of the tomato leaf miner (Tuta absoluta). Insect Biochem Mol Biol. 2012;42(7):506–513. doi: 10.1016/j.ibmb.2012.03.008Zhu J, Chen R, Liu J, et al. Presence of multiple genetic mutations related to insecticide resistance in Chinese field samples of two Phthorimaea pest species. Insects. 2024;15(3):194. doi: 10.3390/insects15030194 EDN: CZCNLQBenito-Murcia M, Martín-Hernández R, Meana A, et al. Study of pyrethroid resistance mutations in populations of Varroa destructor across Spain. Res Vet Sci. 2022;152:34–37. doi: 10.1016/j.rvsc.2022.07.021 EDN: ZPDTQFBenavent-Albarracín L, Alonso M, Catalán J, et al. Mutations in the voltage-gated sodium channel gene associated with deltamethrin resistance in commercially sourced Phytoseiulus persimilis. Insect Mol Biol. 2020;29(4):373–380. doi: 10.1111/imb.12642 EDN: ABJSRVRiebenbauer K, Purkhauser K, Walochnik J, et al. Detection of a knockdown mutation in the voltage-sensitive sodium channel associated with permethrin tolerance in Sarcoptes scabiei var. hominis mites. J Eur Acad Dermatol Venereol. 2023;37(11):2355–2361. doi: 10.1111/jdv.19288 EDN: SADWACCarletto J, Martin T, Vanlerberghe-Masutti F, et al. Insecticide resistance traits differ among and within host races in Aphis gossypii. Pest Manag Sci. 2010;66(3):301–307. doi: 10.1002/ps.1874Wang ZM, Li CX, Xing D, et al. Detection and widespread distribution of sodium channel alleles characteristic of insecticide resistance in Culex pipiens complex mosquitoes in China. Med Vet Entomol. 2012;26(2):228–232. doi: 10.1111/j.1365-2915.2011.00985.xKaratolos N, Gorman K, Williamson MS, et al. Mutations in the sodium channel associated with pyrethroid resistance in the greenhouse whitefly, Trialeurodes vaporariorum. Pest Manag Sci. 2012;68(6):834–838. doi: 10.1002/ps.2334Weston DP, Poynton HC, Wellborn GA, et al. Multiple origins of pyrethroid insecticide resistance across the species complex of a nontarget aquatic crustacean, Hyalella azteca. Proc Natl Acad Sci U S A. 2013;110(41): 16532–16537. doi: 10.1073/pnas.1302023110Morin S, Williamson MS, Goodson SJ, et al. Mutations in the Bemisia tabaci para sodium channel gene associated with resistance to a pyrethroid plus organophosphate mixture. Insect Biochem Mol Biol. 2002;32(12): 1781–1791. doi: 10.1016/S0965-1748(02)00137-6 EDN: BBPWQJBao WX, Sonoda S. Resistance to cypermethrin in melon thrips, Thrips palmi, (Thysanoptera: Thripidae), is conferred by reduced sensitivity of the sodium channel and CYP450-mediated detoxification. Appl Entomol Zool. 2012;47:443–448. doi: 10.1007/s13355-012-0141-7Forcioli D, Frey B, Frey JE. High nucleotide diversity in the para-like voltage-sensitive sodium channel gene sequence in the western flower thrips (Thysanoptera: Thripidae). J Econ Entomol. 2002;95(4):838–848. doi: 10.1603/0022-0493-95.4.838Gao R, Ma S, Geng J, et al. Functional characterization of double mutations T929I/K1774N in the voltage-gated sodium channel of Megalurothrips usitatus (Bagnall) related to pyrethroid resistance. J Agric Food Chem. 2024;72(21):11958–11967. doi: 10.1021/acs.jafc.4c00355 EDN: QRKRUFRinkevich FD, Su C, Lazo TA, et al. Multiple evolutionary origins of knockdown resistance (KDR) in pyrethroid-resistant Colorado potato beetle, Leptinotarsa decemlineata. Pestic Biochem Physiol. 2012;104(3):192–200. doi: 10.1016/j.pestbp.2012.08.001 EDN: YDHZCNAraujo RA, Williamson MS, Bass C, et al. Pyrethroid resistance in Sitophilus zeamais is associated with a mutation (T929I) in the voltage-gated sodium channel. Insect Mol Biol. 2011;20(4):437–445. doi: 10.1111/j.1365-2583.2011.01079.xSingh OP, Dykes CL, Das MK, et al. Presence of two alternative kdr-like mutations, L1014F and L1014S, and a novel mutation, V1010L, in the voltage-gated Na+ channel of Anopheles culicifacies from Orissa, India. Malar J. 2010;9:146. doi: 10.1186/1475-2875-9-146 EDN: MTBYEEFukazawa N, Takahashi R, Matsuda H, et al. Sodium channel mutations (T929I and F1534S) found in pyrethroid-resistant strains of the cigarette beetle, Lasioderma serricorne (Coleoptera: Anobiidae). J Pestic Sci. 2021;46(4):360–365. doi: 10.1584/jpestics.D21-033 EDN: WDJIEEGuan F, Zhang J, Shen H, et al. Whole-genome sequencing to detect mutations associated with resistance to insecticides and Bt proteins in Spodoptera frugiperda. Insect Sci. 2021;28(3):627–638. doi: 10.1111/1744-7917.12838 EDN: BIXGQIRoditakis E, Tsagkarakou A, Vontas J. Identification of mutations in the para sodium channel of Bemisia tabaci from Crete, associated with resistance to pyrethroids. Pestic Biochem Physiol. 2006;85(3):161–166. doi: 10.1016/j.pestbp.2005.11.007Bass C, Schroeder I, Turberg A, et al. Identification of mutations associated with pyrethroid resistance in the para-type sodium channel of the cat flea, Ctenocephalides felis. Insect Biochem Mol Biol. 2004;34(12):1305–1313. doi: 10.1016/j.ibmb.2004.09.002Seidy S, Tavassoli M, Malekifard F. Pyrethroids resistance in Pulex irritans and Ctenocephalides canis in west and northwest Iran. Vet Res Forum. 2022;13(4):529–535. doi: 10.30466/vrf.2021.534642.3215Scott JG. Evolution of resistance to pyrethroid insecticides in Musca domestica. Pest Manag Sci. 2017;73(4):716–722. doi: 10.1002/ps.4328 EDN: WKCFNVFoster SP, Paul VL, Slater R, et al. A mutation (L1014F) in the voltage-gated sodium channel of the grain aphid, Sitobion avenae, is associated with resistance to pyrethroid insecticides. Pest Manag Sci. 2014;70(8):1249–1253. doi: 10.1002/ps.3683Okafor MA, Ekpo ND, Opara KN, et al. Pyrethroid insecticides susceptibility profiles and evaluation of L1014F kdr mutant alleles in Culex quinquefasciatus from lymphatic filariasis endemic communities. Sci Rep. 2023;13(1):18716. doi: 10.1038/s41598-023-44962-2 EDN: EBOPPQWang L, Soto A, Remue L, et al. First report of mutations associated with pyrethroid (L1014F) and organophosphate (G119S) resistance in Belgian Culex (Diptera: Culicidae) mosquitoes. J Med Entomol. 2022;59(6):2072–2079. doi: 10.1093/jme/tjac138 EDN: ZPZQGJSamake JN, Yared S, Getachew D, et al. Detection and population genetic analysis of kdr L1014F variant in eastern Ethiopian Anopheles stephensi. Infect Genet Evol. 2022;99:105235. doi: 10.1016/j.meegid.2022.105235 EDN: LZIAGBCarter TE, Gebresilassie A, Hansel S, et al. Analysis of the knockdown resistance locus (kdr) in Anopheles stephensi, An. arabiensis, and Culex pipiens s. l. for insight into the evolution of target-site pyrethroid resistance in eastern Ethiopia. Am J Trop Med Hyg. 2022;106(2):632–638. doi: 10.4269/ajtmh.20-1357 EDN: UKICOSBadolo A, Okado K, Guelbeogo WM, et al. Development of an allele-specific, loop-mediated, isothermal amplification method (AS-LAMP) to detect the L1014F kdr-w mutation in Anopheles gambiae s. l. Malar J. 2012;11:227. doi: 10.1186/1475-2875-11-227 EDN: UWBLZIBacca T, Haddi K, Pineda M, et al. Pyrethroid resistance is associated with a kdr-type mutation (L1014F) in the potato tuber moth Tecia solanivora. Pest Manag Sci. 2017;73(2):397–403. doi: 10.1002/ps.4414 EDN: YVTZNHSoleño J, Parra-Morales LB, Cichón L, et al. Occurrence of pyrethroid resistance mutation in Cydia pomonella (Lepidoptera: Tortricidae) throughout Argentina. Bull Entomol Res. 2020;110(2):201–206. doi: 10.1017/S0007485319000439 EDN: ZAEYTMZimmer CT, Müller A, Heimbach U, et al. Target-site resistance to pyrethroid insecticides in German populations of the cabbage stem flea beetle, Psylliodes chrysocephala L. (Coleoptera: Chrysomelidae). Pestic Biochem Physiol. 2014;108:1–7. doi: 10.1016/j.pestbp.2013.11.005Olafson PU, Pitzer JB, Kaufman PE. Identification of a mutation associated with permethrin resistance in the para-type sodium channel of the stable fly (Diptera: Muscidae). J Econ Entomol. 2011;104(1):250–257. doi: 10.1603/EC10307Kim H, Baek JH, Lee WJ, et al. Frequency detection of pyrethroid resistance allele in Anopheles sinensis populations by real-time PCR amplification of specific allele (rtPASA). Pestic Biochem Physiol. 2007;87(1):54–61. doi: 10.1016/j.pestbp.2006.06.009Lol JC, Castellanos ME, Liebman KA, et al. Molecular evidence for historical presence of knock-down resistance in Anopheles albimanus, a key malaria vector in Latin America. Parasit Vectors. 2013;6:268. doi: 10.1186/1756-3305-6-268 EDN: SQWYAVLiu N, Pridgeon JW. Metabolic detoxication and the kdr mutation in pyrethroid resistant house flies, Musca domestica (L.). Pestic Biochem Physiol. 2002;73(3):157–163. doi: 10.1016/S0048-3575(02)00101-3 EDN: BCWQFNRinkevich FD, Zhang L, Hamm RL, et al. Frequencies of the pyrethroid resistance alleles of Vssc1 and CYP6D1 in house flies from the eastern United States. Insect Mol Biol. 2006;15(2):157–167. doi: 10.1111/j.1365-2583.2006.00620.xDavies TG, Field LM, Usherwood PN, et al. A comparative study of voltage-gated sodium channels in the Insecta: implications for pyrethroid resistance in Anopheline and other Neopteran species. Insect Mol Biol. 2007;16(3):361–375. doi: 10.1111/j.1365-2583.2007.00733.x EDN: MEMKBFField LM, James AA, Park Y, et al. Voltage-gated sodium channel genes hscp and hDSC1 of Heliothis virescens F. genomic organization. Insect Mol Biol. 1999;8(2):161–170. doi: 10.1046/j.1365-2583.1999.820161.x EDN: ETVUQHHopkins BW, Pietrantonio PV. The Helicoverpa zea (Boddie) (Lepidoptera: Noctuidae) voltage-gated sodium channel and mutations associated with pyrethroid resistance in field-collected adult males. Insect Biochem Mol Biol. 2010;40(5):385–393. doi: 10.1016/j.ibmb.2010.03.004 EDN: NZSTATStump AD, Atieli FK, Vulule JM, et al. Dynamics of the pyrethroid knockdown resistance allele in western Kenyan populations of Anopheles gambiae in response to insecticide-treated bed net trials. Am J Trop Med Hyg. 2004;70(6):591–596.Ranson H, Jensen B, Vulule MJ, et al. Identification of a point mutation in the voltage-gated sodium channel gene of Kenyan Anopheles gambiae associated with resistance to DDT and pyrethroids. Insect Mol Biol. 2000;9(5):491–497. doi: 10.1046/j.1365-2583.2000.00209.xVerhaeghen K, Bortel WV, Trung HD, et al. Knockdown resistance in Anopheles vagus, An. sinensis, An. paraliae and An. peditaeniatus populations of the Mekong region. Parasit Vectors. 2010;3(1):59. doi: 10.1186/1756-3305-3-59 EDN: AZIUKSLüleyap HÜ, Alptekin D, Kasap H, et al. Detection of knockdown resistance mutations in Anopheles sacharovi (Diptera: Culicidae) and genetic distance with Anopheles gambiae (Diptera: Culicidae) using cDNA sequencing of the voltage-gated sodium channel gene. J Med Entomol. 2002;39(6):870–874. doi: 10.1603/0022-2585-39.6.870Chen L, Zhong D, Zhang D, et al. Molecular ecology of pyrethroid knockdown resistance in Culex pipiens pallens mosquitoes. PLoS One. 2010;5(7):e11681. doi: 10.1371/journal.pone.0011681 EDN: NZFUQFMartinez-Torres D, Foster SP, Field LM, et al. A sodium channel point mutation is associated with resistance to DDT and pyrethroid insecticides in the peach-potato aphid, Myzus persicae (Sulzer) (Hemiptera: Aphididae). Insect Mol Biol. 1999;8(3):339–346. doi: 10.1046/j.1365-2583.1999.83121.xTan WL, Li CX, Wang ZM, et al. First detection of multiple knockdown resistance (kdr)-like mutations in voltage-gated sodium channel using three new genotyping methods in Anopheles sinensis from Guangxi Province, China. J Med Entomol. 2014;49(5):1012–1020. doi: 10.1603/ME11266O’Reilly AO, Williamson MS, González-Cabrera J, et al. Predictive 3D modelling of the interactions of pyrethroids with the voltage-gated sodium channels of ticks and mites. Pest Manag Sci. 2014;70(3):369–377. doi: 10.1002/ps.3561Feyereisen R. Insect P450 enzymes. Annu Rev Entomol. 1999;44: 507–533. doi: 10.1146/annurev.ento.44.1.507Liu N, Li M, Gong Y, et al. Cytochrome P450s-their expression, regulation, and role in insecticide resistance. Pestic Biochem Physiol. 2015;120:77–81. doi: 10.1016/j.pestbp.2015.01.006Wang S, Liu X, Tang H, et al. UGT2B13 and UGT2C1 are involved in lambda-cyhalothrin resistance in Rhopalosiphum padi. Pestic Biochem Physiol. 2023;194:105528. doi: 10.1016/j.pestbp.2023.105528 EDN: SMIHGTNdo C, Kopya E, Irving H, et al. Exploring the impact of glutathione S-transferase (GST)-based metabolic resistance to insecticide on vector competence of Anopheles funestus for Plasmodium falciparum. Wellcome Open Res. 2019;4:52. doi: 10.12688/wellcomeopenres.15061.2Du Y, Zhu YC, Portilla M, et al. The mechanisms of metabolic resistance to pyrethroids and neonicotinoids fade away without selection pressure in the tarnished plant bug Lygus lineolaris. Pest Manag Sci. 2023;79(10):3893–3902. doi: 10.1002/ps.7570 EDN: TSCMNDZhang S, Zhang X, Shen J, et al. Susceptibility of field populations of the diamondback moth, Plutella xylostella, to a selection of insecticides in Central China. Pestic Biochem Physiol. 2016;132:38–46. doi: 10.1016/j.pestbp.2016.01.007Wang X, Wang J, Cao X, et al. Long-term monitoring and characterization of resistance to chlorfenapyr in Plutella xylostella (Lepidoptera: Plutellidae) from China. Pest Manag Sci. 2019;75(3):591–597. doi: 10.1002/ps.5222Li X, Liu Z, Lv X, et al. Molecular mechanism of λ-cyhalothrin detoxification by a delta-class glutathione S-transferase (PxGSTD3) from Plutella xylostella. J Agric Food Chem. 2025;73(11):6559–6566. doi: 10.1021/acs.jafc.4c12498 EDN: WELRUGBautista MAM, Bhandary B, Wijeratne AJ, et al. Evidence for trade-offs in detoxification and chemosensation gene signatures in Plutella xylostella. Pest Manag Sci. 2015;71(3):423–432. doi: 10.1002/ps.3822Li R, Zhu B, Shan J, et al. Functional analysis of a carboxylesterase gene involved in beta-cypermethrin and phoxim resistance in Plutella xylostella (L.). Pest Manag Sci. 2021;77(4):2097–2105. doi: 10.1002/ps.6238 EDN: QAZXUCChrustek A, Hołyńska-Iwan I, Dziembowska I, et al. Current research on the safety of pyrethroids used as insecticides. Medicina. 2018;54(4):61. doi: 10.3390/medicina54040061Bradberry SM, Cage SA, Proudfoot AT, et al. Poisoning due to pyrethroids. Toxicol Rev. 2005;24(2):93–106. doi: 10.2165/00139709-200524020-00003Wang M, Zhu B, Zhang L, et al. Influence of seasonal migration on evolution of insecticide resistance in Plutella xylostella. Insect Sci. 2022;29(2):496–504. doi: 10.1111/1744-7917.12987 EDN: LXMNIJ