An C, Lovell S, Kanost M R, Battaile K P, Michel K. 2011. Crystal structure of native Anopheles gambiae serpin-2, a negative regulator of melanization in mosquitoes. Proteins–Structure Function and Bioinformatics, 79, 1999–2003.

Armenteros J J A, Tsirigos K D, Nderby C K S, Petersen T N, Winther O, Brunak S R, von Heijne G, Nielsen H. 2019. SignalP 5.0 improves signal peptide predictions using deep neural networks. Nature Biotechnology, 37, 420–423.

Bao J Q, Chen Y X, Jiang S W, Liu R, Zhang X, Zhang F Z, Chen Z W, Luo C, Kong H L. 2024. Induced CTL-S15 gene expression by Bacillus thuringiensis declines susceptibility in Spodoptera exigua. Journal of Integrative Agriculture, 23, 3078–3088.

Blum M, Chang H, Chuguransky S, Grego T, Kandasaamy S, Mitchell A, Nuka G, Paysan-Lafosse T, Qureshi M, Raj S, Richardson L, Salazar G A, Williams L, Bork P, Bridge A, Gough J, Haft D H, Letunic I, Marchler-Bauer A, Mi H, et al. 2020. The InterPro protein families and domains database: 20 years on. Nucleic Acids Research, 49, D344-D354.

Briscoe J, Thérond P P. 2013. The mechanisms of hedgehog signalling and its roles in development and disease. Nature Reviews Molecular Cell Biology, 14, 416–429.

Calvo E, Mizurini D M, Sá-Nunes A, Ribeiro J M C, Andersen J F, Mans B J, Monteiro R Q, Kotsyfakis M, Francischetti I M B. 2011. Alboserpin, a factor Xa inhibitor from the mosquito vector of yellow fever, binds heparin and membrane phospholipids and exhibits antithrombotic activity. Journal of Biological Chemistry, 286, 27998–28010.

Chen T, Hu N, Tan L, Xiao Q, Dong Z, Chen P, Xu A, Pan M, Lu C. 2019. Resistant silkworm strain block viral infection independent of melanization. Pesticide Biochemistry and Physiology, 154, 88–96.

Christen J M, Hiromasa Y, An C, Kanost M R. 2012. Identification of plasma proteinase complexes with serpin-3 in Manduca sexta. Insect Biochemistry and Molecular Biology, 42, 946–955.

Colinet D, Deleury E, Anselme C, Cazes D, Poulain J, Azema-Dossat C, Belghazi M, Gatti J, Poirié M. 2013. Extensive inter- and intraspecific venom variation in closely related parasites targeting the same host: The case of leptopilina parasitoids of Drosophila. Insect Biochemistry and Molecular Biology, 43, 601–611.

Colinet D, Dubuffet A, Cazes D, Moreau S, Drezen J, Poirié M. 2009. A serpin from the parasitoid wasp Leptopilina boulardi targets the Drosophila phenoloxidase cascade. Developmental and Comparative Immunology, 33, 681–689.

Concordet J, Haeussler M. 2018. CRISPOR: Intuitive guide selection for CRISPR/cas9 genome editing experiments and screens. Nucleic Acids Research, 46, W242–W245.

Danielli A, Barillas-Mury C, Kumar S, Kafatos F C, Loukeris T G. 2005. Overexpression and altered nucleocytoplasmic distribution of anopheles ovalbumin-like SRPN10 serpins in plasmodium-infected midgut cells. Cellular Microbiology, 7, 181–190.

Etebari K, Asgari S. 2013. Conserved microRNA mir-8 blocks activation of the toll pathway by upregulating serpin 27 transcripts. RNA Biology, 10, 1356–1364.

Furlong M J, Wright D J, Dosdall L M. 2013. Diamondback moth ecology and management: Problems, progress, and prospects. Annual Review of Entomology, 58, 517–541.

Gan H, Wang Y, Jiang H, Mita K, Kanost M R. 2001. A bacteria-induced, intracellular serpin in granular hemocytes of Manduca sexta. Insect Biochemistry and Molecular Biology, 31, 887–898.

Garrett M, Fullaondo A, Troxler L, Micklem G, Gubb D. 2009. Identification and analysis of serpin-family genes by homology and synteny across the 12 sequenced drosophilid genomes. BMC Genomics, 10, 489.

Gasteiger E. 2003. ExPASy: The proteomics server for in-depth protein knowledge and analysis. Nucleic Acids Research, 31, 3784–3788.

De Gregorio E, Han S, Lee W, Baek M, Osaki T, Kawabata S, Lee B, Iwanaga S, Lemaitre B, Brey P T. 2002. An immune-responsive serpin regulates the melanization cascade in Drosophila. Developmental Cell, 3, 581–592.

Guo P, Dong Z, Zhao P, Zhang Y, He H, Tan X, Zhang W, Xia Q. 2015. Structural insights into the unique inhibitory mechanism of the silkworm protease inhibitor serpin18. Scientific Reports, 5, 11863.

Han P, Fan J, Liu Y, Cuthbertson A G S, Yan S, Qiu B, Ren S. 2014. RNAi-mediated knockdown of serine protease inhibitor genes increases the mortality of Plutella xylostella challenged by destruxin A. PLoS ONE, 9, e97863.

Hashimoto C, Hudson K L, Anderson K V. 1988. The toll gene of Drosophila, required for dorsal-ventral embryonic polarity, appears to encode a transmembrane protein. Cell, 52, 269–279.

Hashimoto C, Kim D R, Weiss L A, Miller J W, Morisato D. 2003. Spatial regulation of developmental signaling by a serpin. Developmental Cell, 5, 945–950.

Homma S, Murata A, Ikegami M, Kobayashi M, Yamazaki M, Ikeda K, Daimon T, Numata H, Mizoguchi A, Shiomi K. 2022. Circadian clock genes regulate temperature-dependent diapause induction in silkworm Bombyx mori. Frontiers in Physiology, 13, 863380.

Jang I, Chosa N, Kim S, Nam H, Lemaitre B, Ochiai M, Kambris Z, Brun S, Hashimoto C, Ashida M, Brey P T, Lee W. 2006. A spätzle-processing enzyme required for toll signaling activation in Drosophila innate immunity. Developmental Cell, 10, 45–55.

Jiang H, Wang Y, Yu X, Zhu Y, Kanost M. 2003. Prophenoloxidase-activating proteinase-3 (PAP-3) from Manduca sexta hemolymph: A clip-domain serine proteinase regulated by serpin-1J and serine proteinase homologs. Insect Biochemistry and Molecular Biology, 33, 1049–1060.

Karanja F, Sahu S, Weintraub S, Bhandari R, Jaszczak R, Sitt J, Halme A. 2022. Ecdysone exerts biphasic control of regenerative signaling, coordinating the completion of regeneration with developmental progression. Proceedings of the National Academy of Sciences of the United States of America, 119, e2115017119.

Lemaitre B, Hoffmann J. 2007. The host defense of Drosophila melanogaster. Annual Review of Immunology, 25, 697–743.

Letunic I, Bork P. 2006. Interactive tree of life (iTOL): An online tool for phylogenetic tree display and annotation. Bioinformatics, 23, 127–128.

Letunic I, Khedkar S, Bork P. 2020. SMART: Recent updates, new developments and status in 2020. Nucleic Acids Research, 49, D458–D460.

Li S, Xu X, De Mandal S, Shakeel M, Hua Y, Shoukat R F, Fu D, Jin F. 2021. Gut microbiota mediate Plutella xylostella susceptibility to Bt cry1ac protoxin is associated with host immune response. Environmental Pollution, 271, 116271.

Li Z, Feng X, Liu S, You M, Furlong M J. 2016. Biology, ecology, and management of the diamondback moth in China. Annual Review of Entomology, 61, 277–296.

Ligoxygakis P. 2002. A serpin mutant links toll activation to melanization in the host defence of Drosophila. The EMBO Journal, 21, 6330–6337.

Ligoxygakis P, Roth S, Reichhart J. 2003. A serpin regulates dorsal-ventral axis formation in the Drosophila embryo. Current Biology, 13, 2097–2102.

Lin H, Lin X, Zhu J, Yu X, Xia X, Yao F, Yang G, You M. 2017. Characterization and expression profiling of serine protease inhibitors in the diamondback moth, Plutella xylostella (Lepidoptera: Plutellidae). BMC Genomics, 18, 162.

Lin J, Yu X, Wang Q, Tao X, Li J, Zhang S, Xia X, You M. 2020. Immune responses to Bacillus thuringiensis in the midgut of the diamondback moth, Plutella xylostella. Developmental and Comparative Immunology, 107, 103661.

Lu C, Lee R K, Hwu Y, Chu S, Chen Y, Chang W, Lin S, Li S. 2011. SERPINE2, a serine protease inhibitor extensively expressed in adult male mouse reproductive tissues, may serve as a murine sperm decapacitation factor1. Biology of Reproduction, 84, 514–525.

Mantica F, Iñiguez L P, Marquez Y, Permanyer J, Torres-Mendez A, Cruz J, Franch-Marro X, Tulenko F, Burguera D, Bertrand S, Doyle T, Nouzova M, Currie P D, Noriega F G, Escriva H, Arnone M I, Albertin C B, Wotton K R, Almudi I, Martin D, et al. 2024. Evolution of tissue-specific expression of ancestral genes across vertebrates and insects. Nature Ecology & Evolution, 8, 1140–1153.

Meekins D A, Kanost M R, Michel K. 2017. Serpins in arthropod biology. Seminars in Cell & Developmental Biology, 62, 105–119.

Moller S, Croning M D, Apweiler R. 2001. Evaluation of methods for the prediction of membrane spanning regions. Bioinformatics, 17, 646–653.

Mongelli V, Lequime S, Kousathanas A, Gausson V, Blanc H, Nigg J, Quintana-Murci L, Elena S F, Saleh M. 2022. Innate immune pathways act synergistically to constrain RNA virus evolution in Drosophila melanogaster. Nature Ecology & Evolution, 6, 565–578.

Mueller J L, Page J L, Wolfner M F. 2007. An ectopic expression screen reveals the protective and toxic effects of Drosophila seminal fluid proteins. Genetics, 175, 777–783.

Nappi A J, Frey F, Carton Y. 2005. Drosophila serpin 27a is a likely target for immune suppression of the blood cell-mediated melanotic encapsulation response. Journal of Insect Physiology, 51, 197–205.

Owji H, Nezafat N, Negahdaripour M, Hajiebrahimi A, Ghasemi Y. 2018. A comprehensive review of signal peptides: Structure, roles, and applications. European Journal of Cell Biology, 97, 422–441.

Peng L, Zheng J, Liu L, Huang M, Cao M, Cui J, Vasseur L, You M, Zou M. 2025. Identification of seminal fluid proteins and reproductive function of trypsin-1 in male Plutella xylostella. International Journal of Biological Macromolecules, 306, 141450.

Qie X, Yan X, Wang W, Liu Y, Zhang L, Hao C, Lu Z, Ma L. 2023. Serpin-4 negatively regulates prophenoloxidase activation and antimicrobial peptide synthesis in the silkworm, Bombyx mori. International Journal of Molecular Sciences, 25, 313.

Ram K R, Wolfner M F. 2007. Sustained post-mating response in Drosophila melanogaster requires multiple seminal fluid proteins. PLoS Genetics, 3, e238.

Shakeel M, Xu X, De Mandal S, Jin F. 2019. Role of serine protease inhibitors in insect-host-pathogen interactions. Archives of Insect Biochemistry and Physiology, 102, e21556.

Shi G, Zhou Y, Ren F. 2022. Identification and function analysis of BmPxtA in the immune response regulated by PGE2 of silkworm, Bombyx mori. Developmental and Comparative Immunology, 130, 104358.

Shin S W, Park S, Park D, Kim M G, Kim S C, Brey P T, Park H. 1998. Isolation and characterization of immune-related genes from the fall webworm, Hyphantria cunea, using PCR-based differential display and subtractive cloning. Insect Biochemistry and Molecular Biology, 28, 827–837.

Stark K R, James A A. 1998. Isolation and characterization of the gene encoding a novel factor Xa-directed anticoagulant from the yellow fever mosquito, Aedes aegypti. Journal of Biological Chemistry, 273, 20802–20809.

Suwanchaichinda C, Ochieng R, Zhuang S, Kanost M R. 2013. Manduca sexta serpin-7, a putative regulator of hemolymph prophenoloxidase activation. Insect Biochemistry and Molecular Biology, 43, 555–561.

Tamura K, Stecher G, Kumar S. 2021. MEGA11: Molecular evolutionary genetics analysis version 11. Molecular Biology and Evolution, 38, 3022–3027.

Tong Y, Jiang H, Kanost M R. 2005. Identification of plasma proteases inhibited by Manduca sexta serpin-4 and serpin-5 and their association with components of the prophenol oxidase activation pathway. Journal of Biological Chemistry, 280, 14932–14942.

Wang L, Liu H, Fu H, Zhang L, Guo P, Xia Q, Zhao P. 2019. Silkworm serpin32 functions as a negative-regulator in prophenoloxidase activation. Developmental and Comparative Immunology, 91, 123–131.

Watanabe A, Miyazawa S, Kitami M, Tabunoki H, Ueda K, Sato R. 2006. Characterization of a novel c-type lectin, Bombyx mori multibinding protein, from the B. mori hemolymph: Mechanism of wide-range microorganism recognition and role in immunity. Journal of Immunology, 177, 4594–4604.

Waterhouse A, Bertoni M, Bienert S, Studer G, Tauriello G, Gumienny R, Heer F T, de Beer T A P, Rempfer C, Bordoli L, Lepore R, Schwede T. 2018. SWISS-MODEL: Homology modelling of protein structures and complexes. Nucleic Acids Research, 46, W296–W303.

Wei Y, Huang M, Jiang L. 2024. Advancements in serine protease inhibitors: From mechanistic insights to clinical applications. Catalysts, 14, 787.

Wheeler D E, Tuchinskaya I, Buck N A, Tabashnik B E. 2000. Hexameric storage proteins during metamorphosis and egg production in the diamondback moth, Plutella xylostella (Lepidoptera). Journal of Insect Physiology, 46, 951–958.

Wolfner M F, Harada H A, Bertram M J, Stelick T J, Kraus K W, Kalb J M, Lung Y O, Neubaum D M, Park M, Tram U. 1997. New genes for male accessory gland proteins in Drosophila melanogaster. Insect Biochemistry and Molecular Biology, 27, 825–834.

Xu J, Xu X, Shakeel M, Li S, Wang S, Zhou X, Yu J, Xu X, Yu X, Jin F. 2017. The entomopathogenic fungi isaria fumosorosea plays a vital role in suppressing the immune system of Plutella xylostella: RNA-seq and DGE analysis of immunity-related genes. Frontiers in Microbiology, 8, 1421.

Xu Z, Yan Y, Cao J, Zhou Y, Zhang H, Xu Q, Zhou J. 2020a. A family of serine protease inhibitors (serpins) and its expression profiles in the ovaries of Rhipicephalus haemaphysaloide. Infection, Genetics and Evolution, 84, 104346a.

Xu Z, Yan Y, Zhang H, Cao J, Zhou Y, Xu Q, Zhou J. 2020b. A serpin from the tick Rhipicephalus haemaphysaloides: Involvement in vitellogenesis. Veterinary Parasitology, 279,109064b.

Yuan C, Xing L, Wang M, Wang X, Yin M, Wang Q, Hu Z, Zou Z. 2017. Inhibition of melanization by serpin-5 and serpin-9 promotes baculovirus infection in cotton bollworm Helicoverpa armigera. PLoS Pathogens, 13, e1006645.

Zalucki M P, Shabbir A, Silva R, Adamson D, Liu S S, Furlong M J. 2012. 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? Journal of Economic Entomology, 105, 1115–1129.

Zhang Z, Zhu S, De Mandal S, Gao Y, Yu J, Zeng L, Huang J, Zafar J, Jin F, Xu X. 2022. Combined transcriptomic and proteomic analysis of developmental features in the immune system of Plutella xylostella during larva-to-adult metamorphosis. Genomics, 114, 110381.

Zhu Y, Wang Y, Gorman M J, Jiang H, Kanost M R. 2003. Manduca sexta serpin-3 regulates prophenoloxidase activation in response to infection by inhibiting prophenoloxidase-activating proteinases. Journal of Biological Chemistry, 278, 46556–46564.

Zou Z, Jiang H. 2005. Manduca sexta serpin-6 regulates immune serine proteinases PAP-3 and HP8. Journal of Biological Chemistry, 280, 14341–14348.