{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,1,22]],"date-time":"2026-01-22T16:52:14Z","timestamp":1769100734313,"version":"3.49.0"},"reference-count":64,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2019,9,17]],"date-time":"2019-09-17T00:00:00Z","timestamp":1568678400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2019,9,17]],"date-time":"2019-09-17T00:00:00Z","timestamp":1568678400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100004396","name":"Thailand Research Fund","doi-asserted-by":"publisher","award":["IRN61W0001"],"award-info":[{"award-number":["IRN61W0001"]}],"id":[{"id":"10.13039\/501100004396","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004396","name":"Thailand Research Fund","doi-asserted-by":"publisher","award":["RSA6180069"],"award-info":[{"award-number":["RSA6180069"]}],"id":[{"id":"10.13039\/501100004396","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004396","name":"Thailand Research Fund","doi-asserted-by":"publisher","award":["IRN61W0001"],"award-info":[{"award-number":["IRN61W0001"]}],"id":[{"id":"10.13039\/501100004396","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100002873","name":"Chulalongkorn University","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100002873","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title><jats:p>White spot syndrome virus (WSSV) is one of the most lethal viruses severely affecting shrimp industry. This disease can cause 100% mortality of farmed shrimp within a week. This work aims to characterize clathrin assembly proteins in <jats:italic>Penaeus monodon<\/jats:italic> and investigate their roles in WSSV entry. In general, clathrin assembly proteins form complexes with specific receptors and clathrins, leading to clathrin-mediated endocytosis. Adaptor protein 2 (AP-2), which is responsible for endocytosis at plasma membrane, consists of four subunits including \u03b1, \u03b22, \u03bc2 and \u03c32. Knockdown of clathrin coat AP17, or \u03c3 subunit of AP-2 dramatically reduced WSSV infectivity. Similar results were observed, when shrimp were pre-treated with chlorpromazine (CPZ), an inhibitor of clathrin-dependent endocytosis. The complete open reading frames of AP-2\u03b2 and \u03bc subunits of <jats:italic>P<\/jats:italic>. <jats:italic>monodon<\/jats:italic> are reported. <jats:italic>Pm<\/jats:italic>AP-2 \u03b2 was up-regulated about 4-fold at 6 and 36\u2009h post-WSSV infection. Knockdown of <jats:italic>Pm<\/jats:italic>AP-2\u03b2 delayed shrimp mortality during WSSV infection, of which WSSV intermediate early 1 gene expression was also down-regulated. Immunogold-labelling and transmission electron microscopy revealed that <jats:italic>Pm<\/jats:italic>AP-2\u03b2 co-localized with WSSV particles at plasma membrane. In addition, <jats:italic>Pm<\/jats:italic>AP-2\u03b2-silencing significantly affected the expression levels of <jats:italic>Pm<\/jats:italic>STAT, <jats:italic>Pm<\/jats:italic>DOME, <jats:italic>Pm<\/jats:italic>Dorsal and ALF<jats:italic>Pm<\/jats:italic>3 during WSSV infection. It is possible that <jats:italic>Pm<\/jats:italic>AP-2\u03b2 is associated with the JAK\/STAT and the Toll pathway.<\/jats:p>","DOI":"10.1038\/s41598-019-49852-0","type":"journal-article","created":{"date-parts":[[2019,9,17]],"date-time":"2019-09-17T10:04:52Z","timestamp":1568714692000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":10,"title":["Role of Clathrin Assembly Protein-2 Beta Subunit during White Spot Syndrome Virus Infection in Black Tiger Shrimp Penaeus monodon"],"prefix":"10.1038","volume":"9","author":[{"given":"Thapanan","family":"Jatuyosporn","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Pasunee","family":"Laohawutthichai","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Premruethai","family":"Supungul","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5543-6889","authenticated-orcid":false,"given":"Rogerio R.","family":"Sotelo-Mundo","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Adrian","family":"Ochoa-Leyva","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Anchalee","family":"Tassanakajon","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0614-8650","authenticated-orcid":false,"given":"Kuakarun","family":"Krusong","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2019,9,17]]},"reference":[{"key":"49852_CR1","doi-asserted-by":"publisher","first-page":"146","DOI":"10.20506\/rst.16.1.1010","volume":"16","author":"DV Lightner","year":"1997","unstructured":"Lightner, D. V. et al. Risk of spread of penaeid shrimp viruses in the Americas by the international movement of live and frozen shrimp. Rev. Sci. Tech. 16, 146\u2013160 (1997).","journal-title":"Rev. Sci. Tech."},{"key":"49852_CR2","doi-asserted-by":"publisher","first-page":"1655","DOI":"10.1007\/s007050200039","volume":"147","author":"MA Mayo","year":"2002","unstructured":"Mayo, M. A. A summary of taxonomic changes recently approved by ICTV. Arch. Virol. 147, 1655\u20131663 (2002).","journal-title":"Arch. Virol."},{"key":"49852_CR3","doi-asserted-by":"publisher","first-page":"1071","DOI":"10.1007\/s007050200036","volume":"147","author":"MA Mayo","year":"2002","unstructured":"Mayo, M. A. Virus taxonomy - Houston 2002. Arch. Virol. 147, 1071\u20131076 (2002).","journal-title":"Arch. Virol."},{"key":"49852_CR4","doi-asserted-by":"publisher","first-page":"131","DOI":"10.3354\/dao027131","volume":"27","author":"P Chang","year":"1996","unstructured":"Chang, P., Lo, C., Wang, Y. & Kou, G. Identification of white spot syndrome associated baculovirus target organs in the shrimp Penaeus monodon by in situ hybridization. Dis. Aquat. Org. 27, 131\u2013139 (1996).","journal-title":"Dis. Aquat. Org."},{"key":"49852_CR5","doi-asserted-by":"publisher","first-page":"269","DOI":"10.1111\/j.1439-0426.1998.tb00654.x","volume":"14","author":"TW Flegel","year":"1998","unstructured":"Flegel, T. W. & Alday-Sanz, V. The crisis in Asian shrimp aquaculture: current status and future needs. J. Appl. Ichthyol. 14, 269\u2013273 (1998).","journal-title":"J. Appl. Ichthyol."},{"key":"49852_CR6","doi-asserted-by":"publisher","first-page":"71","DOI":"10.3354\/dao034071","volume":"34","author":"JH Park","year":"1998","unstructured":"Park, J. H., Lee, Y. S., Lee, S. & Lee, Y. An infectious viral disease of penaeid shrimp newly found in Korea. Dis. Aquat. Organ. 34, 71\u201375 (1998).","journal-title":"Dis. Aquat. Organ."},{"key":"49852_CR7","doi-asserted-by":"publisher","first-page":"183","DOI":"10.1046\/j.1365-2761.1999.00162.x","volume":"22","author":"KV Rajendran","year":"1999","unstructured":"Rajendran, K. V., Vijayan, K. K., Santiago, T. C. & Krol, R. M. Experimental host range and histopathology of white spot syndrome virus (WSSV) infection in shrimp, prawns, crayfish and lobsters from India. J. Fish Dis. 22, 183\u2013191 (1999).","journal-title":"J. Fish Dis."},{"key":"49852_CR8","doi-asserted-by":"publisher","first-page":"63","DOI":"10.1016\/j.virusres.2004.08.002","volume":"108","author":"X Xie","year":"2005","unstructured":"Xie, X., Li, H., Xu, L. & Yang, F. A simple and efficient method for purification of intact white spot syndrome virus (WSSV) viral particles. Virus Res. 108, 63\u201367 (2005).","journal-title":"Virus Res."},{"key":"49852_CR9","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1006\/viro.2001.1002","volume":"286","author":"MC van Hulten","year":"2001","unstructured":"van Hulten, M. C. et al. The white spot syndrome virus DNA genome sequence. Virology 286, 7\u201322 (2001).","journal-title":"Virology"},{"key":"49852_CR10","doi-asserted-by":"publisher","first-page":"11811","DOI":"10.1128\/JVI.75.23.11811-11820.2001","volume":"75","author":"F Yang","year":"2001","unstructured":"Yang, F. et al. Complete genome sequence of the shrimp white spot bacilliform virus. J. Virol. 75, 11811\u201311820 (2001).","journal-title":"J. Virol."},{"key":"49852_CR11","doi-asserted-by":"publisher","first-page":"136","DOI":"10.1006\/viro.2002.1536","volume":"301","author":"LL Chen","year":"2002","unstructured":"Chen, L. L. et al. Transcriptional analysis of the DNA polymerase gene of shrimp white spot syndrome virus. Virology 301, 136\u2013147 (2002).","journal-title":"Virology"},{"key":"49852_CR12","doi-asserted-by":"publisher","first-page":"309","DOI":"10.1016\/j.fsi.2016.03.007","volume":"52","author":"XF Wang","year":"2016","unstructured":"Wang, X. F., Liu, Q. H., Wu, Y. & Huang, J. Litopenaeus vannamei clathrin coat AP17 involved in white spot syndrome virus infection. Fish. Shellfish. Immunol. 52, 309\u2013316 (2016).","journal-title":"Fish. Shellfish. Immunol."},{"key":"49852_CR13","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1016\/j.virol.2015.08.034","volume":"486","author":"J Huang","year":"2015","unstructured":"Huang, J., Li, F., Wu, J. & Yang, F. White spot syndrome virus enters crayfish hematopoietic tissue cells via clathrin-mediated endocytosis. Virology 486, 35\u201343 (2015).","journal-title":"Virology"},{"key":"49852_CR14","doi-asserted-by":"publisher","DOI":"10.1038\/srep28694","volume":"6","author":"RY Chen","year":"2016","unstructured":"Chen, R. Y. et al. White spot syndrome virus entry is dependent on multiple endocytic routes and strongly facilitated by Cq-GABARAP in a CME-dependent manner. Sci. Rep. 6, 28694 (2016).","journal-title":"Sci. Rep."},{"key":"49852_CR15","doi-asserted-by":"publisher","first-page":"19","DOI":"10.1016\/S0005-2760(98)00051-4","volume":"1393","author":"C Bernardes","year":"1998","unstructured":"Bernardes, C., Antonio, A., Pedroso de Lima, M. C. & Valdeira, M. L. Cholesterol affects African swine fever virus infection. Biochim. Biophys. Acta 1393, 19\u201325 (1998).","journal-title":"Biochim. Biophys. Acta"},{"key":"49852_CR16","doi-asserted-by":"publisher","first-page":"4080","DOI":"10.1128\/JVI.79.7.4080-4089.2005","volume":"79","author":"M Husain","year":"2005","unstructured":"Husain, M. & Moss, B. Role of receptor-mediated endocytosis in the formation of vaccinia virus extracellular enveloped particles. J. Virol. 79, 4080\u20134089 (2005).","journal-title":"J. Virol."},{"key":"49852_CR17","doi-asserted-by":"publisher","first-page":"13047","DOI":"10.1128\/JVI.01744-14","volume":"88","author":"S Wang","year":"2014","unstructured":"Wang, S. et al. Entry of a novel marine DNA virus, Singapore grouper iridovirus, into host cells occurs via clathrin-mediated endocytosis and macropinocytosis in a pH-dependent manner. J. Virol. 88, 13047\u201313063 (2014).","journal-title":"J. Virol."},{"key":"49852_CR18","doi-asserted-by":"publisher","first-page":"13433","DOI":"10.1128\/JVI.77.24.13433-13438.2003","volume":"77","author":"G Simmons","year":"2003","unstructured":"Simmons, G. et al. Folate receptor alpha and caveolae are not required for Ebola virus glycoprotein-mediated viral infection. J. Virol. 77, 13433\u201313438 (2003).","journal-title":"J. Virol."},{"key":"49852_CR19","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.virol.2005.12.027","volume":"348","author":"B Bartosch","year":"2006","unstructured":"Bartosch, B. & Cosset, F. L. Cell entry of hepatitis C virus. Virology 348, 1\u201312 (2006).","journal-title":"Virology"},{"key":"49852_CR20","doi-asserted-by":"publisher","first-page":"11705","DOI":"10.1128\/JVI.79.18.11705-11715.2005","volume":"79","author":"T Suzuki","year":"2005","unstructured":"Suzuki, T. et al. Sialidase activity of influenza A virus in an endocytic pathway enhances viral replication. J. Virol. 79, 11705\u201311715 (2005).","journal-title":"J. Virol."},{"key":"49852_CR21","doi-asserted-by":"publisher","first-page":"e0140824","DOI":"10.1371\/journal.pone.0140824","volume":"10","author":"LE Piccini","year":"2015","unstructured":"Piccini, L. E., Castilla, V. & Damonte, E. B. Dengue-3 Virus Entry into Vero Cells: Role of Clathrin-Mediated Endocytosis in the Outcome of Infection. PLoS One 10, e0140824 (2015).","journal-title":"PLoS One"},{"key":"49852_CR22","doi-asserted-by":"publisher","first-page":"100","DOI":"10.1016\/j.dci.2013.11.017","volume":"44","author":"T Jatuyosporn","year":"2014","unstructured":"Jatuyosporn, T., Supungul, P., Tassanakajon, A. & Krusong, K. The essential role of clathrin-mediated endocytosis in yellow head virus propagation in the black tiger shrimp Penaeus monodon. Dev. Comp. Immunol. 44, 100\u2013110 (2014).","journal-title":"Dev. Comp. Immunol."},{"key":"49852_CR23","doi-asserted-by":"crossref","unstructured":"Park, S. Y. & Guo, X. Adaptor protein complexes and intracellular transport. Biosci. Rep. 34 (2014).","DOI":"10.1042\/BSR20140069"},{"key":"49852_CR24","doi-asserted-by":"publisher","first-page":"313","DOI":"10.1038\/nrm.2017.132","volume":"19","author":"M Kaksonen","year":"2018","unstructured":"Kaksonen, M. & Roux, A. Mechanisms of clathrin-mediated endocytosis. Nat. Rev. Mol. Cell Biol. 19, 313\u2013326 (2018).","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"49852_CR25","doi-asserted-by":"publisher","first-page":"2148","DOI":"10.1093\/emboj\/17.8.2148","volume":"17","author":"I Rapoport","year":"1998","unstructured":"Rapoport, I., Chen, Y. C., Cupers, P., Shoelson, S. E. & Kirchhausen, T. Dileucine-based sorting signals bind to the beta chain of AP-1 at a site distinct and regulated differently from the tyrosine-based motif-binding site. EMBO J. 17, 2148\u20132155 (1998).","journal-title":"EMBO J."},{"key":"49852_CR26","doi-asserted-by":"publisher","first-page":"27160","DOI":"10.1074\/jbc.272.43.27160","volume":"272","author":"RC Aguilar","year":"1997","unstructured":"Aguilar, R. C., Ohno, H., Roche, K. W. & Bonifacino, J. S. Functional domain mapping of the clathrin-associated adaptor medium chains mu1 and mu2. J. Biol. Chem. 272, 27160\u201327166 (1997).","journal-title":"J. Biol. Chem."},{"key":"49852_CR27","doi-asserted-by":"publisher","first-page":"a016790","DOI":"10.1101\/cshperspect.a016790","volume":"5","author":"LM Traub","year":"2013","unstructured":"Traub, L. M. & Bonifacino, J. S. Cargo recognition in clathrin-mediated endocytosis. Cold Spring Harb. Perspect. Biol. 5, a016790 (2013).","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"49852_CR28","doi-asserted-by":"publisher","first-page":"523","DOI":"10.1016\/S0092-8674(02)00735-3","volume":"109","author":"BM Collins","year":"2002","unstructured":"Collins, B. M., McCoy, A. J., Kent, H. M., Evans, P. R. & Owen, D. J. Molecular architecture and functional model of the endocytic AP2 complex. Cell 109, 523\u2013535 (2002).","journal-title":"Cell"},{"key":"49852_CR29","doi-asserted-by":"publisher","first-page":"2612","DOI":"10.1073\/pnas.86.8.2612","volume":"86","author":"T Kirchhausen","year":"1989","unstructured":"Kirchhausen, T. et al. Structural and functional division into two domains of the large (100- to 115-kDa) chains of the clathrin-associated protein complex AP-2. Proc. Natl. Acad. Sci. USA 86, 2612\u20132616 (1989).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"49852_CR30","doi-asserted-by":"publisher","first-page":"1096","DOI":"10.1073\/pnas.97.3.1096","volume":"97","author":"E ter Haar","year":"2000","unstructured":"ter Haar, E., Harrison, S. C. & Kirchhausen, T. Peptide-in-groove interactions link target proteins to the beta-propeller of clathrin. Proc. Natl. Acad. Sci. USA 97, 1096\u20131100 (2000).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"49852_CR31","doi-asserted-by":"publisher","first-page":"459","DOI":"10.1126\/science.1254836","volume":"345","author":"BT Kelly","year":"2014","unstructured":"Kelly, B. T. et al. Clathrin adaptors. AP2 controls clathrin polymerization with a membrane-activated switch. Science 345, 459\u2013463 (2014).","journal-title":"Science"},{"key":"49852_CR32","doi-asserted-by":"publisher","first-page":"896","DOI":"10.1016\/S0960-9822(01)00240-8","volume":"11","author":"O Olusanya","year":"2001","unstructured":"Olusanya, O., Andrews, P. D., Swedlow, J. R. & Smythe, E. Phosphorylation of threonine 156 of the \u03bc2 subunit of the AP2 complex is essential for endocytosis in vitro and in vivo. Curr. Biol. 11, 896\u2013900 (2001).","journal-title":"Curr. Biol."},{"key":"49852_CR33","doi-asserted-by":"publisher","first-page":"921","DOI":"10.1083\/jcb.200108123","volume":"156","author":"SD Conner","year":"2002","unstructured":"Conner, S. D. & Schmid, S. L. Identification of an adaptor-associated kinase, AAK1, as a regulator of clathrin-mediated endocytosis. J. Cell Biol. 156, 921\u2013929 (2002).","journal-title":"J. Cell Biol."},{"key":"49852_CR34","doi-asserted-by":"publisher","first-page":"5476","DOI":"10.1074\/jbc.M009516200","volume":"276","author":"A Fingerhut","year":"2001","unstructured":"Fingerhut, A., von Figura, K. & Honing, S. Binding of AP2 to sorting signals is modulated by AP2 phosphorylation. J. Biol. Chem. 276, 5476\u20135482 (2001).","journal-title":"J. Biol. Chem."},{"key":"49852_CR35","doi-asserted-by":"publisher","first-page":"791","DOI":"10.1083\/jcb.200111068","volume":"156","author":"D Ricotta","year":"2002","unstructured":"Ricotta, D., Conner, S. D., Schmid, S. L., von Figura, K. & Honing, S. Phosphorylation of the AP2 mu subunit by AAK1 mediates high affinity binding to membrane protein sorting signals. J. Cell Biol. 156, 791\u2013795 (2002).","journal-title":"J. Cell Biol."},{"key":"49852_CR36","doi-asserted-by":"publisher","first-page":"1491","DOI":"10.1099\/vir.0.009621-0","volume":"90","author":"S Tharntada","year":"2009","unstructured":"Tharntada, S. et al. Role of anti-lipopolysaccharide factor from the black tiger shrimp, Penaeus monodon, in protection from white spot syndrome virus infection. J. Gen. Virol. 90, 1491\u20131498 (2009).","journal-title":"J. Gen. Virol."},{"key":"49852_CR37","doi-asserted-by":"publisher","first-page":"142","DOI":"10.1016\/j.antiviral.2014.08.005","volume":"110","author":"S Suraprasit","year":"2014","unstructured":"Suraprasit, S. et al. Anti-lipopolysaccharide factor isoform 3 from Penaeus monodon (ALFPm3) exhibits antiviral activity by interacting with WSSV structural proteins. Antiviral Res. 110, 142\u2013150 (2014).","journal-title":"Antiviral Res."},{"key":"49852_CR38","doi-asserted-by":"publisher","first-page":"957","DOI":"10.1016\/j.fsi.2013.07.009","volume":"35","author":"S Ponprateep","year":"2013","unstructured":"Ponprateep, S., Phiwsaiya, K., Tassanakajon, A. & Rimphanitchayakit, V. Interaction between Kazal serine proteinase inhibitor SPIPm2 and viral protein WSV477 reduces the replication of white spot syndrome virus. Fish. Shellfish. Immunol. 35, 957\u2013964 (2013).","journal-title":"Fish. Shellfish. Immunol."},{"key":"49852_CR39","doi-asserted-by":"publisher","first-page":"e10597","DOI":"10.1371\/journal.pone.0010597","volume":"5","author":"E Boucrot","year":"2010","unstructured":"Boucrot, E., Saffarian, S., Zhang, R. & Kirchhausen, T. Roles of AP-2 in clathrin-mediated endocytosis. PLoS One 5, e10597 (2010).","journal-title":"PLoS One"},{"key":"49852_CR40","doi-asserted-by":"publisher","first-page":"707","DOI":"10.1016\/j.fsi.2013.05.028","volume":"35","author":"ZJ Huang","year":"2013","unstructured":"Huang, Z. J., Kang, S. T., Leu, J. H. & Chen, L. L. Endocytic pathway is indicated for white spot syndrome virus (WSSV) entry in shrimp. Fish. Shellfish. Immunol. 35, 707\u2013715 (2013).","journal-title":"Fish. Shellfish. Immunol."},{"key":"49852_CR41","doi-asserted-by":"publisher","first-page":"23","DOI":"10.1016\/j.dci.2016.11.023","volume":"69","author":"T Methatham","year":"2017","unstructured":"Methatham, T., Boonchuen, P., Jaree, P., Tassanakajon, A. & Somboonwiwat, K. Antiviral action of the antimicrobial peptide ALFPm3 from Penaeus monodon against white spot syndrome virus. Dev. Comp. Immunol. 69, 23\u201332 (2017).","journal-title":"Dev. Comp. Immunol."},{"key":"49852_CR42","doi-asserted-by":"publisher","first-page":"480","DOI":"10.1016\/j.fsi.2010.05.009","volume":"29","author":"YM Chai","year":"2010","unstructured":"Chai, Y. M., Yu, S. S., Zhao, X. F., Zhu, Q. & Wang, J. X. Comparative proteomic profiles of the hepatopancreas in Fenneropenaeus chinensis response to white spot syndrome virus. Fish. Shellfish. Immunol. 29, 480\u2013486 (2010).","journal-title":"Fish. Shellfish. Immunol."},{"key":"49852_CR43","doi-asserted-by":"publisher","first-page":"e73218","DOI":"10.1371\/journal.pone.0073218","volume":"8","author":"X Chen","year":"2013","unstructured":"Chen, X. et al. Transcriptome analysis of Litopenaeus vannamei in response to white spot syndrome virus infection. PLoS One 8, e73218 (2013).","journal-title":"PLoS One"},{"key":"49852_CR44","doi-asserted-by":"publisher","first-page":"148","DOI":"10.1016\/j.fsi.2016.08.017","volume":"57","author":"J Feng","year":"2016","unstructured":"Feng, J. et al. Toll receptor response to white spot syndrome virus challenge in giant freshwater prawns (Macrobrachium rosenbergii). Fish. Shellfish. Immunol. 57, 148\u2013159 (2016).","journal-title":"Fish. Shellfish. Immunol."},{"key":"49852_CR45","doi-asserted-by":"publisher","first-page":"e1007109","DOI":"10.1371\/journal.ppat.1007109","volume":"14","author":"H Li","year":"2018","unstructured":"Li, H. et al. RNAi screening identifies a new Toll from shrimp Litopenaeus vannamei that restricts WSSV infection through activating Dorsal to induce antimicrobial peptides. PLoS Pathog. 14, e1007109 (2018).","journal-title":"PLoS Pathog."},{"key":"49852_CR46","doi-asserted-by":"publisher","first-page":"88","DOI":"10.1016\/j.fsi.2017.04.005","volume":"65","author":"S Boonrawd","year":"2017","unstructured":"Boonrawd, S. et al. Characterization of PmSptzle 1 from the black tiger shrimp Peneaus monodon. Fish. Shellfish. Immunol. 65, 88\u201395 (2017).","journal-title":"Fish. Shellfish. Immunol."},{"key":"49852_CR47","doi-asserted-by":"publisher","first-page":"3457","DOI":"10.1242\/jcs.005926","volume":"120","author":"O Devergne","year":"2007","unstructured":"Devergne, O., Ghiglione, C. & Noselli, S. The endocytic control of JAK\/STAT signalling in Drosophila. J. Cell Sci. 120, 3457\u20133464 (2007).","journal-title":"J. Cell Sci."},{"key":"49852_CR48","doi-asserted-by":"publisher","first-page":"354","DOI":"10.1016\/j.fsi.2014.04.002","volume":"38","author":"R Wen","year":"2014","unstructured":"Wen, R., Li, F., Li, S. & Xiang, J. Function of shrimp STAT during WSSV infection. Fish. Shellfish. Immunol. 38, 354\u2013360 (2014).","journal-title":"Fish. Shellfish. Immunol."},{"key":"49852_CR49","doi-asserted-by":"publisher","first-page":"662","DOI":"10.1016\/j.fsi.2015.03.031","volume":"44","author":"X Song","year":"2015","unstructured":"Song, X. et al. A Janus Kinase in the JAK\/STAT signaling pathway from Litopenaeus vannamei is involved in antiviral immune response. Fish. Shellfish. Immunol. 44, 662\u2013673 (2015).","journal-title":"Fish. Shellfish. Immunol."},{"key":"49852_CR50","doi-asserted-by":"publisher","first-page":"289","DOI":"10.1016\/j.dci.2016.07.021","volume":"65","author":"S Wang","year":"2016","unstructured":"Wang, S. et al. Shrimp with knockdown of LvSOCS2, a negative feedback loop regulator of JAK\/STAT pathway in Litopenaeus vannamei, exhibit enhanced resistance against WSSV. Dev. Comp. Immunol. 65, 289\u2013298 (2016).","journal-title":"Dev. Comp. Immunol."},{"key":"49852_CR51","doi-asserted-by":"publisher","first-page":"20","DOI":"10.1016\/j.dci.2018.11.003","volume":"92","author":"W Wang","year":"2019","unstructured":"Wang, W., Pan, C., Huang, Z., Yuan, H. & Chen, J. WSV181 inhibits JAK\/STAT signaling and promotes viral replication in Drosophila. Dev. Comp. Immunol. 92, 20\u201328 (2019).","journal-title":"Dev. Comp. Immunol."},{"key":"49852_CR52","doi-asserted-by":"crossref","unstructured":"Mayor, S., Parton, R. G. & Donaldson, J. G. Clathrin-independent pathways of endocytosis. Cold Spring Harb. Perspect. Biol. 6 (2014).","DOI":"10.1101\/cshperspect.a016758"},{"key":"49852_CR53","doi-asserted-by":"publisher","first-page":"3416","DOI":"10.1091\/mbc.02-05-0071","volume":"13","author":"DS Park","year":"2002","unstructured":"Park, D. S. et al. Caveolin-1-deficient mice show accelerated mammary gland development during pregnancy, premature lactation, and hyperactivation of the Jak-2\/STAT5a signaling cascade. Mol. Biol. Cell 13, 3416\u20133430 (2002).","journal-title":"Mol. Biol. Cell"},{"key":"49852_CR54","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-017-06653-7","volume":"7","author":"K Jaturontakul","year":"2017","unstructured":"Jaturontakul, K. et al. Molecular Characterization of Viral Responsive Protein 15 and Its Possible Role in Nuclear Export of Virus in Black Tiger Shrimp Penaeus monodon. Sci. Rep. 7, 6523 (2017).","journal-title":"Sci. Rep."},{"issue":"9","key":"49852_CR55","doi-asserted-by":"publisher","first-page":"45e","DOI":"10.1093\/nar\/29.9.e45","volume":"29","author":"M. W. Pfaffl","year":"2001","unstructured":"Pfaffl, M. W. A new mathematical model for relative quantification in real-time RT-PCR. Nucleic Acids Res. 29, e45 (2001).","journal-title":"Nucleic Acids Research"},{"key":"49852_CR56","first-page":"84","volume":"14","author":"H Liu","year":"2003","unstructured":"Liu, H., Han, H., Li, J. & Wong, L. An in-silico method for prediction of polyadenylation signals in human sequences. Genome informatics. International Conference on Genome Informatics 14, 84\u201393 (2003).","journal-title":"International Conference on Genome Informatics"},{"key":"49852_CR57","doi-asserted-by":"crossref","unstructured":"Gasteiger, E. et al. Protein identification and analysis tools on the ExPASy server. (Humana Press, 2005).","DOI":"10.1385\/1-59259-890-0:571"},{"key":"49852_CR58","doi-asserted-by":"publisher","first-page":"403","DOI":"10.1016\/S0022-2836(05)80360-2","volume":"215","author":"SF Altschul","year":"1990","unstructured":"Altschul, S. F., Gish, W., Miller, W., Myers, E. W. & Lipman, D. J. Basic local alignment search tool. J. Mol. Biol. 215, 403\u2013410 (1990).","journal-title":"J. Mol. Biol."},{"key":"49852_CR59","doi-asserted-by":"publisher","first-page":"539","DOI":"10.1038\/msb.2011.75","volume":"7","author":"F Sievers","year":"2011","unstructured":"Sievers, F. et al. Fast, scalable generation of high-quality protein multiple sequence alignments using Clustal Omega. Mol. Syst. Biol. 7, 539 (2011).","journal-title":"Mol. Syst. Biol."},{"key":"49852_CR60","doi-asserted-by":"publisher","first-page":"W242","DOI":"10.1093\/nar\/gkw290","volume":"44","author":"I Letunic","year":"2016","unstructured":"Letunic, I. & Bork, P. Interactive tree of life (iTOL) v3: an online tool for the display and annotation of phylogenetic and other trees. Nucleic Acids Res. 44, W242\u2013245 (2016).","journal-title":"Nucleic Acids Res."},{"key":"49852_CR61","doi-asserted-by":"publisher","first-page":"W389","DOI":"10.1093\/nar\/gkv332","volume":"43","author":"A Drozdetskiy","year":"2015","unstructured":"Drozdetskiy, A., Cole, C., Procter, J. & Barton, G. J. JPred4: a protein secondary structure prediction server. Nucleic Acids Res. 43, W389\u2013394 (2015).","journal-title":"Nucleic Acids Res."},{"key":"49852_CR62","doi-asserted-by":"publisher","first-page":"D344","DOI":"10.1093\/nar\/gks1067","volume":"41","author":"CJ Sigrist","year":"2013","unstructured":"Sigrist, C. J. et al. New and continuing developments at PROSITE. Nucleic Acids Res. 41, D344\u2013347 (2013).","journal-title":"Nucleic Acids Res."},{"key":"49852_CR63","doi-asserted-by":"publisher","first-page":"D211","DOI":"10.1093\/nar\/gkn785","volume":"37","author":"S Hunter","year":"2009","unstructured":"Hunter, S. et al. InterPro: the integrative protein signature database. Nucleic Acids Res. 37, D211\u2013215 (2009).","journal-title":"Nucleic Acids Res."},{"key":"49852_CR64","doi-asserted-by":"publisher","first-page":"402","DOI":"10.1006\/meth.2001.1262","volume":"25","author":"KJ Livak","year":"2001","unstructured":"Livak, K. J. & Schmittgen, T. D. Analysis of relative gene expression data using real-time quantitative PCR and the 2(-Delta Delta C(T)) Method. Methods 25, 402\u2013408 (2001).","journal-title":"Methods"}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-019-49852-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-019-49852-0","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-019-49852-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,12,17]],"date-time":"2022-12-17T13:38:20Z","timestamp":1671284300000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-019-49852-0"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2019,9,17]]},"references-count":64,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2019,12]]}},"alternative-id":["49852"],"URL":"https:\/\/doi.org\/10.1038\/s41598-019-49852-0","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2019,9,17]]},"assertion":[{"value":"5 June 2019","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"2 September 2019","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"17 September 2019","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing Interests"}}],"article-number":"13489"}}