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(Landmark Ed)"],"abstract":"<jats:p>Nematodes are ubiquitous metazoans that occupy diverse ecological niches, exposing them to a wide range of viruses and positioning them as both viral hosts and vectors. Free-living nematodes, particularly Caenorhabditis elegans and Caenorhabditis briggsae, provide powerful genetic models for studying antiviral immune mechanisms. Natural infections with noda-like viruses have revealed powerful antiviral defense strategies in Caenorhabditis species, including RNA interference, the Intracellular Pathogen Response, Signal Transducers and Activators of Transcription (STAT)-like transcriptional regulation, terminal RNA uridylation, and structural barriers to viral entry and egress. These mechanisms demonstrate robust antiviral mechanisms independent of canonical interferon signaling while relying on conserved principles of viral RNA recognition, signal amplification, and coordinated transcriptional responses. Beyond free-living species, parasitic nematodes harbor a diverse and largely unexplored virome. Plant-parasitic nematodes host multiple RNA viruses and act as vectors for economically and agriculturally important plant pathogens, while animal-parasitic nematodes carry persistent viral infections that may influence the immune responses and disease outcomes of their animal hosts. Evidence for functional antiviral RNA interference in parasitic nematodes suggests partial conservation of immune mechanisms, although functional data remain limited. Together, these findings define current knowledge of viral infections and antiviral responses in nematodes and emphasize the need for deeper mechanistic studies, particularly in parasitic species, where the molecular basis of viral recognition and immune defense remains poorly characterized.<\/jats:p>","DOI":"10.31083\/fbl51009","type":"journal-article","created":{"date-parts":[[2026,7,30]],"date-time":"2026-07-30T12:17:49Z","timestamp":1785413869000},"source":"Crossref","is-referenced-by-count":0,"title":["Viruses and Antiviral Responses of Nematodes"],"prefix":"10.31083","volume":"31","author":[{"ORCID":"https:\/\/orcid.org\/0009-0006-8739-8139","authenticated-orcid":true,"given":"Dustin T.","family":"Howard","sequence":"first","affiliation":[{"name":"Department of Biological Sciences, Columbian College of Arts and Sciences, The George Washington University, Washington, DC 20052, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5825-6584","authenticated-orcid":true,"given":"Vladimir","family":"La\u017eeti\u0107","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences, Columbian College of Arts and Sciences, The George Washington University, Washington, DC 20052, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"16242","published-online":{"date-parts":[[2026,7,27]]},"reference":[{"key":"ref1","doi-asserted-by":"publisher","unstructured":"van den Hoogen J, Geisen S, Routh D, Ferris H, Traunspurger W, Wardle DA, et al. 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Global, regional, and national incidence, prevalence, and years lived with disability for 328 diseases and injuries for 195 countries, 1990-2016: a systematic analysis for the Global Burden of Disease Study 2016. Lancet. 2017; 390: 1211\u20131259.","DOI":"10.1016\/S0140-6736(17)32154-2"},{"key":"ref14","doi-asserted-by":"publisher","unstructured":"Chariou PL, Steinmetz NF. Delivery of Pesticides to Plant Parasitic Nematodes Using Tobacco Mild Green Mosaic Virus as a Nanocarrier. ACS Nano. 2017; 11: 4719\u20134730.","DOI":"10.1021\/acsnano.7b00823"},{"key":"ref15","doi-asserted-by":"publisher","unstructured":"Singh S, Singh B, Singh AP. Nematodes: A Threat to Sustainability of Agriculture. Procedia Environmental Sciences. 2015; 29: 215\u2013216.","DOI":"10.1016\/j.proenv.2015.07.270"},{"key":"ref16","doi-asserted-by":"publisher","unstructured":"Garcia S, Hily JM, Komar V, Gertz C, Demangeat G, Lemaire O, et al. 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Journal of Virology. 2014; 88: 11774\u201311781.","DOI":"10.1128\/JVI.01630-14"},{"key":"ref42","doi-asserted-by":"publisher","unstructured":"Hirokawa N, Tilney LG, Fujiwara K, Heuser JE. Organization of actin, myosin, and intermediate filaments in the brush border of intestinal epithelial cells. The Journal of Cell Biology. 1982; 94: 425\u2013443.","DOI":"10.1083\/jcb.94.2.425"},{"key":"ref43","doi-asserted-by":"publisher","unstructured":"Fath KR, Burgess DR. Microvillus assembly. Not actin alone. Current Biology: CB. 1995; 5: 591\u2013593.","DOI":"10.1016\/s0960-9822(95)00117-5"},{"key":"ref44","doi-asserted-by":"publisher","unstructured":"Castiglioni VG, Olmo-Uceda MJ, Mart\u00edn S, F\u00e9lix MA, Gonz\u00e1lez R, Elena SF. Experimental evolution of an RNA virus in Caenorhabditis elegans. Infection, Genetics and Evolution: Journal of Molecular Epidemiology and Evolutionary Genetics in Infectious Diseases. 2024; 123: 105623.","DOI":"10.1016\/j.meegid.2024.105623"},{"key":"ref45","doi-asserted-by":"publisher","unstructured":"Ranganathan S, Aruscavage PJ, Bass BL. Characterization of Orsay virus replication intermediates in Caenorhabditis elegans reveals links to antiviral RNA interference. Molecular Biology of the Cell. 2026; 37: ar35.","DOI":"10.1091\/mbc.E26-01-0031"},{"key":"ref46","doi-asserted-by":"publisher","unstructured":"Castiglioni VG, Villena-Gim\u00e9nez A, Herek D, Gonz\u00e1lez-S\u00e1nchez A, Toft C, G\u00f3mez GG, et al. Latent infection of Caenorhabditis elegans by Orsay virus induces age-dependent immunity and cross-protection. Nature Communications. 2025; 16: 7123.","DOI":"10.1038\/s41467-025-62522-2"},{"key":"ref47","doi-asserted-by":"publisher","unstructured":"Sowa JN, Jiang H, Somasundaram L, Tecle E, Xu G, Wang D, et al. The Caenorhabditis elegans RIG-I Homolog DRH-1 Mediates the Intracellular Pathogen Response upon Viral Infection. Journal of Virology. 2020; 94:  https:\/\/doi.org\/10.1128\/JVI.01173-19","DOI":"10.1128\/jvi.01173-19"},{"key":"ref48","doi-asserted-by":"publisher","unstructured":"Jiang H, Chen K, Sandoval LE, Leung C, Wang D. An Evolutionarily Conserved Pathway Essential for Orsay Virus Infection of Caenorhabditis elegans. mbio. 2017; 8:  https:\/\/doi.org\/10.1128\/mBio.00940-17","DOI":"10.1128\/mbio.00940-17"},{"key":"ref49","doi-asserted-by":"publisher","unstructured":"Wilkins C, Dishongh R, Moore SC, Whitt MA, Chow M, Machaca K. RNA interference is an antiviral defence mechanism in Caenorhabditis elegans. Nature. 2005; 436: 1044\u20131047.","DOI":"10.1038\/nature03957"},{"key":"ref50","doi-asserted-by":"publisher","unstructured":"Gammon DB, Ishidate T, Li L, Gu W, Silverman N, Mello CC. 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Journal of Virology. 2019; 93:  https:\/\/doi.org\/10.1128\/JVI.01400-18","DOI":"10.1128\/jvi.01400-18"},{"key":"ref57","doi-asserted-by":"publisher","unstructured":"Yokoyama N, Miller WT. Biochemical properties of the Cdc42-associated tyrosine kinase ACK1. Substrate specificity, authphosphorylation, and interaction with Hck. The Journal of Biological Chemistry. 2003; 278: 47713\u201347723.","DOI":"10.1074\/jbc.M306716200"},{"key":"ref58","doi-asserted-by":"publisher","unstructured":"Rohatgi R, Nollau P, Ho HY, Kirschner MW, Mayer BJ. Nck and phosphatidylinositol 4,5-bisphosphate synergistically activate actin polymerization through the N-WASP-Arp2\/3 pathway. The Journal of Biological Chemistry. 2001; 276: 26448\u201326452.","DOI":"10.1074\/jbc.M103856200"},{"key":"ref59","doi-asserted-by":"publisher","unstructured":"Consalvo CD, Aderounmu AM, Donelick HM, Aruscavage PJ, Eckert DM, Shen PS, et al. Caenorhabditis elegans Dicer acts with the RIG-I-like helicase DRH-1 and RDE-4 to cleave dsRNA. elife. 2024; 13: RP93979.","DOI":"10.7554\/eLife.93979"},{"key":"ref60","doi-asserted-by":"publisher","unstructured":"Tabara H, Yigit E, Siomi H, Mello CC. The dsRNA binding protein RDE-4 interacts with RDE-1, DCR-1, and a DExH-box helicase to direct RNAi in C. elegans. Cell. 2002; 109: 861\u2013871.","DOI":"10.1016\/s0092-8674(02)00793-6"},{"key":"ref61","doi-asserted-by":"publisher","unstructured":"La\u017eeti\u0107 V, Batachari LE, Russell AB, Troemel ER. Similarities in the induction of the intracellular pathogen response in Caenorhabditis elegans and the type I interferon response in mammals. BioEssays: News and Reviews in Molecular, Cellular and Developmental Biology. 2023; 45: e2300097.","DOI":"10.1002\/bies.202300097"},{"key":"ref62","doi-asserted-by":"publisher","unstructured":"Reddy KC, Dror T, Sowa JN, Panek J, Chen K, Lim ES, et al. An Intracellular Pathogen Response Pathway Promotes Proteostasis in C. elegans. Current Biology: CB. 2017; 27: 3544\u20133553.e5.","DOI":"10.1016\/j.cub.2017.10.009"},{"key":"ref63","doi-asserted-by":"publisher","unstructured":"La\u017eeti\u0107 V, Wu F, Cohen LB, Reddy KC, Chang YT, Gang SS, et al. The transcription factor ZIP-1 promotes resistance to intracellular infection in Caenorhabditis elegans. Nature Communications. 2022; 13: 17.","DOI":"10.1038\/s41467-021-27621-w"},{"key":"ref64","doi-asserted-by":"publisher","unstructured":"Tabuchi TM, Rechtsteiner A, Strome S, Hagstrom KA. Opposing Activities of DRM and MES-4 Tune Gene Expression and X-Chromosome Repression in Caenorhabditis elegans Germ Cells. G3 Genes|Genomes|Genetics. 2014; 4: 143\u2013153.","DOI":"10.1534\/g3.113.007849"},{"key":"ref65","doi-asserted-by":"publisher","unstructured":"Cockrum CS, Strome S. 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The Journal of Clinical Investigation. 2006; 116: 3006\u20133014.","DOI":"10.1172\/JCI29832"},{"key":"ref94","doi-asserted-by":"publisher","unstructured":"Reich NC. Too much of a good thing: Detrimental effects of interferon. Seminars in Immunology. 2019; 43: 101282.","DOI":"10.1016\/j.smim.2019.101282"},{"key":"ref95","doi-asserted-by":"publisher","unstructured":"Gao X, Sun Y. Unraveling the Biology of Interferon-Stimulated Genes: Mechanisms, Functions, and Clinical Implications. Infection and Drug Resistance. 2026; 19: 565301.","DOI":"10.2147\/IDR.S565301"},{"key":"ref96","doi-asserted-by":"publisher","unstructured":"Bakowski MA, Desjardins CA, Smelkinson MG, Dunbar TL, Lopez-Moyado IF, Rifkin SA, et al. Ubiquitin-mediated response to microsporidia and virus infection in C. elegans. PLoS Pathogens. 2014; 10: e1004200.","DOI":"10.1371\/journal.ppat.1004200"},{"key":"ref97","doi-asserted-by":"publisher","unstructured":"Nayak S, Santiago FE, Jin H, Lin D, Schedl T, Kipreos ET. The Caenorhabditis elegans Skp1-related gene family: diverse functions in cell proliferation, morphogenesis, and meiosis. Current Biology: CB. 2002; 12: 277\u2013287.","DOI":"10.1016\/s0960-9822(02)00682-6"},{"key":"ref98","doi-asserted-by":"publisher","unstructured":"Leyva-D\u00edaz E, Stefanakis N, Carrera I, Glenwinkel L, Wang G, Driscoll M, et al. Silencing of Repetitive DNA Is Controlled by a Member of an Unusual Caenorhabditis elegans Gene Family. Genetics. 2017; 207: 529\u2013545.","DOI":"10.1534\/genetics.117.300134"},{"key":"ref99","doi-asserted-by":"publisher","unstructured":"La\u017eeti\u0107 V, Troemel ER. Conservation lost: host-pathogen battles drive diversification and expansion of gene families. The FEBS Journal. 2021; 288: 5289\u20135299.","DOI":"10.1111\/febs.15627"},{"key":"ref100","doi-asserted-by":"publisher","unstructured":"Reddy KC, Dror T, Underwood RS, Osman GA, Elder CR, Desjardins CA, et al. Antagonistic paralogs control a switch between growth and pathogen resistance in C. elegans. PLoS Pathogens. 2019; 15: e1007528.","DOI":"10.1371\/journal.ppat.1007528"},{"key":"ref101","doi-asserted-by":"publisher","unstructured":"La\u017eeti\u0107 V, Blanchard MJ, Bui T, Troemel ER. Multiple pals gene modules control a balance between immunity and development in Caenorhabditis elegans. PLoS Pathogens. 2023; 19: e1011120.","DOI":"10.1371\/journal.ppat.1011120"},{"key":"ref102","doi-asserted-by":"publisher","unstructured":"Fay DS, Yochem J. The SynMuv genes of Caenorhabditis elegans in vulval development and beyond. Developmental Biology. 2007; 306: 1\u20139.","DOI":"10.1016\/j.ydbio.2007.03.016"},{"key":"ref103","doi-asserted-by":"publisher","unstructured":"Ferguson EL, Horvitz HR. The multivulva phenotype of certain Caenorhabditis elegans mutants results from defects in two functionally redundant pathways. Genetics. 1989; 123: 109\u2013121.","DOI":"10.1093\/genetics\/123.1.109"},{"key":"ref104","doi-asserted-by":"publisher","unstructured":"Gal C, Carelli FN, Appert A, Cerrato C, Huang N, Dong Y, et al. DREAM represses distinct targets by cooperating with different THAP domain proteins. Cell Reports. 2021; 37: 109835.","DOI":"10.1016\/j.celrep.2021.109835"},{"key":"ref105","doi-asserted-by":"publisher","unstructured":"Lehner B, Calixto A, Crombie C, Tischler J, Fortunato A, Chalfie M, et al. Loss of LIN-35, the Caenorhabditis elegans ortholog of the tumor suppressor p105Rb, results in enhanced RNA interference. Genome Biology. 2006; 7: R4.","DOI":"10.1186\/gb-2006-7-1-r4"},{"key":"ref106","doi-asserted-by":"publisher","unstructured":"Andersen EC, Lu X, Horvitz HR. C. elegans ISWI and NURF301 antagonize an Rb-like pathway in the determination of multiple cell fates. Development (Cambridge, England). 2006; 133: 2695\u20132704.","DOI":"10.1242\/dev.02444"},{"key":"ref107","doi-asserted-by":"publisher","unstructured":"Rajopadhye S, La\u017eeti\u0107 V, Rodriguez-Crespo D, Troemel E, Meister P, Wicky C. The chromatin remodeler LET-418\/Mi-2 regulates the intracellular pathogen response in the C. elegans intestine. BMC Genomics. 2025; 26: 938.","DOI":"10.1186\/s12864-025-12153-0"},{"key":"ref108","doi-asserted-by":"publisher","unstructured":"Wernet ND, Tecle E, Sarmiento MB, Kuo CJ, Chhan CB, Baick I, et al. Adenosine deaminase and deoxyadenosine regulate intracellular immune response in C. elegans. iscience. 2025; 28: 111950.","DOI":"10.1016\/j.isci.2025.111950"},{"key":"ref109","doi-asserted-by":"publisher","unstructured":"Au-Yeung N, Mandhana R, Horvath CM. Transcriptional regulation by STAT1 and STAT2 in the interferon JAK-STAT pathway. JAK-STAT. 2013; 2: e23931.","DOI":"10.4161\/jkst.23931"},{"key":"ref110","doi-asserted-by":"publisher","unstructured":"Wang WB, Levy DE, Lee CK. STAT3 negatively regulates type I IFN-mediated antiviral response. Journal of Immunology (Baltimore, Md.: 1950). 2011; 187: 2578\u20132585.","DOI":"10.4049\/jimmunol.1004128"},{"key":"ref111","doi-asserted-by":"publisher","unstructured":"Dierking K, Polanowska J, Omi S, Engelmann I, Gut M, Lembo F, et al. Unusual regulation of a STAT protein by an SLC6 family transporter in C. elegans epidermal innate immunity. Cell Host & Microbe. 2011; 9: 425\u2013435.","DOI":"10.1016\/j.chom.2011.04.011"},{"key":"ref112","doi-asserted-by":"publisher","unstructured":"Liu S, Liao Y, Chen B, Chen Y, Yu Z, Wei H, et al. Critical role of Syk-dependent STAT1 activation in innate antiviral immunity. Cell Reports. 2021; 34: 108627.","DOI":"10.1016\/j.celrep.2020.108627"},{"key":"ref113","doi-asserted-by":"publisher","unstructured":"Lim J, Ha M, Chang H, Kwon SC, Simanshu DK, Patel DJ, et al. Uridylation by TUT4 and TUT7 marks mRNA for degradation. Cell. 2014; 159: 1365\u20131376.","DOI":"10.1016\/j.cell.2014.10.055"},{"key":"ref114","doi-asserted-by":"publisher","unstructured":"Zhou Y, Chen H, Zhong W, Tao YJ. Collagen and actin network mediate antiviral immunity against Orsay virus in C. elegans intestinal cells. PLoS Pathogens. 2024; 20: e1011366.","DOI":"10.1371\/journal.ppat.1011366"},{"key":"ref115","doi-asserted-by":"publisher","unstructured":"Stein LD, Bao Z, Blasiar D, Blumenthal T, Brent MR, Chen N, et al. The genome sequence of Caenorhabditis briggsae: a platform for comparative genomics. PLoS Biology. 2003; 1: E45.","DOI":"10.1371\/journal.pbio.0000045"},{"key":"ref116","doi-asserted-by":"crossref","unstructured":"Gupta BP, Johnsen R, Chen N. Genomics and biology of the nematode Caenorhabditis briggsae. 2007. Available at: https:\/\/www.ncbi.nlm.nih.gov\/books\/NBK19675\/ (Accessed: 2 January 2026).","DOI":"10.1895\/wormbook.1.136.1"},{"key":"ref117","doi-asserted-by":"publisher","unstructured":"Guo YR, Fan Y, Zhou Y, Jin M, Zhang JL, Jiang H, et al. Orsay Virus CP-\u03b4 Adopts a Novel \u03b2-Bracelet Structural Fold and Incorporates into Virions as a Head Fiber. Journal of Virology. 2020; 94:  https:\/\/doi.org\/10.1128\/JVI.01560-20","DOI":"10.1128\/jvi.01560-20"},{"key":"ref118","doi-asserted-by":"publisher","unstructured":"Castiglioni VG, Olmo-Uceda MJ, Villena-Gim\u00e9nez A, Mu\u00f1oz-S\u00e1nchez JC, Legarda EG, Elena SF. Story of an infection: Viral dynamics and host responses in the Caenorhabditis elegans-Orsay virus pathosystem. Science Advances. 2024; 10: eadn5945.","DOI":"10.1126\/sciadv.adn5945"},{"key":"ref119","doi-asserted-by":"publisher","unstructured":"Williams SH, Che X, Oleynik A, Garcia JA, Muller D, Zabka TS, et al. Discovery of two highly divergent negative-sense RNA viruses associated with the parasitic nematode, Capillaria hepatica, in wild Mus musculus from New York City. The Journal of General Virology. 2019; 100: 1350\u20131362.","DOI":"10.1099\/jgv.0.001315"},{"key":"ref120","doi-asserted-by":"publisher","unstructured":"Parrado LM, Quintanilla M. Plant-parasitic nematode disease complexes as overlooked challenges to crop production. Frontiers in Plant Science. 2024; 15: 1439951.","DOI":"10.3389\/fpls.2024.1439951"},{"key":"ref121","doi-asserted-by":"publisher","unstructured":"Mendoza-de Gives P. Soil-Borne Nematodes: Impact in Agriculture and Livestock and Sustainable Strategies of Prevention and Control with Special Reference to the Use of Nematode Natural Enemies. Pathogens (Basel, Switzerland). 2022; 11: 640.","DOI":"10.3390\/pathogens11060640"},{"key":"ref122","unstructured":"Brown DJF, Halbrendt TM, Jones AT, Taylor CE, Lamberti F. An appraisal of some aspects of the ecology of nematode vectors of plant viruses. Nematologia Mediterranea. 1994; 22: 253\u2013263."},{"key":"ref123","doi-asserted-by":"publisher","unstructured":"LOEWENBERG JR, SULLIVAN T, SCHUSTER ML. A virus disease of Meloidogyne incognita incognita, the southern root knot nematode. Nature. 1959; 184: 1896.","DOI":"10.1038\/1841896a0"},{"key":"ref124","doi-asserted-by":"publisher","unstructured":"Davis EL, Tylka GL. Soybean cyst nematode disease. The Plant Health Instructor. 2021.","DOI":"10.1094\/PHI-I-2000-0725-01"},{"key":"ref125","unstructured":"Rupprecht CE, Baron S. Medical Microbiology. 4th edn. University of Texas Medical Branch at Galveston: Galveston (TX). 1996."},{"key":"ref126","unstructured":"Wolford RW, Schaefer TJ. StatPearls. StatPearls Publishing: Treasure Island (FL). 2025."},{"key":"ref127","doi-asserted-by":"publisher","unstructured":"Pepin M, Bouloy M, Bird BH, Kemp A, Paweska J. Rift Valley fever virus(Bunyaviridae: Phlebovirus): an update on pathogenesis, molecular epidemiology, vectors, diagnostics and prevention. Veterinary Research. 2010; 41: 61.","DOI":"10.1051\/vetres\/2010033"},{"key":"ref128","doi-asserted-by":"publisher","unstructured":"Ruark CL, Koenning SR, Davis EL, Opperman CH, Lommel SA, Mitchum MG, et al. Soybean cyst nematode culture collections and field populations from North Carolina and Missouri reveal high incidences of infection by viruses. PloS One. 2017; 12: e0171514.","DOI":"10.1371\/journal.pone.0171514"},{"key":"ref129","doi-asserted-by":"publisher","unstructured":"Ruark-Seward CL, Davis EL, Sit TL. Localization of viral and host RNA within soybean cyst nematode via fluorescence in situ hybridization. Experimental Parasitology. 2020; 211: 107866.","DOI":"10.1016\/j.exppara.2020.107866"},{"key":"ref130","doi-asserted-by":"publisher","unstructured":"Overmars H, Mobach M, van de Ruitenbeek SJS, Willig JJ, van Steenbrugge JJM, Goverse A, et al. Plant parasitic cyst nematodes respond to viral infection through RNA interference. bioRxiv. 2025.  (preprint)","DOI":"10.64898\/2025.12.02.691819"},{"key":"ref131","doi-asserted-by":"publisher","unstructured":"Maule AG, McVeigh P, Dalzell JJ, Atkinson L, Mousley A, Marks NJ. An eye on RNAi in nematode parasites. Trends in Parasitology. 2011; 27: 505\u2013513.","DOI":"10.1016\/j.pt.2011.07.004"},{"key":"ref132","doi-asserted-by":"publisher","unstructured":"Urwin PE, Lilley CJ, Atkinson HJ. Ingestion of double-stranded RNA by preparasitic juvenile cyst nematodes leads to RNA interference. Molecular Plant-microbe Interactions: MPMI. 2002; 15: 747\u2013752.","DOI":"10.1094\/MPMI.2002.15.8.747"},{"key":"ref133","doi-asserted-by":"publisher","unstructured":"Yadav BC, Veluthambi K, Subramaniam K. Host-generated double stranded RNA induces RNAi in plant-parasitic nematodes and protects the host from infection. Molecular and Biochemical Parasitology. 2006; 148: 219\u2013222.","DOI":"10.1016\/j.molbiopara.2006.03.013"},{"key":"ref134","doi-asserted-by":"publisher","unstructured":"Huang G, Allen R, Davis EL, Baum TJ, Hussey RS. Engineering broad root-knot resistance in transgenic plants by RNAi silencing of a conserved and essential root-knot nematode parasitism gene. Proceedings of the National Academy of Sciences of the United States of America. 2006; 103: 14302\u201314306.","DOI":"10.1073\/pnas.0604698103"},{"key":"ref135","doi-asserted-by":"publisher","unstructured":"Iqbal S, Fosu-Nyarko J, Jones MGK. Attempt to Silence Genes of the RNAi Pathways of the Root-Knot Nematode, Meloidogyne incognita Results in Diverse Responses Including Increase and No Change in Expression of Some Genes. Frontiers in Plant Science. 2020; 11: 328.","DOI":"10.3389\/fpls.2020.00328"},{"key":"ref136","doi-asserted-by":"publisher","unstructured":"Vos T, Lim SS, Abbafati C, Abbas KM, Abbasi M, Abbasifard M, et al. Global burden of 369 diseases and injuries in 204 countries and territories, 1990-2019: a systematic analysis for the Global Burden of Disease Study 2019. Lancet. 2020; 396: 1204\u20131222.","DOI":"10.1016\/S0140-6736(20)30925-9"},{"key":"ref137","unstructured":"de Lima Corvino DF, Horrall S. StatPearls. StatPearls Publishing: Treasure Island (FL). 2025."},{"key":"ref138","doi-asserted-by":"publisher","unstructured":"Servi\u00e1n A, Repetto SA, Lorena Zonta M, Navone GT. Human hookworms from Argentina: Differential diagnosis of Necator americanus and Ancylostoma duodenale in endemic populations from Buenos Aires and Misiones. Revista Argentina De Microbiologia. 2022; 54: 268\u2013281.","DOI":"10.1016\/j.ram.2022.05.005"},{"key":"ref139","doi-asserted-by":"publisher","unstructured":"Loukas A, Hotez PJ, Diemert D, Yazdanbakhsh M, McCarthy JS, Correa-Oliveira R, et al. Hookworm infection. Nature Reviews. Disease Primers. 2016; 2: 16088.","DOI":"10.1038\/nrdp.2016.88"},{"key":"ref140","unstructured":"Goldin J, Juergens AL. StatPearls. StatPearls Publishing: Treasure Island (FL). 2025."},{"key":"ref141","doi-asserted-by":"publisher","unstructured":"Manguin S, Bangs MJ, Pothikasikorn J, Chareonviriyaphap T. Review on global co-transmission of human Plasmodium species and Wuchereria bancrofti by Anopheles mosquitoes. Infection, Genetics and Evolution: Journal of Molecular Epidemiology and Evolutionary Genetics in Infectious Diseases. 2010; 10: 159\u2013177.","DOI":"10.1016\/j.meegid.2009.11.014"},{"key":"ref142","doi-asserted-by":"publisher","unstructured":"Molyneux D. Lymphatic Filariasis (Elephantiasis) Elimination: A public health success and development opportunity. Filaria Journal. 2003; 2: 13.","DOI":"10.1186\/1475-2883-2-13"},{"key":"ref143","doi-asserted-by":"publisher","unstructured":"Loghry HJ, Kwon H, Smith RC, Sondjaja NA, Minkler SJ, Young S, et al. Extracellular vesicles secreted by Brugia malayi microfilariae modulate the melanization pathway in the mosquito host. Scientific Reports. 2023; 13: 8778.","DOI":"10.1038\/s41598-023-35940-9"},{"key":"ref144","unstructured":"Gyasi ME, Okonkwo ON, Tripathy K. StatPearls. StatPearls Publishing: Treasure Island (FL). 2025."},{"key":"ref145","doi-asserted-by":"publisher","unstructured":"van Sluijs L, Dietz C, van Noort F, Helder J, Zwart MP, Mason-Jones K. Nematodes vector bacteriophages in compost and soil. Soil Biology and Biochemistry. 2025; 205.","DOI":"10.1016\/j.soilbio.2025.109785"},{"key":"ref146","unstructured":"Karanastasi E, Wyss U, Brown D. Observations on the feeding behaviour of Paratrichodorus anemones in relation to tobravirus transmission. Mededelingen (Rijksuniversiteit Te Gent. Fakulteit Van De Landbouwkundige en Toegepaste Biologische Wetenschappen). 2001; 66: 599\u2013608."},{"key":"ref147","doi-asserted-by":"crossref","unstructured":"Singh S, Awasthi LP, Jangre A, Nirmalkar VK. Transmission of plant viruses through soil-inhabiting nematode vectors. Applied Plant Virology (pp. 291\u2013300). Academic Press: Cambridge, MA. 2020.","DOI":"10.1016\/B978-0-12-818654-1.00022-0"},{"key":"ref148","unstructured":"Sirca S, Stare BG, Plesko IM, Marn MV, Urek G. First record of Longidorus juvenilis and L. leptocephalus (Nematoda: Dorylaimida) in Slovenia and their morphometrical and ribosomal DNA sequence analysis. Russian Journal of Nematology. 2007; 15: 1."},{"key":"ref149","unstructured":"Georgi LL. Transmission of Tomato Ringspot Virus by Xiphinema americanum and X. rivesi from New York Apple Orchards. Journal of Nematology. 1988; 20: 304\u2013308."},{"key":"ref150","doi-asserted-by":"publisher","unstructured":"Pinkerton JN, Kraus J, Martin RR, Schreiner RP. Epidemiology of Xiphinema americanum and Tomato ringspot virus on Red Raspberry, Rubus idaeus. Plant Disease. 2008; 92: 364\u2013371.","DOI":"10.1094\/PDIS-92-3-0364"},{"key":"ref151","unstructured":"Gallitelli D, Vovlas C, Martelli GP, Rana GL. Viruses of Globe Artichoke: An Overview. Journal of Plant Pathology. 2004; 86: 267\u2013281."},{"key":"ref152","doi-asserted-by":"publisher","unstructured":"Taylor CE, Robertson WM. Sites of virus retention in the alimentary tract of the nematode vectors, Xiphinema diversicaudatum (Micol.) and X. index (Thorne and Allen). The Annals of Applied Biology. 1970; 66: 375\u2013380.","DOI":"10.1111\/j.1744-7348.1970.tb04616.x"},{"key":"ref153","doi-asserted-by":"publisher","unstructured":"Wang S, Gergerich RC, Wickizer SL, Kim KS. Localization of Transmissible and Nontransmissible Viruses in the Vector Nematode Xiphinema americanum. Phytopathology. 2002; 92: 646\u2013653.","DOI":"10.1094\/PHYTO.2002.92.6.646"},{"key":"ref154","doi-asserted-by":"publisher","unstructured":"Brown DJ, Robertson WM, Trudgill DL. Transmission of viruses by plant nematodes. Annual Review of Phytopathology. 1995; 33: 223\u2013249.","DOI":"10.1146\/annurev.py.33.090195.001255"},{"key":"ref155","unstructured":"Harrison BD, Robertson WM, Taylor CE. Specificity of retention and transmission of viruses by nematodes. Journal of Nematology. 1974; 6: 155\u2013164."},{"key":"ref156","doi-asserted-by":"publisher","unstructured":"Robertson W, Henry CE. An association of carbohydrates with particles of arabis mosaic virus retained within Xiphinema diversicaudatum. Annals of Applied Biology. 1986; 109: 299\u2013305.","DOI":"10.1111\/j.1744-7348.1986.tb05321.x"},{"key":"ref157","doi-asserted-by":"publisher","unstructured":"Demangeat G, Voisin R, Minot JC, Bosselut N, Fuchs M, Esmenjaud D. Survival of Xiphinema index in Vineyard Soil and Retention of Grapevine fanleaf virus Over Extended Time in the Absence of Host Plants. Phytopathology. 2005; 95: 1151\u20131156.","DOI":"10.1094\/PHYTO-95-1151"},{"key":"ref158","doi-asserted-by":"publisher","unstructured":"Das S, Raski DJ. Vector-Efficiency of Xiphinema Index in the Transmission of Grapevine Fanleaf Virus. Nematologica. 1968; 14: 55\u201362.","DOI":"10.1163\/187529268X00642"},{"key":"ref159","doi-asserted-by":"publisher","unstructured":"Goverse A, Mitchum MG. At the molecular plant-nematode interface: New players and emerging paradigms. Current Opinion in Plant Biology. 2022; 67: 102225.","DOI":"10.1016\/j.pbi.2022.102225"},{"key":"ref160","doi-asserted-by":"crossref","unstructured":"Raski DJ, Goheen AC, Lider LA, Meredith CP. Strategies Against Grapevine Fanleaf Virus and Its Nematode Vector. Plant Disease. 1983; 63: 335\u2013339.","DOI":"10.1094\/PD-67-335"},{"key":"ref161","unstructured":"Brown DJF, Weischer B. Specificity, Exclusivity and Complementarity in the Transmission of Plant Viruses by Plant Parasitic Nematodes: an annotated terminology. Fundamental and Applied Nematology. 1998; 21: 1\u201311."},{"key":"ref162","doi-asserted-by":"publisher","unstructured":"Kungu Gbulu R, De Wilde B, Lubbers C, Falay DS, Siewe Fodjo JN, Colebunders R. Exploring potential pathogenesis mechanisms of onchocerciasis-associated epilepsy through cerebrospinal fluid analysis of children with complex febrile seizures: a pilot study in Aketi, Democratic Republic of the Congo. BMJ Open. 2026; 16: e105724.","DOI":"10.1136\/bmjopen-2025-105724"},{"key":"ref163","doi-asserted-by":"publisher","unstructured":"Andret-Link P, Schmitt-Keichinger C, Demangeat G, Komar V, Fuchs M. The specific transmission of Grapevine fanleaf virus by its nematode vector Xiphinema index is solely determined by the viral coat protein. Virology. 2004; 320: 12\u201322.","DOI":"10.1016\/j.virol.2003.11.022"},{"key":"ref164","doi-asserted-by":"publisher","unstructured":"Hadermann A, Amaral LJ, Van Cutsem G, Siewe Fodjo JN, Colebunders R. Onchocerciasis-associated epilepsy: an update and future perspectives. 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