{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,24]],"date-time":"2026-10-24T03:42:44Z","timestamp":1792813364296,"version":"4.5.0"},"reference-count":46,"publisher":"Public Library of Science (PLoS)","issue":"9","license":[{"start":{"date-parts":[[2026,9,8]],"date-time":"2026-09-08T00:00:00Z","timestamp":1788825600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Shenzhen Medical Research Fund","award":["A2304001"],"award-info":[{"award-number":["A2304001"]}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["82130066"],"award-info":[{"award-number":["82130066"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["82572589"],"award-info":[{"award-number":["82572589"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Science and Technology Project of Guangdong Province","award":["2025A1515010675"],"award-info":[{"award-number":["2025A1515010675"]}]}],"content-domain":{"domain":["www.plospathogens.org"],"crossmark-restriction":false},"short-container-title":["PLoS Pathog"],"abstract":"<jats:p>\n                    Tripartite motif-containing 21 (TRIM21) is an E3 ubiquitin ligase that binds viral and host proteins to mediate antiviral defense. Its role in\n                    <jats:italic>Mycobacterium tuberculosis<\/jats:italic>\n                    (\n                    <jats:italic>Mtb<\/jats:italic>\n                    ) infection, however, is unclear. Here, our analysis of human TB transcriptomic datasets showed that TRIM21 expression levels correlate with TB progression in human patients. We further demonstrated that TRIM21 deficiency suppresses\n                    <jats:italic>Mtb<\/jats:italic>\n                    growth in both macrophages and a murine model of tuberculosis (TB). Mechanistically, TRIM21 limits host resistance to\n                    <jats:italic>Mtb<\/jats:italic>\n                    through a non-canonical pathway distinct with its E3 ligase activity. It physically interacts with the RNA helicase DEAD-box helicase family member DDX3X and disrupts phase separation dynamics required for stress granule (SG) assembly. Impaired SG formation compromises innate immune responses, thereby facilitating intracellular bacterial growth and creating a permissive cellular environment for\n                    <jats:italic>Mtb<\/jats:italic>\n                    growth. Consistent with these genetic findings, pharmacological inhibition of TRIM21 restricted\n                    <jats:italic>Mtb<\/jats:italic>\n                    growth in experimental macrophage and murine models. Together, these findings uncover a non-canonical role of TRIM21 in\n                    <jats:italic>Mtb<\/jats:italic>\n                    infection and highlight its potential as a therapeutic target for host-directed therapy against TB.\n                  <\/jats:p>","DOI":"10.1371\/journal.ppat.1014553","type":"journal-article","created":{"date-parts":[[2026,9,8]],"date-time":"2026-09-08T17:47:48Z","timestamp":1788889668000},"page":"e1014553","update-policy":"https:\/\/doi.org\/10.1371\/journal.ppat.corrections_policy","source":"Crossref","is-referenced-by-count":1,"title":["TRIM21 blocks DDX3X-driven stress granule formation during Mycobacterium tuberculosis infection via a non-canonical E3 ligase mechanism"],"prefix":"10.1371","volume":"22","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3005-1728","authenticated-orcid":true,"given":"Ruiyao","family":"Xu","sequence":"first","affiliation":[{"name":"School of Medicine, Shenzhen University","place":["Shenzhen, China"],"department":["Guangdong Provincial Key Laboratory of Infection Immunity and Inflammation, Department of Pathogen Biology"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mengru","family":"Lu","sequence":"additional","affiliation":[{"name":"Guangzhou Medical Research Institute of Infectious Diseases, Infectious Disease Center, Guangzhou Eighth People\u2019s Hospital, Guangzhou Medical University","place":["Guangzhou, Guangdong, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gaoyan","family":"Wang","sequence":"additional","affiliation":[{"name":"Guangzhou Medical Research Institute of Infectious Diseases, Infectious Disease Center, Guangzhou Eighth People\u2019s Hospital, Guangzhou Medical University","place":["Guangzhou, Guangdong, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ximeng","family":"Zhang","sequence":"additional","affiliation":[{"name":"School of Medicine, Shenzhen University","place":["Shenzhen, China"],"department":["Guangdong Provincial Key Laboratory of Infection Immunity and Inflammation, Department of Pathogen Biology"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuxing","family":"Huang","sequence":"additional","affiliation":[{"name":"Guangzhou Medical Research Institute of Infectious Diseases, Infectious Disease Center, Guangzhou Eighth People\u2019s Hospital, Guangzhou Medical University","place":["Guangzhou, Guangdong, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yu","family":"Wang","sequence":"additional","affiliation":[{"name":"School of Medicine, Shenzhen University","place":["Shenzhen, China"],"department":["Guangdong Provincial Key Laboratory of Infection Immunity and Inflammation, Department of Pathogen Biology"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qing","family":"Yu","sequence":"additional","affiliation":[{"name":"Guangzhou Medical Research Institute of Infectious Diseases, Infectious Disease Center, Guangzhou Eighth People\u2019s Hospital, Guangzhou Medical University","place":["Guangzhou, Guangdong, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yaxuan","family":"Wang","sequence":"additional","affiliation":[{"name":"Guangzhou Medical Research Institute of Infectious Diseases, Infectious Disease Center, Guangzhou Eighth People\u2019s Hospital, Guangzhou Medical University","place":["Guangzhou, Guangdong, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dan","family":"Chen","sequence":"additional","affiliation":[{"name":"School of Medicine, Shenzhen University","place":["Shenzhen, China"],"department":["Guangdong Provincial Key Laboratory of Infection Immunity and Inflammation, Department of Pathogen Biology"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yunlong","family":"Hu","sequence":"additional","affiliation":[{"name":"School of Medicine, Shenzhen University","place":["Shenzhen, China"],"department":["Guangdong Provincial Key Laboratory of Infection Immunity and Inflammation, Department of Pathogen Biology"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chenyan","family":"Shi","sequence":"additional","affiliation":[{"name":"School of Medicine, Shenzhen University","place":["Shenzhen, China"],"department":["Guangdong Provincial Key Laboratory of Infection Immunity and Inflammation, Department of Pathogen Biology"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yi","family":"Cai","sequence":"additional","affiliation":[{"name":"School of Medicine, Shenzhen University","place":["Shenzhen, China"],"department":["Guangdong Provincial Key Laboratory of Infection Immunity and Inflammation, Department of Pathogen Biology"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Li","family":"Fu","sequence":"additional","affiliation":[{"name":"School of Medicine","place":["Shenzhen, China"],"department":["Guangdong Provincial Key Laboratory of Regional Immunity and Diseases, Department of Pharmacology and International Cancer Center"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xinchun","family":"Chen","sequence":"additional","affiliation":[{"name":"School of Medicine, Shenzhen University","place":["Shenzhen, China"],"department":["Guangdong Provincial Key Laboratory of Infection Immunity and Inflammation, Department of Pathogen Biology"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Youchao","family":"Dai","sequence":"additional","affiliation":[{"name":"Guangzhou Medical Research Institute of Infectious Diseases, Infectious Disease Center, Guangzhou Eighth People\u2019s Hospital, Guangzhou Medical University","place":["Guangzhou, Guangdong, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2026,9,8]]},"reference":[{"key":"ppat.1014553.ref001","volume-title":"Global tuberculosis report 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Autoimmunity"},{"issue":"6","key":"ppat.1014553.ref005","article-title":"TRIM21 restricts influenza A virus replication by ubiquitination-dependent degradation of M1","volume":"19","author":"L Lin","year":"2023","journal-title":"PLoS Pathog"},{"issue":"14","key":"ppat.1014553.ref006","article-title":"TRIM21 Promotes Innate Immune Response to RNA Viral Infection through Lys27-Linked Polyubiquitination of MAVS","volume":"92","author":"B Xue","year":"2018","journal-title":"J Virol"},{"issue":"4","key":"ppat.1014553.ref007","doi-asserted-by":"crossref","first-page":"114095","DOI":"10.1016\/j.celrep.2024.114095","article-title":"Viral infection and spread are inhibited by the polyubiquitination and downregulation of TRPV2 channel by the interferon-stimulated gene TRIM21","volume":"43","author":"Y-Y Guo","year":"2024","journal-title":"Cell Rep"},{"issue":"9","key":"ppat.1014553.ref008","doi-asserted-by":"crossref","first-page":"2456","DOI":"10.4049\/jimmunol.2000048","article-title":"TRIM21 Is Targeted for Chaperone-Mediated Autophagy during Salmonella Typhimurium Infection","volume":"205","author":"NJ Hos","year":"2020","journal-title":"J Immunol"},{"issue":"4","key":"ppat.1014553.ref009","doi-asserted-by":"crossref","first-page":"943","DOI":"10.1080\/15548627.2023.2213983","article-title":"Mycobacterium tuberculosis induces host autophagic ferritin degradation for enhanced iron bioavailability and bacterial growth","volume":"20","author":"Y Dai","year":"2024","journal-title":"Autophagy"},{"issue":"8","key":"ppat.1014553.ref010","article-title":"Mycobacterium tuberculosis hijacks host TRIM21- and NCOA4-dependent ferritinophagy to enhance intracellular growth","volume":"133","author":"Y Dai","year":"2023","journal-title":"J Clin Invest"},{"key":"ppat.1014553.ref011","doi-asserted-by":"crossref","first-page":"156394","DOI":"10.1016\/j.metabol.2025.156394","article-title":"Emerging roles of TRIM in metabolic regulation","volume":"174","author":"J Wang","year":"2026","journal-title":"Metabolism"},{"issue":"1","key":"ppat.1014553.ref012","doi-asserted-by":"crossref","first-page":"495","DOI":"10.1038\/s41419-025-07800-z","article-title":"Stress granules and cell death: crosstalk and potential therapeutic strategies in infectious diseases","volume":"16","author":"H-S Huang","year":"2025","journal-title":"Cell Death Dis"},{"issue":"1","key":"ppat.1014553.ref013","doi-asserted-by":"crossref","first-page":"2308","DOI":"10.1038\/s41467-018-04579-w","article-title":"A modular transcriptional signature identifies phenotypic heterogeneity of human tuberculosis infection","volume":"9","author":"A Singhania","year":"2018","journal-title":"Nat Commun"},{"issue":"9","key":"ppat.1014553.ref014","doi-asserted-by":"crossref","first-page":"1198","DOI":"10.1164\/rccm.201711-2340OC","article-title":"Four-gene pan-African blood signature predicts progression to tuberculosis","volume":"197","author":"S Suliman","year":"2018","journal-title":"American 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response","volume":"207","author":"JM Cliff","year":"2013","journal-title":"J Infect Dis"},{"issue":"7989","key":"ppat.1014553.ref018","doi-asserted-by":"crossref","first-page":"1062","DOI":"10.1038\/s41586-023-06726-w","article-title":"Stress granules plug and stabilize damaged endolysosomal membranes","volume":"623","author":"C Bussi","year":"2023","journal-title":"Nature"},{"issue":"2","key":"ppat.1014553.ref019","doi-asserted-by":"crossref","DOI":"10.1016\/j.cell.2020.03.046","article-title":"G3BP1 Is a Tunable Switch that Triggers Phase Separation to Assemble Stress Granules","volume":"181","author":"P Yang","year":"2020","journal-title":"Cell"},{"issue":"10","key":"ppat.1014553.ref020","doi-asserted-by":"crossref","first-page":"1814","DOI":"10.1261\/rna.1684009","article-title":"Metazoan stress granule assembly is mediated by P-eIF2alpha-dependent and -independent mechanisms","volume":"15","author":"NG Farny","year":"2009","journal-title":"RNA"},{"key":"ppat.1014553.ref021","doi-asserted-by":"crossref","first-page":"108528","DOI":"10.1016\/j.micpath.2026.108528","article-title":"Mycobacterium tuberculosis Rv3875 upregulates TRIM21 expression through the type I interferon\/STAT1 pathway and promotes intracellular growth","volume":"216","author":"R Xu","year":"2026","journal-title":"Microb Pathog"},{"key":"ppat.1014553.ref022","doi-asserted-by":"crossref","first-page":"135251","DOI":"10.1016\/j.ijbiomac.2024.135251","article-title":"Cell-free analysis reveals the role of RG\/RGG motifs in DDX3X phase separation and their potential link to cancer pathogenesis","volume":"279","author":"H Chen","year":"2024","journal-title":"Int J Biol Macromol"},{"issue":"7775","key":"ppat.1014553.ref023","doi-asserted-by":"crossref","first-page":"590","DOI":"10.1038\/s41586-019-1551-2","article-title":"DDX3X acts as a live-or-die checkpoint in stressed cells by regulating NLRP3 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Oncol"},{"issue":"6","key":"ppat.1014553.ref030","doi-asserted-by":"crossref","first-page":"563","DOI":"10.1016\/j.chom.2012.04.012","article-title":"Amino acid starvation induced by invasive bacterial pathogens triggers an innate host defense program","volume":"11","author":"I Tattoli","year":"2012","journal-title":"Cell Host Microbe"},{"issue":"1","key":"ppat.1014553.ref031","article-title":"The PERK\/PKR-eIF2\u03b1 Pathway Negatively Regulates Porcine Hemagglutinating Encephalomyelitis Virus Replication by Attenuating Global Protein Translation and Facilitating Stress Granule Formation","volume":"96","author":"J Shi","year":"2022","journal-title":"J Virol"},{"issue":"11","key":"ppat.1014553.ref032","article-title":"Reprogramming the unfolded protein response for replication by porcine reproductive and respiratory syndrome virus","volume":"15","author":"P Gao","year":"2019","journal-title":"PLoS Pathog"},{"issue":"12","key":"ppat.1014553.ref033","article-title":"Reactive oxygen species-mediated endoplasmic reticulum stress response induces apoptosis of Mycobacterium avium-infected macrophages by activating regulated IRE1-dependent decay pathway","volume":"21","author":"D Go","year":"2019","journal-title":"Cell Microbiol"},{"issue":"3","key":"ppat.1014553.ref034","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1038\/s41419-018-0427-y","article-title":"Inhibition of Sec61-dependent translocation by mycolactone uncouples the integrated stress response from ER stress, driving cytotoxicity via translational activation of ATF4","volume":"9","author":"J Ogbechi","year":"2018","journal-title":"Cell Death Dis"},{"issue":"4","key":"ppat.1014553.ref035","doi-asserted-by":"crossref","first-page":"859","DOI":"10.1016\/j.cell.2013.07.031","article-title":"Inhibition of mTORC1 by astrin and stress granules prevents apoptosis in cancer cells","volume":"154","author":"K Thedieck","year":"2013","journal-title":"Cell"},{"issue":"6","key":"ppat.1014553.ref036","doi-asserted-by":"crossref","first-page":"2450","DOI":"10.1128\/MCB.25.6.2450-2462.2005","article-title":"Sequestration of TRAF2 into stress granules interrupts tumor necrosis factor signaling under stress conditions","volume":"25","author":"WJ Kim","year":"2005","journal-title":"Molecular and Cellular Biology"},{"issue":"16","key":"ppat.1014553.ref037","doi-asserted-by":"crossref","first-page":"6430","DOI":"10.1074\/jbc.RA118.005868","article-title":"The stress granule protein G3BP1 binds viral dsRNA and RIG-I to enhance interferon-\u03b2 response","volume":"294","author":"SS-Y Kim","year":"2019","journal-title":"J Biol Chem"},{"issue":"5","key":"ppat.1014553.ref038","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1016\/j.chom.2007.08.006","article-title":"Inhibition of cytoplasmic mRNA stress granule formation by a viral proteinase","volume":"2","author":"JP 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Rabouw","year":"2016","journal-title":"PLoS Pathog"},{"issue":"10","key":"ppat.1014553.ref042","doi-asserted-by":"crossref","DOI":"10.1371\/journal.ppat.1006677","article-title":"KSHV inhibits stress granule formation by viral ORF57 blocking PKR activation","volume":"13","author":"NR Sharma","year":"2017","journal-title":"PLoS Pathog"},{"issue":"7","key":"ppat.1014553.ref043","doi-asserted-by":"crossref","first-page":"1934","DOI":"10.1080\/15548627.2022.2164427","article-title":"Stress granule homeostasis is modulated by TRIM21-mediated ubiquitination of G3BP1 and autophagy-dependent elimination of stress granules","volume":"19","author":"C Yang","year":"2023","journal-title":"Autophagy"},{"issue":"3","key":"ppat.1014553.ref044","doi-asserted-by":"crossref","first-page":"323","DOI":"10.1111\/joim.12375","article-title":"Reduced expression of TRIM21\/Ro52 predicts poor prognosis in diffuse large B-cell lymphoma patients with and without rheumatic disease","volume":"278","author":"S Brauner","year":"2015","journal-title":"J Intern Med"},{"issue":"4","key":"ppat.1014553.ref045","doi-asserted-by":"crossref","DOI":"10.1128\/mBio.01949-19","article-title":"CD157 Confers Host Resistance to Mycobacterium tuberculosis via TLR2-CD157-PKCzeta-Induced Reactive Oxygen Species Production","volume":"10","author":"Q Yang","year":"2019","journal-title":"mBio"},{"key":"ppat.1014553.ref046","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1016\/j.ymeth.2017.04.020","article-title":"Isolation of yeast and mammalian stress granule cores","volume":"126","author":"JR Wheeler","year":"2017","journal-title":"Methods"}],"container-title":["PLOS 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