{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,29]],"date-time":"2026-08-29T09:52:44Z","timestamp":1787997164946,"version":"build-2784847793"},"reference-count":54,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2025,12,19]],"date-time":"2025-12-19T00:00:00Z","timestamp":1766102400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Immunol."],"abstract":"<jats:sec>\n                    <jats:title>Background<\/jats:title>\n                    <jats:p>Tuberculosis remains a leading cause of death from infectious diseases globally, underscoring the need to boost innate responses in monocytes and macrophages to enhance early control of Mycobacterium tuberculosis infection. Trained immunity, a form of innate immune memory, enhances macrophage responsiveness through epigenet ic and metabolic reprogramming, offering a promising approach to strengthen host defenses against M. tuberculosis.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>This study evaluated the immunomodulatory potential of pharmaceutical-grade bacterial suspension (BS) and bacterial lysates (BL) in human monocyte-derived macrophages (MDM) and their role in innate response to M. tuberculosis infection. MDMs were stimulated with M. bovis BCG, BS, and BL following a training protocol described for BCG-dependent trained immunity.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>We observed that BS and BL induced sustained cytokine responses and a metabolic transcriptional profile upon secondary stimulation with M. tuberculosis. BS and BL promoted increased IL-1b production in M. tuberculosis-infected MDMs. Additionally, the expression of surface markers shifted to high levels of CD80, CD86, HLA-DR, TLR2, and CD16 and low expression of CD163, TLR9, CCR2, and TLR4, consistent with an M1 phenotype. Moreover, BS and BL upregulated antimicrobial transcriptional signatures, including autophagy-related MAP1LC3 and ATG16L1.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Discussion<\/jats:title>\n                    <jats:p>These findings indicate that BS and BL engage training-associated transcriptional and phenotypic changes, providing new adjunctive strategies to boost innate responses in tuberculosis and other chronic infections.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.3389\/fimmu.2025.1664444","type":"journal-article","created":{"date-parts":[[2025,12,19]],"date-time":"2025-12-19T06:26:12Z","timestamp":1766125572000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":4,"title":["Immunomodulation by bacterial products promotes innate signatures favorable to macrophage responses in tuberculosis infection"],"prefix":"10.3389","volume":"16","author":[{"given":"D\u00e1maris P.","family":"Romero-Rodr\u00edguez","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Eduardo","family":"Montes","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"H\u00e9ctor Isaac","family":"Rocha-Gonz\u00e1lez","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Martha","family":"Torres","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Joaqu\u00edn","family":"Z\u00fa\u00f1iga","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Esmeralda","family":"Ju\u00e1rez","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2025,12,19]]},"reference":[{"key":"B1","article-title":"Global tuberculosis report","year":"2024"},{"key":"B2","doi-asserted-by":"publisher","first-page":"e0004921","DOI":"10.1128\/cmr.00049-21","article-title":"Trained immunity as a preventive measure for surgical site infections","volume":"34","author":"Ter Steeg","year":"2021","journal-title":"Clin Microbiol Rev"},{"key":"B3","doi-asserted-by":"publisher","first-page":"427","DOI":"10.1126\/science.aaf1098","article-title":"Trained immunity: A program of innate immune memory in health and disease","volume":"352","author":"Netea","year":"2016","journal-title":"Science"},{"key":"B4","doi-asserted-by":"publisher","DOI":"10.1016\/j.chom.2011.04.006","article-title":"Trained immunity: A memory for innate host defense","volume":"9","author":"Netea","year":"2011","journal-title":"Cell Host Microbe"},{"key":"B5","doi-asserted-by":"publisher","first-page":"113821","DOI":"10.1016\/j.addr.2021.05.031","article-title":"Nanoparticles and trained immunity: Glimpse into the future","volume":"175","author":"Magad\u00e1n","year":"2021","journal-title":"Adv Drug Deliv Rev"},{"key":"B6","doi-asserted-by":"publisher","first-page":"176","DOI":"10.1016\/j.cell.2017.12.031","article-title":"BCG educates hematopoietic stem cells to generate protective innate immunity against tuberculosis","volume":"172","author":"Kaufmann","year":"2018","journal-title":"Cell"},{"key":"B7","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2016.11.011","article-title":"Immunometabolic pathways in BCG-induced trained immunity","volume":"17","author":"Arts","year":"2016","journal-title":"Cell Rep"},{"key":"B8","doi-asserted-by":"publisher","DOI":"10.1038\/s41577-020-0285-6","article-title":"Defining trained immunity and its role in health and disease","volume":"20","author":"Netea","year":"2020","journal-title":"Nat Rev Immunol"},{"key":"B9","doi-asserted-by":"publisher","DOI":"10.1002\/JLB.MR0318-104R","article-title":"Long-term reprogramming of the innate immune system","volume":"105","author":"Dominguez-Andres","year":"2019","journal-title":"J Leuk Biol"},{"key":"B10","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2018.00298","article-title":"The potential role of trained immunity in autoimmune and autoinflammatory disorders","volume":"9","author":"Arts","year":"2018","journal-title":"Front Immunol"},{"key":"B11","doi-asserted-by":"publisher","DOI":"10.1016\/j.cmet.2016.06.004","article-title":"Itaconate links inhibition of succinate dehydrogenase with macrophage metabolic remodeling and regulation of inflammation","volume":"24","author":"Lampropoulou","year":"2016","journal-title":"Cell Metab"},{"key":"B12","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-018-0052-z","article-title":"Electrophilic properties of itaconate and derivatives regulate the I\u03baB\u03b6-ATF3 inflammatory axis","volume":"556","author":"Bambouskova","year":"2018","journal-title":"Nature"},{"key":"B13","doi-asserted-by":"publisher","first-page":"112658","DOI":"10.1016\/j.celrep.2023.112658","article-title":"Dimethyl itaconate induces long-term innate immune responses and confers protection against infection","volume":"42","author":"Ferreira","year":"2023","journal-title":"Cell Rep"},{"key":"B14","doi-asserted-by":"publisher","DOI":"10.1016\/j.cmet.2016.10.008","article-title":"Glutaminolysis and fumarate accumulation integrate immunometabolic and epigenetic programs in trained immunity","volume":"24","author":"Arts","year":"2016","journal-title":"Cell Metab"},{"key":"B15","doi-asserted-by":"publisher","DOI":"10.2147\/JIR.S363995","article-title":"Induction of trained immunity protects neonatal mice against microbial sepsis by boosting both the inflammatory response and antimicrobial activity","volume":"15","author":"Zhou","year":"2022","journal-title":"J Inflammation Res"},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.2100107","article-title":"\u03b2-glucan induces distinct and protective innate immune memory in differentiated macrophages","volume":"207","author":"Stothers","year":"2021","journal-title":"J Immunol"},{"key":"B17","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1016\/j.chom.2017.12.010","article-title":"BCG Vaccination Protects against Experimental Viral Infection in Humans through the Induction of Cytokines Associated with Trained Immunity","volume":"23","author":"Arts","year":"2018","journal-title":"Cell Host Microbe"},{"key":"B18","doi-asserted-by":"publisher","DOI":"10.1080\/22221751.2021.1899771","article-title":"Trained immunity contributes to the prevention of Mycobacterium tuberculosis infection, a novel role of autophagy","volume":"10","author":"Zhou","year":"2021","journal-title":"Emerg Microbes Infect"},{"key":"B19","article-title":"Mecanismos moleculares de la respuesta inmune en la tuberculosis pulmonar humana","volume-title":"Rev Inst Nal Enf Resp Mex","author":"Herrera","year":""},{"key":"B20","doi-asserted-by":"publisher","DOI":"10.3109\/1040841X.2012.706249","article-title":"Recognition of bacterial infection by innate immune sensors","volume":"39","author":"Kumar","year":"2013","journal-title":"Crit Rev Microbiol"},{"key":"B21","doi-asserted-by":"publisher","first-page":"102","DOI":"10.3390\/pathogens9020102","article-title":"Effect and analysis of bacterial lysates for the treatment of recurrent urinary tract infections in adults","volume":"9","author":"Ahumada-Cota","year":"2020","journal-title":"Pathogens"},{"key":"B22","doi-asserted-by":"publisher","DOI":"10.1007\/s00405-023-08035-4","article-title":"Bacterial immunotherapy is highly effective in reducing recurrent upper respiratory tract infections in children: a prospective observational study","volume":"280","author":"Rebolledo","year":"2023","journal-title":"Eur Arch Otolaryngol"},{"key":"B23","doi-asserted-by":"publisher","DOI":"10.1177\/20503121241309514","article-title":"Long-term sublingual bacterial immunotherapy prevents ear, nose and throat infections: A real-life study","volume":"13","author":"Aarestrup","year":"2025","journal-title":"SAGE Open Med"},{"key":"B24","doi-asserted-by":"publisher","first-page":"111284","DOI":"10.1016\/j.isci.2024.111284","article-title":"Staphylococcus aureus induced trained immunity in macrophages confers heterologous protection against gram-negative bacterial infection","volume":"27","author":"Carlile","year":"2024","journal-title":"iScience"},{"key":"B25","doi-asserted-by":"publisher","DOI":"10.1164\/rccm.200503-411OC","article-title":"Mycobacterium tuberculosis growth control by lung macrophages and CD8 cells from patient contacts","volume":"173","author":"Carranza","year":"2006","journal-title":"Am J Respir Crit Care Med"},{"key":"B26","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2022.747799","article-title":"Macrophage: A cell with many faces and functions in tuberculosis","volume":"13","author":"Ahmad","year":"2022","journal-title":"Front Immunol"},{"key":"B27","doi-asserted-by":"publisher","first-page":"e2400413121","DOI":"10.1073\/pnas.2400413121","article-title":"High-throughput screen identifies non inflammatory small molecule inducers of trained immunity","volume":"121","author":"Knight","year":"2024","journal-title":"Proc Natl Acad Sci U.S.A"},{"key":"B28","doi-asserted-by":"publisher","first-page":"151951","DOI":"10.1016\/j.imbio.2020.151951","article-title":"Natural and trained innate immunity against Mycobacterium tuberculosis","volume":"225","author":"Ferluga","year":"2020","journal-title":"Immunobiology"},{"key":"B29","doi-asserted-by":"publisher","first-page":"2109","DOI":"10.3390\/cells9092109","article-title":"Lessons from bacillus calmette-gu\u00e9rin: harnessing trained immunity for vaccine development","volume":"9","author":"Pasco","year":"2020","journal-title":"Cells"},{"key":"B30","doi-asserted-by":"publisher","DOI":"10.1016\/j.cmi.2019.02.015","article-title":"Trained innate immunity and resistance to Mycobacterium tuberculosis infection","volume":"25","author":"Koeken","year":"2019","journal-title":"Clin Microbiol Infect"},{"key":"B31","doi-asserted-by":"publisher","first-page":"109208","DOI":"10.1016\/j.clim.2022.109208","article-title":"The impact of BCG dose and revaccination on trained immunity","volume":"246","author":"Debisarun","year":"2023","journal-title":"Clin Immunol"},{"key":"B32","doi-asserted-by":"publisher","first-page":"107634","DOI":"10.1016\/j.celrep.2020.107634","article-title":"\u03b2-Glucan Induces Protective Trained Immunity against Mycobacterium tuberculosis Infection: A Key Role for IL-1","volume":"31","author":"Moorlag","year":"2020","journal-title":"Cell Rep"},{"key":"B33","doi-asserted-by":"publisher","DOI":"10.1080\/22221751.2021.1899771","article-title":"Trained immunity contributes to the prevention of Mycobacterium tuberculosis infection, a novel role of autophagy","volume":"10","author":"Zhou","year":"2021","journal-title":"Emerg Microbes Infect"},{"key":"B34","doi-asserted-by":"publisher","first-page":"206","DOI":"10.29262\/ram.v70i3.1292","article-title":"Efectividad de la suspensi\u00f3n bacteriana sublingual en la prevenci\u00f3n de infecciones de v\u00edas a\u00e9reas de repetici\u00f3n en pediatr\u00eda","volume":"70","author":"Cervantes-Trujano","year":"2023","journal-title":"Rev allerg Mex"},{"key":"B35","doi-asserted-by":"publisher","DOI":"10.1016\/j.jval.2014.08.373","article-title":"The effect of bacterial lysates on patients with recurrent respiratory tract infections: A meta-analysis","volume":"17","author":"Ma","year":"2014","journal-title":"Val Health"},{"key":"B36","article-title":"Bacterial lysate in the prevention of acute exacerbation of COPD and in respiratory recurrent infections","volume":"2","author":"Braido","year":"2007","journal-title":"Int J COPD"},{"key":"B37","doi-asserted-by":"publisher","first-page":"1","DOI":"10.3390\/ijms21228462","article-title":"Exposome and immunity training: How pathogen exposure order influences innate immune cell lineage commitment and function","volume":"21","author":"Adams","year":"2020","journal-title":"Int J Mol Sci"},{"key":"B38","doi-asserted-by":"publisher","DOI":"10.1590\/s0100-879x1999000100002","article-title":"Metabolic fate of glutamine in immune cells","volume":"32","author":"Curi","year":"1999","journal-title":"Braz J Med Biol Res"},{"key":"B39","doi-asserted-by":"publisher","DOI":"10.1016\/j.smim.2016.09.002","article-title":"Immunometabolic circuits in trained immunity","volume":"28","author":"Arts","year":"2016","journal-title":"Semin Immunol"},{"key":"B40","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.198.Supp.208.8","article-title":"Macrophage heterogeneity in human macrophages correlates with autophagy-dependent anti-mycobacterial activity and ability to present antigen to CD4 T cells","volume":"198","author":"Khan","year":"2017","journal-title":"J Immunol"},{"key":"B41","doi-asserted-by":"publisher","first-page":"480","DOI":"10.1038\/s42003-022-03387-9","article-title":"Human M1 macrophages express unique innate immune response genes after mycobacterial infection to defend against tuberculosis","volume":"5","author":"Khan","year":"2022","journal-title":"Commun Biol"},{"key":"B42","doi-asserted-by":"publisher","DOI":"10.2147\/JIR.S448372","article-title":"Identification and validation of genes related to macrophage polarization and cell death modes under mycobacterium tuberculosis infection","volume":"17","author":"Yang","year":"2024","journal-title":"J Inflammation Res"},{"key":"B43","doi-asserted-by":"publisher","first-page":"151951","DOI":"10.1016\/j.imbio.2020.151951","article-title":"Natural and trained innate immunity against Mycobacterium tuberculosis","volume":"225","author":"Ferluga","year":"2020","journal-title":"Immunobiology"},{"key":"B44","doi-asserted-by":"publisher","DOI":"10.1038\/s42255-021-00402-x","article-title":"Pre-operative exercise therapy triggers anti-inflammatory trained immunity of Kupffer cells through metabolic reprogramming","volume":"3","author":"Zhang","year":"2021","journal-title":"Nat Metab"},{"key":"B45","doi-asserted-by":"publisher","first-page":"112471","DOI":"10.1016\/j.celrep.2023.112471","article-title":"The glycolysis\/HIF-1\u03b1 axis defines the inflammatory role of IL-4-primed macrophages","volume":"42","author":"Dang","year":"2023","journal-title":"Cell Rep"},{"key":"B46","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1301686","article-title":"All- Trans Retinoic Acid\u2013Triggered Antimicrobial Activity against Mycobacterium tuberculosis Is Dependent on NPC2","volume":"192","author":"Wheelwright","year":"2014","journal-title":"J Immunol"},{"key":"B47","doi-asserted-by":"publisher","DOI":"10.1002\/eji.201142105","article-title":"NOD2 enhances the innate response of alveolar macrophages to Mycobacterium tuberculosis in humans","volume":"42","author":"Ju\u00e1rez","year":"2012","journal-title":"Eur J Immunol"},{"key":"B48","doi-asserted-by":"publisher","DOI":"10.1128\/IAI.01218-07","article-title":"Expression of cathelicidin LL-37 during Mycobacterium tuberculosis infection in human alveolar macrophages, monocytes, neutrophils, and epithelial cells","volume":"76","author":"Rivas-Santiago","year":"2008","journal-title":"Infect Immun"},{"key":"B49","doi-asserted-by":"publisher","first-page":"268","DOI":"10.3390\/biom12020268","article-title":"High vitamin D concentrations restore the ability to express LL37 by M. tuberculosis-infected human macrophages","volume":"12","author":"Herrera","year":"2022","journal-title":"Biomolecules"},{"key":"B50","doi-asserted-by":"publisher","first-page":"152","DOI":"10.1186\/1471-2466-14-152","article-title":"Nucleotide-oligomerizing domain-1 (NOD1) receptor activation induces pro-inflammatory responses and autophagy in human alveolar macrophages","volume":"14","author":"Ju\u00e1rez","year":"2014","journal-title":"BMC Pulm Med"},{"key":"B51","doi-asserted-by":"publisher","DOI":"10.1128\/IAI.00672-12","article-title":"Cathelicidin antimicrobial peptide expression is not induced or required for bacterial clearance during Salmonella enterica infection of human monocyte-derived macrophages","volume":"80","author":"Strandberg","year":"2012","journal-title":"Infect Immun"},{"key":"B52","doi-asserted-by":"publisher","first-page":"107889","DOI":"10.1016\/j.isci.2023.107889","article-title":"BCG revaccination in adults enhances pro-inflammatory markers of trained immunity along with anti-inflammatory pathways","volume":"26","author":"Ahmed","year":"2023","journal-title":"iScience"},{"key":"B53","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1202870109","article-title":"Bacille Calmette-Gu\u00e9rin induces NOD2-dependent nonspecific protection from reinfection via epigenetic reprogramming of monocytes","volume":"109","author":"Kleinnijenhuis","year":"2012","journal-title":"Proc Natl Acad Sci U.S.A"},{"key":"B54","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2016.11.011","article-title":"Immunometabolic pathways in BCG-induced trained immunity","volume":"17","author":"Arts","year":"2016","journal-title":"Cell Rep"}],"container-title":["Frontiers in Immunology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2025.1664444\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,12,19]],"date-time":"2025-12-19T06:26:15Z","timestamp":1766125575000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2025.1664444\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,12,19]]},"references-count":54,"alternative-id":["10.3389\/fimmu.2025.1664444"],"URL":"https:\/\/doi.org\/10.3389\/fimmu.2025.1664444","relation":{},"ISSN":["1664-3224"],"issn-type":[{"value":"1664-3224","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,12,19]]},"article-number":"1664444"}}