{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,10,30]],"date-time":"2025-10-30T06:26:30Z","timestamp":1761805590849,"version":"build-2065373602"},"reference-count":106,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2025,10,30]],"date-time":"2025-10-30T00:00:00Z","timestamp":1761782400000},"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>Introduction<\/jats:title>\n                    <jats:p>\n                      Different innate immune cell types are known to release extracellular traps (ETs) in response to invasive pathogens, including parasites. These ETs function to trap, immobilize, and eventually kill pathogens. In line with this, monocytes and macrophages have been shown to release ETs, known as monocyte\/macrophage extracellular traps (METs).\n                      <jats:italic>Toxoplasma gondii<\/jats:italic>\n                      (\n                      <jats:italic>T. gondii<\/jats:italic>\n                      ) is an apicomplexan zoonotic parasite that infects humans and homeothermic animals. While most studies have focused on prolonged exposure of immune cells to\n                      <jats:italic>T. gondii<\/jats:italic>\n                      , this study characterized the early innate immune reaction of mononuclear phagocytes to vital\n                      <jats:italic>T. gondii<\/jats:italic>\n                      tachyzoites.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>\n                      Primary human and bovine monocytes, monocytic THP-1 cells, and THP-1 cell-derived macrophages (M0-, M1-, and M2-like) were exposed to\n                      <jats:italic>T. gondii<\/jats:italic>\n                      tachyzoites for 4\u00a0h. Scanning electron microscopy (SEM), transmission electron microscopy (TEM), immunofluorescencemicroscopy, and confocal microscopy were used to visualize cell activation and the presence of METs. Additionally, the release of pro-inflammatory cytokines interleukin (IL)-1\u03b2 and IL-6, and expression of Toll-like receptor (TLR) 2 and TLR4 were analyzed.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results and discussion<\/jats:title>\n                    <jats:p>\n                      Microscopic analysis illustrated the activation of all cell types tested within 4\u00a0h of exposure to\n                      <jats:italic>T. gondii<\/jats:italic>\n                      tachyzoites. Numerous tachyzoites were found intracellularly in THP-1 cell-derived M1-like macrophages. Furthermore, the co-localization of extracellular DNA (extDNA) and histones in extracellular web-like fibers proved classical characteristics of extruded\n                      <jats:italic>T. gondii<\/jats:italic>\n                      -induced METs, although this was a rare event. In primary human monocytes, an increased release of IL-1\u03b2 and IL-6 was observed following exposure to\n                      <jats:italic>T. gondii<\/jats:italic>\n                      tachyzoites. When co-stimulated with lipopolysaccharide (LPS), primary human monocytes showed an enhanced release of IL-1\u03b2 and IL-6 in response to\n                      <jats:italic>T. gondii<\/jats:italic>\n                      . In contrast to monocytic THP-1 cells, THP-1 cell-derived M1-like macrophages released IL-1\u03b2 in response to\n                      <jats:italic>T. gondii<\/jats:italic>\n                      tachyzoite exposure. When additionally stimulated by LPS, all THP-1 cell-derived macrophages showed an enhanced release of IL-1\u03b2, and monocytic THP-1 cells an increased release of IL-6 in response to\n                      <jats:italic>T. gondii<\/jats:italic>\n                      tachyzoites. This study provides insights into the early innate immune response of human and bovine mononuclear phagocytes to\n                      <jats:italic>T. gondii<\/jats:italic>\n                      . While cytokine secretion was prominent, MET formation was rare in the early response (i.e. &amp;lt; 4\u00a0h of exposure) to\n                      <jats:italic>T. gondii<\/jats:italic>\n                      tachyzoites.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.3389\/fimmu.2025.1683634","type":"journal-article","created":{"date-parts":[[2025,10,30]],"date-time":"2025-10-30T06:22:22Z","timestamp":1761805342000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Early responses of primary human and bovine monocytes, monocytic THP-1 cells and THP-1 cell-derived macrophages to vital Toxoplasma gondii tachyzoites"],"prefix":"10.3389","volume":"16","author":[{"given":"Dominik","family":"Hanke","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Zahady D.","family":"Vel\u00e1squez","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Kathrin","family":"B\u00fcttner","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Andreas","family":"Krueger","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ralf","family":"Ross","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Andreas","family":"Hecker","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Sybille","family":"Mazurek","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Veronika","family":"Grau","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Anja","family":"Taubert","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Carlos","family":"Hermosilla","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Katrin","family":"Richter","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Iv\u00e1n","family":"Conejeros","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1965","published-online":{"date-parts":[[2025,10,30]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","DOI":"10.1128\/microbiolspec.PFS-0011-2015","article-title":"Toxoplasma gondii as a parasite in food: analysis and control","volume":"4","author":"Hill","year":"2016","journal-title":"Microbiol Spectr"},{"key":"B2","doi-asserted-by":"publisher","first-page":"263","DOI":"10.1186\/s13071-021-04769-4","article-title":"Outbreaks of clinical toxoplasmosis in humans: five decades of personal experience, perspectives and lessons learned","volume":"14","author":"Dubey","year":"2021","journal-title":"Parasit Vectors"},{"key":"B3","doi-asserted-by":"publisher","DOI":"10.1128\/CMR.11.2.267","article-title":"Structures of Toxoplasma gondii tachyzoites, bradyzoites, and sporozoites and biology and development of tissue cysts","volume":"11","author":"Dubey","year":"1998","journal-title":"Clin Microbiol Rev"},{"key":"B4","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2019.00235","article-title":"Long-term impact of Toxoplasma gondii infection on human monocytes","volume":"9","author":"Ehmen","year":"2019","journal-title":"Front Cell Infect Microbiol"},{"key":"B5","doi-asserted-by":"publisher","first-page":"588","DOI":"10.1186\/s13071-020-04445-z","article-title":"The life-cycle of Toxoplasma gondii reviewed using animations","volume":"13","author":"Attias","year":"2020","journal-title":"Parasit Vectors"},{"key":"B6","unstructured":"Pro-inflammatory responses in macrophages during Toxoplasma gondii infection\n          \n          \n            \n              Masek\n              KS\n            \n            \n              Hunter\n              CA\n            \n          \n          Austin, TX, USA\n          Landes Bioscience\n          Madame Curie Bioscience Database\n          \n          2013"},{"key":"B7","doi-asserted-by":"publisher","DOI":"10.1016\/j.pt.2023.07.007","article-title":"Monocytes as primary defenders against Toxoplasma gondii infection","volume":"39","author":"Orchanian","year":"2023","journal-title":"Trends Parasitol"},{"key":"B8","volume-title":"Ultrastrukturelle, immunzytochemische und glykohistochemische Untersuchungen am Blut des Rindes (Bos tauris)","author":"Bitzmann","year":"2009"},{"key":"B9","doi-asserted-by":"publisher","first-page":"595","DOI":"10.1016\/j.immuni.2018.10.005","article-title":"Developmental and functional heterogeneity of monocytes","volume":"49","author":"Guilliams","year":"2018","journal-title":"Immunity"},{"key":"B10","doi-asserted-by":"publisher","DOI":"10.1038\/nri3712","article-title":"Dendritic cells, monocytes and macrophages: a unified nomenclature based on ontogeny","volume":"14","author":"Guilliams","year":"2014","journal-title":"Nat Rev Immunol"},{"key":"B11","doi-asserted-by":"publisher","DOI":"10.1089\/jir.2008.0027","article-title":"Monocyte chemoattractant protein-1 (MCP-1): an overview","volume":"29","author":"Deshmane","year":"2009","journal-title":"J Interferon Cytokine Res"},{"key":"B12","doi-asserted-by":"publisher","DOI":"10.7243\/2055-2394-1-1","article-title":"Sex differences in monocytes and TLR4 associated immune responses; implications for systemic lupus erythematosus (SLE)","volume":"1","author":"Jiang","year":"2014","journal-title":"J Immunother Appl"},{"key":"B13","doi-asserted-by":"publisher","DOI":"10.1097\/MOT.0000000000000313","article-title":"Role of monocytes and macrophages in regulating immune response following lung transplantation","volume":"21","author":"Chiu","year":"2016","journal-title":"Curr Opin Organ Transplant"},{"key":"B14","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.autrev.2015.08.009","article-title":"The critical role of Toll-like receptors - from microbial recognition to autoimmunity: a comprehensive review","volume":"15","author":"Jim\u00e9nez-Dalmaroni","year":"2016","journal-title":"Autoimmun Rev"},{"key":"B15","doi-asserted-by":"publisher","first-page":"125","DOI":"10.1186\/s12967-020-02290-3","article-title":"TLR expression in peripheral monocyte subsets of patients with idiopathic inflammatory myopathies: association with clinical and immunological features","volume":"18","author":"Torres-Ruiz","year":"2020","journal-title":"J Transl Med"},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2023.1140592","article-title":"Activation of endothelial NO synthase and P2X7 receptor modification mediates the cholinergic control of ATP-induced interleukin-1\u03b2 release by mononuclear phagocytes","volume":"14","author":"Richter","year":"2023","journal-title":"Front Immunol"},{"key":"B17","doi-asserted-by":"publisher","DOI":"10.1128\/IAI.06120-11","article-title":"Mannheimia haemolytica and its leukotoxin cause macrophage extracellular trap formation by bovine macrophages","volume":"80","author":"Aulik","year":"2012","journal-title":"Infect Immun"},{"key":"B18","doi-asserted-by":"publisher","DOI":"10.1016\/j.dci.2015.02.002","article-title":"Harbour seal (Phoca vitulina) PMN and monocytes release extracellular traps to capture the apicomplexan parasite Toxoplasma gondii","volume":"50","author":"Reichel","year":"2015","journal-title":"Dev Comp Immunol"},{"key":"B19","doi-asserted-by":"publisher","DOI":"10.1016\/j.vetpar.2016.07.028","article-title":"Eimeria ninakohlyakimovae induces NADPH oxidase-dependent monocyte extracellular trap formation and upregulates IL-12 and TNF-\u03b1, IL-6 and CCL2 gene transcription","volume":"227","author":"P\u00e9rez","year":"2016","journal-title":"Vet Parasitol"},{"key":"B20","doi-asserted-by":"publisher","DOI":"10.1189\/jlb.3MA0916-411R","article-title":"Human blood monocytes are able to form extracellular traps","volume":"102","author":"Granger","year":"2017","journal-title":"J Leukoc Biol"},{"key":"B21","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2017.02016","article-title":"Caprine monocytes release extracellular traps against Neospora caninum in vitro","volume":"8","author":"Yang","year":"2018","journal-title":"Front Immunol"},{"key":"B22","doi-asserted-by":"publisher","DOI":"10.1155\/2022\/7050807","article-title":"Macrophage extracellular traps: current opinions and the state of research regarding various diseases","volume":"2022","author":"Weng","year":"2022","journal-title":"J Immunol Res"},{"key":"B23","doi-asserted-by":"publisher","DOI":"10.3390\/biology12070961","article-title":"MCT-dependent Cryptosporidium parvum-induced bovine monocyte extracellular traps (METs) under physioxia","volume":"12","author":"Hasheminasab","year":"2023","journal-title":"Biology"},{"key":"B24","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41420-024-02034-y","article-title":"Human monocyte subsets differ in their capacity to form extracellular traps","volume":"10","author":"Ibrahim","year":"2024","journal-title":"Cell Death Discov"},{"key":"B25","doi-asserted-by":"publisher","DOI":"10.1126\/science.1092385","article-title":"Neutrophil extracellular traps kill bacteria","volume":"303","author":"Brinkmann","year":"2004","journal-title":"Science"},{"key":"B26","doi-asserted-by":"publisher","DOI":"10.1007\/s00436-014-4094-3","article-title":"Besnoitia besnoiti tachyzoites induce monocyte extracellular trap formation","volume":"113","author":"Mu\u00f1oz-Caro","year":"2014","journal-title":"Parasitol Res"},{"key":"B27","doi-asserted-by":"publisher","DOI":"10.1080\/19396368.2019.1624873","article-title":"Monocyte-derived extracellular trap (MET) formation induces aggregation and affects motility of human spermatozoa","volume":"65","author":"Schulz","year":"2019","journal-title":"vitro Syst Biol Reprod Med"},{"key":"B28","doi-asserted-by":"publisher","DOI":"10.3390\/biology9020034","article-title":"Extracellular traps: an ancient weapon of multiple kingdoms","volume":"9","author":"Neumann","year":"2020","journal-title":"Biology"},{"key":"B29","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1021\/nl902830x","article-title":"Phagocytosis independent extracellular nanoparticle clearance by human immune cells","volume":"10","author":"Bartneck","year":"2010","journal-title":"Nano Lett"},{"key":"B30","doi-asserted-by":"publisher","first-page":"37","DOI":"10.1016\/j.intimp.2014.08.002","article-title":"THP-1 cell line: an in vitro cell model for immune modulation approach","volume":"23","author":"Chanput","year":"2014","journal-title":"Int Immunopharmacol"},{"key":"B31","doi-asserted-by":"publisher","first-page":"577","DOI":"10.1186\/s12885-015-1546-9","article-title":"M1 and M2 macrophages derived from THP-1 cells differentially modulate the response of cancer cells to etoposide","volume":"15","author":"Genin","year":"2015","journal-title":"BMC Cancer"},{"key":"B32","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1016\/j.diff.2022.10.001","article-title":"Macrophage polarization in THP-1 cell line and primary monocytes: a systematic review","volume":"128","author":"Mohd Yasin","year":"2022","journal-title":"Differentiation"},{"key":"B33","doi-asserted-by":"publisher","first-page":"723","DOI":"10.1038\/nri3073","article-title":"Protective and pathogenic functions of macrophage subsets","volume":"11","author":"Murray","year":"2011","journal-title":"Nat Rev Immunol"},{"key":"B34","doi-asserted-by":"publisher","DOI":"10.1002\/cbin.10396","article-title":"Tunicamycin-induced neutrophil extracellular trap (NET)-like structures in cultured human myeloid cell lines","volume":"39","author":"Nakayama","year":"2015","journal-title":"Cell Biol Int"},{"key":"B35","doi-asserted-by":"publisher","DOI":"10.1099\/mic.0.000717","article-title":"Utilization of macrophage extracellular trap nucleotides by Mycoplasma hyopneumoniae","volume":"164","author":"Henthorn","year":"2018","journal-title":"Microbiol Read Engl"},{"key":"B36","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1016\/j.freeradbiomed.2023.03.025","article-title":"New insight into the composition of extracellular traps released by macrophages exposed to different types of inducers","volume":"202","author":"Jensen","year":"2023","journal-title":"Free Radic Biol Med"},{"key":"B37","doi-asserted-by":"publisher","DOI":"10.1007\/s00436-018-6139-5","article-title":"Trichomonas vaginalis triggers the release of THP-1 extracellular traps","volume":"118","author":"Fei","year":"2019","journal-title":"Parasitol Res"},{"key":"B38","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2021.780210","article-title":"Extracellular CIRP induces macrophage extracellular trap formation via gasdermin D activation","volume":"12","author":"Lee","year":"2021","journal-title":"Front Immunol"},{"key":"B39","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2017.00053","article-title":"Aflatoxin B1 induces reactive oxygen species-mediated autophagy and extracellular trap formation in macrophages","volume":"7","author":"An","year":"2017","journal-title":"Front Cell Infect Microbiol"},{"key":"B40","doi-asserted-by":"publisher","DOI":"10.1128\/mBio.02084-18","article-title":"Streptococcus agalactiae induces placental macrophages to release extracellular traps loaded with tissue remodeling enzymes via an oxidative burst-dependent mechanism","volume":"9","author":"Doster","year":"2018","journal-title":"mBio"},{"key":"B41","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0155685","article-title":"Mycobacterium massiliense induces macrophage extracellular traps with facilitating bacterial growth","volume":"11","author":"Je","year":"2016","journal-title":"PloS One"},{"key":"B42","doi-asserted-by":"publisher","DOI":"10.1186\/1742-2094-6-9","article-title":"Lipopolysaccharide-induced interleukin-6 production is controlled by glycogen synthase kinase-3 and STAT3 in the brain","volume":"6","author":"Beurel","year":"2009","journal-title":"J Neuroinflamm"},{"key":"B43","doi-asserted-by":"publisher","DOI":"10.1038\/ni.1863","article-title":"The role of pattern-recognition receptors in innate immunity: update on Toll-like receptors","volume":"11","author":"Kawai","year":"2010","journal-title":"Nat Immunol"},{"key":"B44","doi-asserted-by":"publisher","first-page":"8","DOI":"10.1111\/imr.12621","article-title":"Overview of the IL-1 family in innate inflammation and acquired immunity","volume":"281","author":"Dinarello","year":"2018","journal-title":"Immunol Rev"},{"key":"B45","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2019.03.012","article-title":"Interleukin-1 and related cytokines in the regulation of inflammation and immunity","volume":"50","author":"Mantovani","year":"2019","journal-title":"Immunity"},{"key":"B46","doi-asserted-by":"publisher","DOI":"10.1038\/nri.2016.58","article-title":"Inflammasomes: mechanism of assembly, regulation and signalling","volume":"16","author":"Broz","year":"2016","journal-title":"Nat Rev Immunol"},{"key":"B47","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijpara.2005.12.003","article-title":"Toll-like receptor recognition of Toxoplasma gondii","volume":"36","author":"Yarovinsky","year":"2006","journal-title":"Int J Parasitol"},{"key":"B48","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M110.157586","article-title":"Dendritic cell activation prevents MHC class II ubiquitination and promotes MHC class II survival regardless of the activation stimulus","volume":"285","author":"Walseng","year":"2010","journal-title":"J Biol Chem"},{"key":"B49","doi-asserted-by":"publisher","DOI":"10.1016\/j.pt.2005.08.001","article-title":"Differences among the three major strains of Toxoplasma gondii and their specific interactions with the infected host","volume":"21","author":"Saeij","year":"2005","journal-title":"Trends Parasitol"},{"key":"B50","doi-asserted-by":"publisher","DOI":"10.1080\/21505594.2021.2012346","article-title":"The pathogenicity and virulence of Toxoplasma gondii","volume":"12","author":"Sanchez","year":"2021","journal-title":"Virulence"},{"key":"B51","doi-asserted-by":"publisher","DOI":"10.1038\/nature05395","article-title":"Toxoplasma co-opts host gene expression by injection of a polymorphic kinase homologue","volume":"445","author":"Saeij","year":"2007","journal-title":"Nature"},{"key":"B52","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1084\/jem.20100717","article-title":"Strain-specific activation of the NF-\u03baB pathway by GRA15, a novel Toxoplasma gondii dense granule protein","volume":"208","author":"Rosowski","year":"2011","journal-title":"J Exp Med"},{"key":"B53","doi-asserted-by":"publisher","DOI":"10.1128\/mBio.00255-13","article-title":"Human innate immunity to Toxoplasma gondii is mediated by host caspase-1 and ASC and parasite GRA15","volume":"4","author":"Gov","year":"2013","journal-title":"mBio"},{"key":"B54","doi-asserted-by":"publisher","DOI":"10.14411\/fp.2022.020","article-title":"Polarisation of human macrophages towards an M1 subtype triggered by an atypical Brazilian strain of Toxoplasma gondii results in a reduction in parasite burden","volume":"69","author":"Guimaraes Gois","year":"2022","journal-title":"Folia Parasitol (Praha)"},{"key":"B55","doi-asserted-by":"publisher","first-page":"2855","DOI":"10.4049\/jimmunol.1700245","article-title":"NLRP3 and potassium efflux drive rapid IL-1\u03b2 release from primary human monocytes during Toxoplasma gondii infection","volume":"199","author":"Gov","year":"2017","journal-title":"J Immunol Baltim Md 1950"},{"key":"B56","doi-asserted-by":"publisher","DOI":"10.1128\/mBio.02027-17","article-title":"Evasion of human neutrophil-mediated host defense during Toxoplasma gondii infection","volume":"9","author":"Lima","year":"2018","journal-title":"mBio"},{"key":"B57","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1007923","article-title":"Toxoplasma gondii activates a Syk-CARD9-NF-\u03baB signaling axis and gasdermin D-independent release of IL-1\u03b2 during infection of primary human monocytes","volume":"15","author":"Pandori","year":"2019","journal-title":"PloS Pathog"},{"key":"B58","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.2200513","article-title":"Role for caspase-8 in the release of IL-1\u03b2 and active caspase-1 from viable human monocytes during Toxoplasma gondii infection","volume":"212","author":"Pandori","year":"2024","journal-title":"J Immunol"},{"key":"B59","doi-asserted-by":"publisher","DOI":"10.1093\/infdis\/169.5.1181","article-title":"Interleukin-6 is secreted by human monocytes after stimulation with anti-Toxoplasma gondii sera","volume":"169","author":"Pelloux","year":"1994","journal-title":"J Infect Dis"},{"key":"B60","doi-asserted-by":"publisher","DOI":"10.1177\/1753425918798450","article-title":"Toxoplasma gondii profilin does not stimulate an innate immune response through bovine or human TLR5","volume":"24","author":"Tomb\u00e1cz","year":"2018","journal-title":"Innate Immun"},{"key":"B61","doi-asserted-by":"publisher","DOI":"10.1159\/000362367","article-title":"Toxoplasma gondii-derived profilin triggers human Toll-like receptor 5-dependent cytokine production","volume":"6","author":"Salazar Gonzalez","year":"2014","journal-title":"J Innate Immun"},{"key":"B62","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-2249.1993.tb05896.x","article-title":"Differential cytokine gene expression and secretion after phagocytosis by a human monocytic cell line of Toxoplasma gondii compared with Mycobacterium tuberculosis","volume":"91","author":"Friedland","year":"1993","journal-title":"Clin Exp Immunol"},{"key":"B63","doi-asserted-by":"publisher","DOI":"10.1016\/j.placenta.2012.12.006","article-title":"Trophoblast cells are able to regulate monocyte activity to control Toxoplasma gondii infection","volume":"34","author":"Castro","year":"2013","journal-title":"Placenta"},{"key":"B64","doi-asserted-by":"publisher","DOI":"10.3347\/kjp.2016.54.6.711","article-title":"Production of IL-1\u03b2 and inflammasome with up-regulated expressions of NOD-like receptor related genes in Toxoplasma gondii-infected THP-1 macrophages","volume":"54","author":"Chu","year":"2016","journal-title":"Korean J Parasitol"},{"key":"B65","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0008668","article-title":"The identification of markers of macrophage differentiation in PMA-stimulated THP-1 cells and monocyte-derived macrophages","volume":"5","author":"Daigneault","year":"2010","journal-title":"PloS One"},{"key":"B66","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2019.00012","article-title":"Serum and serum albumin inhibit in vitro formation of neutrophil extracellular traps (NETs)","volume":"10","author":"Neubert","year":"2019","journal-title":"Front Immunol"},{"key":"B67","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2024.1388366","article-title":"Human dendritic cell interactions with the zoonotic parasite Cryptosporidium parvum result in activation and maturation","volume":"15","author":"Ross","year":"2024","journal-title":"Front Immunol"},{"key":"B68","doi-asserted-by":"publisher","DOI":"10.1016\/0022-1759(86)90354-6","article-title":"Improved methods for purification and depletion of monocytes from bovine peripheral blood mononuclear cells. Functional evaluation of monocytes in responses to lectins","volume":"89","author":"Goddeeris","year":"1986","journal-title":"J Immunol Methods"},{"key":"B69","doi-asserted-by":"publisher","first-page":"22","DOI":"10.1016\/j.ymeth.2020.06.017","article-title":"Methods in isolation and characterization of bovine monocytes and macrophages","volume":"186","author":"Ceciliani","year":"2021","journal-title":"Methods San Diego Calif"},{"key":"B70","doi-asserted-by":"publisher","DOI":"10.1038\/nmeth.2019","article-title":"Fiji: an open-source platform for biological-image analysis","volume":"9","author":"Schindelin","year":"2012","journal-title":"Nat Methods"},{"key":"B71","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1016\/j.vetpar.2007.11.013","article-title":"PMN-mediated immune reactions against Eimeria bovis","volume":"151","author":"Behrendt","year":"2008","journal-title":"Vet Parasitol"},{"key":"B72","doi-asserted-by":"publisher","DOI":"10.1016\/0008-8749(77)90230-1","article-title":"Human monocyte cytoplasmic spreading in vitro: early kinetics and scanning electron microscopy","volume":"34","author":"Bumol","year":"1977","journal-title":"Cell Immunol"},{"key":"B73","doi-asserted-by":"publisher","DOI":"10.1111\/cmi.12145","article-title":"Rapid cytoskeleton remodelling in dendritic cells following invasion by Toxoplasma gondii coincides with the onset of a hypermigratory phenotype","volume":"15","author":"Weidner","year":"2013","journal-title":"Cell Microbiol"},{"key":"B74","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2019.00073","article-title":"Toxoplasma-induced hypermigration of primary cortical microglia implicates GABAergic signaling","volume":"9","author":"Bhandage","year":"2019","journal-title":"Front Cell Infect Microbiol"},{"key":"B75","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2019.00915","article-title":"Differential responses of bovine monocyte-derived macrophages to infection by Neospora caninum isolates of high and low virulence","volume":"10","author":"Garc\u00eda-S\u00e1nchez","year":"2019","journal-title":"Front Immunol"},{"key":"B76","doi-asserted-by":"publisher","DOI":"10.1155\/2011\/282739","article-title":"Macrophage autophagy and oxidative stress: an ultrastructural and immunoelectron microscopical study","volume":"2011","author":"Perrotta","year":"2011","journal-title":"Oxid Med Cell Longev"},{"key":"B77","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2014.00538","article-title":"Cytokine secretion in macrophages: SNAREs, Rabs, and membrane trafficking","volume":"5","author":"Murray","year":"2014","journal-title":"Front Immunol"},{"key":"B78","doi-asserted-by":"publisher","DOI":"10.3390\/pathogens10020118","article-title":"The role of TLR2 and TLR4 in recognition and uptake of the apicomplexan parasite Eimeria bovis and their effects on NET formation","volume":"10","author":"Mu\u00f1oz-Caro","year":"2021","journal-title":"Pathogens"},{"key":"B79","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.179.2.1129","article-title":"Activation of TLR2 and TLR4 by glycosylphosphatidylinositols derived from Toxoplasma gondii","volume":"179","author":"Debierre-Grockiego","year":"2007","journal-title":"J Immunol Baltim Md 1950"},{"key":"B80","doi-asserted-by":"publisher","DOI":"10.1111\/cmi.12239","article-title":"Real-time imaging of Toxoplasma-infected human monocytes under fluidic shear stress reveals rapid translocation of intracellular parasites across endothelial barriers","volume":"16","author":"Ueno","year":"2014","journal-title":"Cell Microbiol"},{"key":"B81","doi-asserted-by":"publisher","DOI":"10.1126\/science.1178331","article-title":"Development of monocytes, macrophages, and dendritic cells","volume":"327","author":"Geissmann","year":"2010","journal-title":"Science"},{"key":"B82","doi-asserted-by":"publisher","first-page":"478","DOI":"10.1016\/j.chom.2019.09.008","article-title":"In vivo CRISPR screen identifies TgWIP as a Toxoplasma modulator of dendritic cell migration","volume":"26","author":"Sangar\u00e9","year":"2019","journal-title":"Cell Host Microbe"},{"key":"B83","doi-asserted-by":"publisher","DOI":"10.1155\/2016\/5898074","article-title":"Far beyond phagocytosis: phagocyte-derived extracellular traps act efficiently against protozoan parasites in vitro and in vivo","volume":"2016","author":"Silva","year":"2016","journal-title":"Mediators Inflammation"},{"key":"B84","doi-asserted-by":"publisher","DOI":"10.1016\/j.chom.2011.04.015","article-title":"Toxoplasma polymorphic effectors determine macrophage polarization and intestinal inflammation","volume":"9","author":"Jensen","year":"2011","journal-title":"Cell Host Microbe"},{"key":"B85","doi-asserted-by":"publisher","DOI":"10.1111\/pim.12712","article-title":"The role of macrophages in protective and pathological responses to Toxoplasma gondii","volume":"42","author":"Park","year":"2020","journal-title":"Parasite Immunol"},{"key":"B86","doi-asserted-by":"publisher","DOI":"10.1182\/blood-2007-07-104018","article-title":"Phagocytosis-independent antimicrobial activity of mast cells by means of extracellular trap formation","volume":"111","author":"von K\u00f6ckritz-Blickwede","year":"2008","journal-title":"Blood"},{"key":"B87","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2023.1125667","article-title":"Dynamics of cell cycle proteins involved in Toxoplasma gondii-induced bovine NET formation","volume":"14","author":"Vel\u00e1squez","year":"2023","journal-title":"Front Immunol"},{"key":"B88","doi-asserted-by":"publisher","DOI":"10.1134\/S0006297918030021","article-title":"Plasticity of human THP-1 cell phagocytic activity during macrophagic differentiation","volume":"83","author":"Kurynina","year":"2018","journal-title":"Biochem Biokhimiia"},{"key":"B89","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2019.02210","article-title":"mir-124-5p regulates phagocytosis of human macrophages by targeting the actin cytoskeleton via the ARP2\/3 complex","volume":"10","author":"Herdoiza Padilla","year":"2019","journal-title":"Front Immunol"},{"key":"B90","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0286056","article-title":"Optimization of differentiation and transcriptomic profile of THP-1 cells into macrophage by PMA","volume":"18","author":"Liu","year":"2023","journal-title":"PloS One"},{"key":"B91","doi-asserted-by":"publisher","DOI":"10.1093\/biolre\/ioad068","article-title":"Sperm induce macrophage extracellular trap formation via phagocytosis-dependent mechanism","volume":"109","author":"Lu","year":"2023","journal-title":"Biol Reprod"},{"key":"B92","doi-asserted-by":"publisher","DOI":"10.1016\/S0580-9517(10)37007-3","article-title":"Visualization and functional evaluation of phagocyte extracellular traps. San Diego (CA, USA): Methods in Microbiology","volume":"37","author":"von K\u00f6ckritz-Blickwede","year":"2010","journal-title":"Immunol Infection"},{"key":"B93","doi-asserted-by":"publisher","first-page":"19","DOI":"10.1006\/excr.1994.1003","article-title":"Different mode of cell death induced by calcium ionophore in human leukemia cell lines: possible role of constitutive endonuclease","volume":"210","author":"Matsubara","year":"1994","journal-title":"Exp Cell Res"},{"key":"B94","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2020.559561","article-title":"Trypanosoma brucei brucei induces polymorphonuclear neutrophil activation and neutrophil extracellular traps release","volume":"11","author":"Grob","year":"2020","journal-title":"Front Immunol"},{"key":"B95","doi-asserted-by":"publisher","DOI":"10.1128\/IAI.05730-11","article-title":"Toxoplasma gondii triggers release of human and mouse neutrophil extracellular traps","volume":"80","author":"Abi Abdallah","year":"2012","journal-title":"Infect Immun"},{"key":"B96","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.vetimm.2017.05.005","article-title":"Role of NETs in the difference in host susceptibility to Toxoplasma gondii between sheep and cattle","volume":"189","author":"Yildiz","year":"2017","journal-title":"Vet Immunol Immunopathol"},{"key":"B97","doi-asserted-by":"publisher","DOI":"10.1016\/j.exppara.2019.107770","article-title":"Toxoplasma gondii induces extracellular traps release in cat neutrophils","volume":"207","author":"Lacerda","year":"2019","journal-title":"Exp Parasitol"},{"key":"B98","doi-asserted-by":"publisher","DOI":"10.1016\/j.dci.2019.103527","article-title":"Dolphin-derived NETosis results in rapid Toxoplasma gondii tachyzoite ensnarement and different phenotypes of NETs","volume":"103","author":"Imlau","year":"2020","journal-title":"Dev Comp Immunol"},{"key":"B99","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2020.00429","article-title":"Toxoplasma gondii triggers neutrophil extracellular traps release in dogs","volume":"10","author":"Wei","year":"2020","journal-title":"Front Cell Infect Microbiol"},{"key":"B100","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1016\/j.toxlet.2009.08.025","article-title":"Lipopolysaccharide triggers macrophage activation of inflammatory cytokine expression, chemotaxis, phagocytosis, and oxidative ability via a Toll-like receptor 4-dependent pathway: validated by RNA interference","volume":"191","author":"Wu","year":"2009","journal-title":"Toxicol Lett"},{"key":"B101","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1501558","article-title":"The IL-12 response of primary human dendritic cells and monocytes to Toxoplasma gondii is stimulated by phagocytosis of live parasites rather than host cell invasion","volume":"196","author":"Tosh","year":"2016","journal-title":"J Immunol Baltim Md 1950"},{"key":"B102","doi-asserted-by":"publisher","DOI":"10.3347\/kjp.2008.46.3.145","article-title":"Suppressed production of pro-inflammatory cytokines by LPS-activated macrophages after treatment with Toxoplasma gondii lysate","volume":"46","author":"Lee","year":"2008","journal-title":"Korean J Parasitol"},{"key":"B103","doi-asserted-by":"publisher","DOI":"10.1189\/jlb.4A1116-496R","article-title":"Phenotypic and functional consequences of different isolation protocols on skin mononuclear phagocytes","volume":"101","author":"Botting","year":"2017","journal-title":"J Leukoc Biol"},{"key":"B104","doi-asserted-by":"publisher","DOI":"10.1016\/j.tibs.2016.09.002","article-title":"Mechanism and regulation of NLRP3 inflammasome activation","volume":"41","author":"He","year":"2016","journal-title":"Trends Biochem Sci"},{"key":"B105","doi-asserted-by":"publisher","first-page":"6662","DOI":"10.1038\/s41467-024-50771-6","article-title":"High-affinity agonism at the P2X7 receptor is mediated by three residues outside the orthosteric pocket","volume":"15","author":"Oken","year":"2024","journal-title":"Nat Commun"},{"key":"B106","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2013.05.016","article-title":"K+ efflux is the common trigger of NLRP3 inflammasome activation by bacterial toxins and particulate matter","volume":"38","author":"Mu\u00f1oz-Planillo","year":"2013","journal-title":"Immunity"}],"container-title":["Frontiers in Immunology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2025.1683634\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,30]],"date-time":"2025-10-30T06:22:24Z","timestamp":1761805344000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2025.1683634\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,10,30]]},"references-count":106,"alternative-id":["10.3389\/fimmu.2025.1683634"],"URL":"https:\/\/doi.org\/10.3389\/fimmu.2025.1683634","relation":{},"ISSN":["1664-3224"],"issn-type":[{"value":"1664-3224","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,10,30]]},"article-number":"1683634"}}