{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,9]],"date-time":"2026-04-09T09:56:58Z","timestamp":1775728618747,"version":"3.50.1"},"reference-count":50,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,3,18]],"date-time":"2026-03-18T00:00:00Z","timestamp":1773792000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,4,9]],"date-time":"2026-04-09T00:00:00Z","timestamp":1775692800000},"content-version":"vor","delay-in-days":22,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100008977","name":"Universit\u00e4t Ulm","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100008977","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Parasites Vectors"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:sec>\n                    <jats:title>Background<\/jats:title>\n                    <jats:p>\n                      Cystic echinococcosis (CE) and alveolar echinococcosis (AE) are zoonotic diseases caused by the larval stages of\n                      <jats:italic>Echinococcus granulosus<\/jats:italic>\n                      and\n                      <jats:italic>E. multilocularis<\/jats:italic>\n                      . Prior studies have identified small particles from the outer layer of parasitic lesions termed spegs (from\n                      <jats:italic>E. granulosus<\/jats:italic>\n                      ) and spems (from\n                      <jats:italic>E. multilocularis<\/jats:italic>\n                      ), which appear to interact with the human immune system, leading to lymph node enlargement.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>\n                      We analyzed lymph nodes from CE (\n                      <jats:italic>n<\/jats:italic>\n                      \u2009=\u20093) and AE (\n                      <jats:italic>n<\/jats:italic>\n                      \u2009=\u20093) patients by immunohistochemical staining for spegs\/spems and CD57. Automated image analysis revealed a significantly increased number of CD57-positive cells in\n                      <jats:italic>Echinococcus<\/jats:italic>\n                      particle-positive germinal centers compared with speg-\/spem-negative germinal centers.\n                    <\/jats:p>\n                    <jats:p>Double immunofluorescence staining was used to characterize CD57-positive cells with markers for CD2, CD3, CD4, CD8, PD-1, Granzyme B, Perforin, and T-cell intracellular antigen 1 (TIA1). Additionally, IgG4 and IgE immunostaining was performed on liver lesions and lymph nodes containing spegs and spems and compared with unaffected controls.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>Spegs and spems were found to accumulate in the germinal centers of the lymph nodes. Affected germinal centers showed a higher number of CD57-positive cells, which coexpressed CD2, CD3, CD4, and PD-1 but lacked cytotoxic markers (CD8, Granzyme B, Perforin, TIA1), identifying them as follicular helper T cells (Tfh). Furthermore, the speg- and spem-affected liver and lymph node sections exhibited significantly increased numbers of IgG4-, IgE-, and CD57-positive cells.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusions<\/jats:title>\n                    <jats:p>Spegs and spems are associated with changes in the host immune response, characterized by an increased accumulation of Tfh in germinal centers. This finding is accompanied by the presence of IgE and IgG4 plasma cells in the lymph node and the perilesional area, suggesting parasite-associated immunomodulatory processes in echinococcosis.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Graphical Abstract<\/jats:title>\n                  <\/jats:sec>","DOI":"10.1186\/s13071-026-07321-4","type":"journal-article","created":{"date-parts":[[2026,3,18]],"date-time":"2026-03-18T12:45:23Z","timestamp":1773837923000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Small particles of Echinococcus granulosus (spegs) and Echinococcus multilocularis (spems) promote follicular T helper cell expansion and are associated with IgE and IgG4 class switching in human lymph nodes"],"prefix":"10.1186","volume":"19","author":[{"given":"Leonard","family":"Schreiber","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Linyun","family":"Zhang","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Johannes","family":"Grimm","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Kevin","family":"Mellert","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Juliane","family":"Nell","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Peter","family":"M\u00f6ller","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Thomas F. E.","family":"Barth","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2026,3,18]]},"reference":[{"key":"7321_CR1","unstructured":"Thompson A. Echinococcus and Echinococcosis, Part B: Echinococcus and Echinococcosis, Part B. Issn Ser, v.Volume 96. Saint Louis: Elsevier Science; 2017. 422 p."},{"key":"7321_CR2","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1128\/CMR.17.1.107-135.2004","volume":"17","author":"J Eckert","year":"2004","unstructured":"Eckert J, Deplazes P. Biological, epidemiological, and clinical aspects of echinococcosis, a zoonosis of increasing concern. Clin Microbiol Rev. 2004;17:107\u201335.","journal-title":"Clin Microbiol Rev"},{"key":"7321_CR3","doi-asserted-by":"publisher","first-page":"264","DOI":"10.1016\/j.pt.2011.01.008","volume":"27","author":"A D\u00edaz","year":"2011","unstructured":"D\u00edaz A, Casaravilla C, Allen JE, Sim RB, Ferreira AM. Understanding the laminated layer of larval Echinococcus II: immunology. Trends Parasitol. 2011;27:264\u201373.","journal-title":"Trends Parasitol"},{"key":"7321_CR4","first-page":"265","volume-title":"WHO\/OIE manual on echinococcosis in humans and animals: A zoonosis of global concern","author":"J Eckert","year":"2001","unstructured":"Eckert J, Gemmell MA. WHO\/OIE manual on echinococcosis in humans and animals: A zoonosis of global concern. Paris: World Organisation for Animal Health; 2001. p. 265."},{"key":"7321_CR5","doi-asserted-by":"publisher","DOI":"10.1128\/iai.00031-23","volume":"91","author":"AA Barrios","year":"2023","unstructured":"Barrios AA, Mouhape C, Schreiber L, Zhang L, Nell J, Su\u00e1rez-Martins M, et al. Mucins shed from the laminated layer in cystic echinococcosis are captured by Kupffer cells via the lectin receptor Clec4F. Infect Immun. 2023;91:e0003123.","journal-title":"Infect Immun"},{"key":"7321_CR6","doi-asserted-by":"publisher","first-page":"39204","DOI":"10.1038\/srep39204","volume":"6","author":"PI Seoane","year":"2016","unstructured":"Seoane PI, R\u00fcckerl D, Casaravilla C, Barrios AA, Pittini \u00c1, MacDonald AS, et al. Particles from the Echinococcus granulosus laminated layer inhibit IL-4 and growth factor-driven Akt phosphorylation and proliferative responses in macrophages. Sci Rep. 2016;6:39204.","journal-title":"Sci Rep"},{"key":"7321_CR7","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pntd.0001877","volume":"6","author":"TFE Barth","year":"2012","unstructured":"Barth TFE, Herrmann TS, Tappe D, Stark L, Gr\u00fcner B, Buttenschoen K, et al. Sensitive and specific immunohistochemical diagnosis of human alveolar echinococcosis with the monoclonal antibody Em2G11. PLoS Negl Trop Dis. 2012;6:e1877.","journal-title":"PLoS Negl Trop Dis"},{"key":"7321_CR8","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1371\/journal.pntd.0008921","volume":"14","author":"J Grimm","year":"2020","unstructured":"Grimm J, Nell J, Hillenbrand A, Henne-Bruns D, Schmidberger J, Kratzer W, et al. Immunohistological detection of small particles of Echinococcus multilocularis and Echinococcus granulosus in lymph nodes is associated with enlarged lymph nodes in alveolar and cystic echinococcosis. PLoS Negl Trop Dis. 2020;14:1\u201316.","journal-title":"PLoS Negl Trop Dis"},{"key":"7321_CR9","first-page":"6449","volume":"365","author":"Rick M Maizels","year":"1993","unstructured":"Maizels Rick M, Bundy Don A. P, Selkirk Murray E, Smith Deborah F, Anderson Roy M. Immunological modulation and evasion by helminth parasites in human populations. Nature Publishing Group. 1993;365:6449.","journal-title":"Nature Publishing Group"},{"key":"7321_CR10","doi-asserted-by":"publisher","first-page":"225","DOI":"10.1016\/0014-4894(74)90026-5","volume":"35","author":"EA Coltorii","year":"1974","unstructured":"Coltorii EA, Varela-D\u00edaz VM. Echinococcus granulosus: penetration of macromolecules and their localization on the parasite membranes of cysts. Exp Parasitol. 1974;35:225\u201331.","journal-title":"Exp Parasitol"},{"key":"7321_CR11","doi-asserted-by":"publisher","first-page":"1162530","DOI":"10.3389\/fcimb.2023.1162530","volume":"13","author":"PA Kronenberg","year":"2023","unstructured":"Kronenberg PA, Reinehr M, Eichenberger RM, Hasler S, Laurim\u00e4e T, Weber A, et al. Monoclonal antibody-based localization of major diagnostic antigens in metacestode tissue, excretory\/secretory products, and extracellular vesicles of Echinococcus species. Front Cell Infect Microbiol. 2023;13:1162530.","journal-title":"Front Cell Infect Microbiol"},{"key":"7321_CR12","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pntd.0009155","volume":"15","author":"J Grimm","year":"2021","unstructured":"Grimm J, Krickl J, Beck A, Nell J, Bergmann M, Tappe D, et al. Establishing and evaluation of a polymerase chain reaction for the detection of Echinococcus multilocularis in human tissue. PLoS Negl Trop Dis. 2021;15:e0009155.","journal-title":"PLoS Negl Trop Dis"},{"key":"7321_CR13","doi-asserted-by":"publisher","first-page":"422","DOI":"10.3389\/fimmu.2013.00422","volume":"4","author":"CM Nielsen","year":"2013","unstructured":"Nielsen CM, White MJ, Goodier MR, Riley EM. Functional significance of CD57 expression on human NK cells and relevance to disease. Front Immunol. 2013;4:422.","journal-title":"Front Immunol"},{"key":"7321_CR14","doi-asserted-by":"publisher","first-page":"441","DOI":"10.1007\/s00262-016-1803-z","volume":"65","author":"H Kared","year":"2016","unstructured":"Kared H, Martelli S, Ng TP, Pender SLF, Larbi A. CD57 in human natural killer cells and T-lymphocytes. Cancer Immunol Immunother. 2016;65:441\u201352.","journal-title":"Cancer Immunol Immunother"},{"key":"7321_CR15","doi-asserted-by":"crossref","unstructured":"Frimpong A, Kusi KA, Adu-Gyasi D, Amponsah J, Ofori MF, Ndifon W. Phenotypic Evidence of T Cell Exhaustion and Senescence During Symptomatic Plasmodium falciparum Malaria 2019 Front Immunol 10:1345","DOI":"10.3389\/fimmu.2019.01345"},{"key":"7321_CR16","doi-asserted-by":"publisher","first-page":"2711","DOI":"10.1182\/blood-2002-07-2103","volume":"101","author":"JM Brenchley","year":"2003","unstructured":"Brenchley JM, Karandikar NJ, Betts MR, Ambrozak DR, Hill BJ, Crotty LE, et al. Expression of CD57 defines replicative senescence and antigen-induced apoptotic death of CD8+ T cells. Blood. 2003;101:2711\u201320.","journal-title":"Blood"},{"key":"7321_CR17","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1038\/ni.1679","volume":"10","author":"SD Blackburn","year":"2009","unstructured":"Blackburn SD, Shin H, Haining WN, Zou T, Workman CJ, Polley A, et al. Coregulation of CD8+ T cell exhaustion by multiple inhibitory receptors during chronic viral infection. Nat Immunol. 2009;10:29\u201337.","journal-title":"Nat Immunol"},{"key":"7321_CR18","doi-asserted-by":"publisher","first-page":"173439","DOI":"10.1155\/2009\/173439","volume":"2009","author":"R Garc\u00eda-Mu\u00f1oz","year":"2009","unstructured":"Garc\u00eda-Mu\u00f1oz R, Rodr\u00edguez-Otero P, Galar A, Merino J, Beunza JJ, P\u00e1ramo JA, et al. Expansion of CD8+CD57+ T Cells in an Immunocompetent Patient with Acute Toxoplasmosis. Adv Hematol. 2009;2009:173439.","journal-title":"Adv Hematol"},{"key":"7321_CR19","doi-asserted-by":"publisher","DOI":"10.1016\/j.intimp.2022.109596","volume":"114","author":"Z Alizadeh","year":"2023","unstructured":"Alizadeh Z, Omidnia P, Altalbawy FMA, Gabr GA, Obaid RF, Rostami N, et al. Unraveling the role of natural killer cells in leishmaniasis. Int Immunopharmacol. 2023;114:109596.","journal-title":"Int Immunopharmacol"},{"key":"7321_CR20","doi-asserted-by":"publisher","first-page":"1272084","DOI":"10.3389\/fimmu.2024.1272084","volume":"15","author":"RV Motta","year":"2024","unstructured":"Motta RV, Culver EL. IgG4 autoantibodies and autoantigens in the context of IgG4-autoimmune disease and IgG4-related disease. Front Immunol. 2024;15:1272084.","journal-title":"Front Immunol"},{"key":"7321_CR21","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1097\/PAS.0000000000001374","volume":"44","author":"M Reinehr","year":"2020","unstructured":"Reinehr M, Micheloud C, Grimm F, Kronenberg PA, Grimm J, Beck A, et al. Pathology of Echinococcosis: A Morphologic and Immunohistochemical Study on 138 Specimens With Focus on the Differential Diagnosis Between Cystic and Alveolar Echinococcosis. Am J Surg Pathol. 2020;44:43\u201354.","journal-title":"Am J Surg Pathol"},{"key":"7321_CR22","doi-asserted-by":"publisher","first-page":"433","DOI":"10.1186\/s12859-021-04344-9","volume":"22","author":"DR Stirling","year":"2021","unstructured":"Stirling DR, Swain-Bowden MJ, Lucas AM, Carpenter AE, Cimini BA, Goodman A. Cell Profiler 4: improvements in speed, utility and usability. BMC Bioinformatics. 2021;22:433.","journal-title":"BMC Bioinformatics"},{"key":"7321_CR23","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1186\/s40164-015-0022-1","volume":"4","author":"A Sattarzadeh","year":"2015","unstructured":"Sattarzadeh A, Diepstra A, Rutgers B, van den Berg A, Visser L. CD57+ T-cells are a subpopulation of T-follicular helper cells in nodular lymphocyte predominant Hodgkin lymphoma. Exp Hematol Oncol. 2015;4:27.","journal-title":"Exp Hematol Oncol"},{"key":"7321_CR24","doi-asserted-by":"publisher","first-page":"124","DOI":"10.1038\/s41408-023-00899-3","volume":"13","author":"Z-Z Yang","year":"2023","unstructured":"Yang Z-Z, Kim HJ, Wu H, Tang X, Yu Y, Krull J, et al. T-cell phenotype including CD57+ T follicular helper cells in the tumor microenvironment correlate with a poor outcome in follicular lymphoma. Blood Cancer J. 2023;13:124.","journal-title":"Blood Cancer J"},{"key":"7321_CR25","doi-asserted-by":"publisher","first-page":"102421","DOI":"10.1016\/j.coi.2024.102421","volume":"87","author":"SA Wellford","year":"2024","unstructured":"Wellford SA, Schwartzberg PL. Help me help you: emerging concepts in T follicular helper cell differentiation, identity, and function. Curr Opin Immunol. 2024;87:102421.","journal-title":"Curr Opin Immunol"},{"key":"7321_CR26","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1186\/s13071-016-1350-7","volume":"9","author":"V Silva-\u00c1lvarez","year":"2016","unstructured":"Silva-\u00c1lvarez V, Folle AM, Ramos AL, Kitano ES, Iwai LK, Corraliza I, et al. Echinococcus granulosus antigen B binds to monocytes and macrophages modulating cell response to inflammation. Parasit Vectors. 2016;9:69.","journal-title":"Parasit Vectors"},{"key":"7321_CR27","first-page":"2","volume":"14","author":"\u00c1 D\u00edaz","year":"2022","unstructured":"D\u00edaz \u00c1, Barrios AA, Grezzi L, Mouhape C, Jenkins SJ, Allen JE, et al. Immunology of a unique biological structure: the Echinococcus laminated layer. Protein & Cell. 2022;14:2.","journal-title":"Protein & Cell"},{"key":"7321_CR28","doi-asserted-by":"publisher","first-page":"272","DOI":"10.1016\/j.immuni.2021.12.015","volume":"55","author":"C-H Yeh","year":"2022","unstructured":"Yeh C-H, Finney J, Okada T, Kurosaki T, Kelsoe G. Primary germinal center-resident T follicular helper cells are a physiologically distinct subset of CXCR5hiPD-1hi T follicular helper cells. Immunity. 2022;55:272-289.e7.","journal-title":"Immunity"},{"key":"7321_CR29","doi-asserted-by":"crossref","unstructured":"Crotty S 2011 Follicular helper CD4 T cells [TFH] 29 1 621 663","DOI":"10.1146\/annurev-immunol-031210-101400"},{"key":"7321_CR30","doi-asserted-by":"publisher","first-page":"1178792","DOI":"10.3389\/fimmu.2023.1178792","volume":"14","author":"J Qi","year":"2023","unstructured":"Qi J, Liu C, Bai Z, Li X, Yao G. T follicular helper cells and T follicular regulatory cells in autoimmune diseases. Front Immunol. 2023;14:1178792.","journal-title":"Front Immunol"},{"key":"7321_CR31","doi-asserted-by":"publisher","first-page":"335","DOI":"10.1146\/annurev-immunol-041015-055605","volume":"34","author":"CG Vinuesa","year":"2016","unstructured":"Vinuesa CG, Linterman MA, Di Yu, MacLennan ICM. Follicular Helper T Cells. Annu Rev Immunol. 2016;34:335\u201368.","journal-title":"Annu Rev Immunol"},{"key":"7321_CR32","doi-asserted-by":"publisher","first-page":"58","DOI":"10.1038\/ni.2182","volume":"13","author":"H-E Liang","year":"2011","unstructured":"Liang H-E, Reinhardt RL, Bando JK, Sullivan BM, Ho I-C, Locksley RM. Divergent expression patterns of IL-4 and IL-13 define unique functions in allergic immunity. Nat Immunol. 2011;13:58\u201366.","journal-title":"Nat Immunol"},{"key":"7321_CR33","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.jaci.2016.04.021","volume":"139","author":"T Kobayashi","year":"2017","unstructured":"Kobayashi T, Iijima K, Dent AL, Kita H. Follicular helper T cells mediate IgE antibody response to airborne allergens. J Allergy Clin Immunol. 2017;139:1.","journal-title":"J Allergy Clin Immunol"},{"key":"7321_CR34","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1093\/oxfimm\/iqab008","volume":"2","author":"MSF Soon","year":"2021","unstructured":"Soon MSF, Nalubega M, Boyle MJ. T-follicular helper cells in malaria infection and roles in antibody induction. Oxf Open Immunol. 2021;2:1.","journal-title":"Oxf Open Immunol"},{"key":"7321_CR35","first-page":"247","volume":"71","author":"DA Vuitton","year":"1998","unstructured":"Vuitton DA, BRESSON-HADNI S, Lenys D, Flausse F, Liance M, Wattre P, et al. IgE-dependent humoral immune response in Echinococcus multilocularis infection: circulating and basophil-bound specific IgE against Echinococcus antigens in patients with alveolar echinococcosis. Clin Exp Immunol. 1998;71:247\u201352.","journal-title":"Clin Exp Immunol"},{"key":"7321_CR36","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pntd.0005636","volume":"11","author":"FJ Ricken","year":"2017","unstructured":"Ricken FJ, Nell J, Gr\u00fcner B, Schmidberger J, Kaltenbach T, Kratzer W, et al. Albendazole increases the inflammatory response and the amount of Em2-positive small particles of Echinococcus multilocularis [spems] in human hepatic alveolar echinococcosis lesions. PLoS Negl Trop Dis. 2017;11:e0005636.","journal-title":"PLoS Negl Trop Dis"},{"key":"7321_CR37","doi-asserted-by":"publisher","first-page":"3164","DOI":"10.1128\/IAI.01959-14","volume":"82","author":"C Casaravilla","year":"2014","unstructured":"Casaravilla C, Pittini A, R\u00fcckerl D, Seoane PI, Jenkins SJ, MacDonald AS, et al. Unconventional maturation of dendritic cells induced by particles from the laminated layer of larval Echinococcus granulosus. Infect Immun. 2014;82:3164\u201376.","journal-title":"Infect Immun"},{"key":"7321_CR38","doi-asserted-by":"publisher","first-page":"1334211","DOI":"10.3389\/fcimb.2024.1334211","volume":"14","author":"M Chop","year":"2024","unstructured":"Chop M, Ledo C, Nicolao MC, Loos J, Cumino A, Rodriguez Rodrigues C. Hydatid fluid from Echinococcus granulosus induces autophagy in dendritic cells and promotes polyfunctional T-cell responses. Front Cell Infect Microbiol. 2024;14:1334211.","journal-title":"Front Cell Infect Microbiol"},{"key":"7321_CR39","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pntd.0010721","volume":"16","author":"G Bewket","year":"2022","unstructured":"Bewket G, Kiflie A, Tajebe F, Abate E, Sch\u00f6n T, Blomgran R. Helminth species dependent effects on Th1 and Th17 cytokines in active tuberculosis patients and healthy community controls. PLoS Negl Trop Dis. 2022;16:e0010721.","journal-title":"PLoS Negl Trop Dis"},{"key":"7321_CR40","doi-asserted-by":"crossref","unstructured":"Thompson R, Lymbery AJ, Constantine CC 2017 Variation in Echinococcus: Towards a Taxonomic Revision of the Genus. In: Thompson A, Deplazes P, Lymbery AJ, editors. Echinococcus and Echinococcosis, Part B. Issn Ser. v.Volume 96. Saint Louis: Elsevier Science 145\u201375.","DOI":"10.1016\/S0065-308X(08)60071-8"},{"key":"7321_CR41","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pntd.0001516","volume":"6","author":"JK Nono","year":"2012","unstructured":"Nono JK, Pletinckx K, Lutz MB, Brehm K. Excretory\/secretory-products of Echinococcus multilocularis larvae induce apoptosis and tolerogenic properties in dendritic cells in vitro. PLoS Negl Trop Dis. 2012;6:e1516.","journal-title":"PLoS Negl Trop Dis"},{"key":"7321_CR42","doi-asserted-by":"publisher","first-page":"657","DOI":"10.1007\/s00436-010-1912-0","volume":"107","author":"NW Brattig","year":"2010","unstructured":"Brattig NW, Tenner-Racz K, Korten S, Hoerauf A, B\u00fcttner DW. Immunohistology of ectopic secondary lymph follicles in subcutaneous nodules from patients with hyperreactive onchocerciasis [sowda]. Parasitol Res. 2010;107:657\u201366.","journal-title":"Parasitol Res"},{"key":"7321_CR43","doi-asserted-by":"publisher","DOI":"10.1002\/iid3.1329","volume":"12","author":"S Mahmoudzadeh","year":"2024","unstructured":"Mahmoudzadeh S, Dizaji Asl K, Nozad Charoudeh H, Rahbarghazi R, Ahmadi M, Heidarzadeh M, et al. Toxoplasma gondii suppress human cord blood cell differentiation to the NK cell population. Immun Inflamm Dis. 2024;12:e1329.","journal-title":"Immun Inflamm Dis"},{"key":"7321_CR44","doi-asserted-by":"publisher","first-page":"1341","DOI":"10.1086\/586709","volume":"197","author":"S Zhang","year":"2008","unstructured":"Zhang S, H\u00fce S, S\u00e8ne D, Penfornis A, Bresson-Hadni S, Kantelip B, et al. Expression of major histocompatibility complex class I chain-related molecule A, NKG2D, and transforming growth factor-beta in the liver of humans with alveolar echinococcosis: new actors in the tolerance to parasites? J Infect Dis. 2008;197:1341\u20139.","journal-title":"J Infect Dis"},{"key":"7321_CR45","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1007\/s12016-004-0004-2","volume":"26","author":"DA Vuitton","year":"2004","unstructured":"Vuitton DA. Echinococcosis and allergy. Clin Rev Allergy Immunol. 2004;26:93\u2013104.","journal-title":"Clin Rev Allergy Immunol"},{"key":"7321_CR46","first-page":"61","volume":"5","author":"CM Fitzsimmons","year":"2014","unstructured":"Fitzsimmons CM, Falcone FH, Dunne DW. Helminth Allergens. Parasite-specific IgE and its pro role in human. Immun Front Immunol. 2014;5:61.","journal-title":"Parasite-specific IgE and its pro role in human. Immun Front Immunol"},{"key":"7321_CR47","doi-asserted-by":"publisher","first-page":"585","DOI":"10.1128\/CMR.05040-11","volume":"25","author":"HJ McSorley","year":"2012","unstructured":"McSorley HJ, Maizels RM. Helminth infections and host immune regulation. Clin Microbiol Rev. 2012;25:585\u2013608.","journal-title":"Clin Microbiol Rev"},{"key":"7321_CR48","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1016\/S0001-706X(99)00027-3","volume":"73","author":"N Wellinghausen","year":"1999","unstructured":"Wellinghausen N, Gebert P, Kern P. Interleukin [IL]-4, IL-10 and IL-12 profile in serum of patients with alveolar echinococcosis. Acta Trop. 1999;73:165\u201374.","journal-title":"Acta Trop"},{"key":"7321_CR49","unstructured":"Mejri N, Hemphill A, Gottstein B Triggering and modulation of the host-parasite interplay by Echinococcus multilocularis: a review 2024."},{"key":"7321_CR50","unstructured":"Zentrale Ethikkommission bei der Bundes\u00e4rztekammer. Die [Weiter-]Verwendung von menschlichen K\u00f6rpermaterialien f\u00fcr Zwecke medizinischer Forschung [20.02.2003]."}],"container-title":["Parasites &amp; Vectors"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s13071-026-07321-4","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s13071-026-07321-4.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s13071-026-07321-4.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,4,9]],"date-time":"2026-04-09T09:02:48Z","timestamp":1775725368000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1186\/s13071-026-07321-4"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,3,18]]},"references-count":50,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["7321"],"URL":"https:\/\/doi.org\/10.1186\/s13071-026-07321-4","relation":{},"ISSN":["1756-3305"],"issn-type":[{"value":"1756-3305","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,3,18]]},"assertion":[{"value":"8 December 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"16 February 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"18 March 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"Paraffin blocks of tissue samples used in this study were drawn from the archives of the Institute of Pathology, University of Ulm and dated back to 1989. In compliance with the German law for correct usage of archival tissue for clinical research [\n                      \n                      ], the used blocks were anonymized. This study was approved by the Ethics Committee of the University Ulm (ref. no. 440\/15 and 116\/13) and was conducted in accordance with the Declaration of Helsinki.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval and consent to participate"}},{"value":"All tissue samples included in this study were fully anonymized prior to analysis, and no identifiable personal data were retained. In accordance with ethical guidelines, informed consent for the use of tissue and clinical information for research purposes was obtained from all tissue donors (or their legal representatives). Consequently, the study complies with all requirements regarding the use of individual patient material in scientific publications.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"The authors declare no competing interests.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"151"}}