{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,11]],"date-time":"2026-06-11T09:00:01Z","timestamp":1781168401146,"version":"3.54.1"},"reference-count":57,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,5,28]],"date-time":"2026-05-28T00:00:00Z","timestamp":1779926400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,6,11]],"date-time":"2026-06-11T00:00:00Z","timestamp":1781136000000},"content-version":"vor","delay-in-days":14,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100006196","name":"University of Oulu","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100006196","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["BMC Med"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:sec>\n                    <jats:title>Background<\/jats:title>\n                    <jats:p>While most clinical microbiome research has investigated gut microbiota, less is known about extracellular vesicles\u00a0(EVs) produced by microbiota. Microbiota-derived EVs form a distinct taxonomic entity from the gut microbiota. Recently, microbiota-derived EVs from maternal microbiota have been shown to reach the fetus, which could be an important mechanism of microbiota\u2013host interaction during the fetal period. We hypothesized that maternal factors could change the composition of microbiota-derived EVs during pregnancy.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>We compared the influence of antibiotics and maternal weight on microbiota-derived EVs secreted by maternal microbiota during pregnancy. We collected fecal samples from 60 pregnant women and 18 amniotic fluid samples from those undergoing C-section. Microbiota-derived EVs were isolated from the samples using size-exclusion chromatography and density-gradient ultracentrifugation and characterized using transmission electron microscopy (TEM) and nanoparticle tracking analysis (NTA). RNA was isolated from the microbiota-derived EVs and converted to complementary DNA (cDNA), and 16\u00a0S rRNA gene sequencing was performed.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>Altogether, 18 (30%) women had received antibiotics during pregnancy and 37 (62%) during delivery. Twenty (33%) women were lean, 27 (45%) had overweight and 13 had obesity (22%) during the first trimester. Taxonomic diversity of microbiota-secreted microbiota-derived EVs was lower in women exposed to antibiotics during pregnancy. When women exposed to antibiotics during delivery were excluded, the differences were not statistically significant. The microbiota-derived EVs differed in the amniotic fluid and maternal feces in women with overweight and obesity, gestational diabetes (GDM), and those who gained\u2009&gt;\u200915\u00a0kg during pregnancy, as many bacterial origins of microbiota-derived EVs were depleted in these women. A beta diversity analysis of microbiota-derived EVs from fecal samples showed significant differences between the overweight and obesity groups. Diversity analysis showed no differences in various health factors during pregnancy, including asthma, allergies, or smoking.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusions<\/jats:title>\n                    <jats:p>Maternal factors influence the composition of microbiota-derived extracellular vesicles in feces and amniotic fluid of pregnant women, which may change host-microbiota interaction in the fetal period.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.1186\/s12916-026-04960-3","type":"journal-article","created":{"date-parts":[[2026,5,28]],"date-time":"2026-05-28T14:56:30Z","timestamp":1779980190000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Maternal factors associate with microbiota-derived extracellular vesicle profiles in pregnancy: a clinical cohort study"],"prefix":"10.1186","volume":"24","author":[{"given":"Jenni","family":"Turunen","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kaisu","family":"Kyr\u00f6","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Janica","family":"Virta","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marko","family":"Suokas","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jenni","family":"Hekkala","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sonja","family":"Karikka","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anna","family":"Kaisanlahti","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Justus","family":"Reunanen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Niko","family":"Paalanne","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mysore V.","family":"Tejesvi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Terhi","family":"Ruuska-Loewald","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,5,28]]},"reference":[{"key":"4960_CR1","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1038\/s41579-018-0112-2","volume":"17","author":"M Toyofuku","year":"2019","unstructured":"Toyofuku M, Nomura N, Eberl L. Types and origins of bacterial membrane vesicles. Nat Rev Microbiol. 2019;17:13\u201324. https:\/\/doi.org\/10.1038\/s41579-018-0112-2.","journal-title":"Nat Rev Microbiol"},{"key":"4960_CR2","doi-asserted-by":"publisher","first-page":"375","DOI":"10.1038\/nri3837","volume":"15","author":"M Kaparakis-Liaskos","year":"2015","unstructured":"Kaparakis-Liaskos M, Ferrero RL. Immune modulation by bacterial outer membrane vesicles. Nat Rev Immunol. 2015;15:375\u201387. https:\/\/doi.org\/10.1038\/nri3837.","journal-title":"Nat Rev Immunol"},{"key":"4960_CR3","doi-asserted-by":"publisher","DOI":"10.1128\/msystems.00311-25","author":"A Kaisanlahti","year":"2025","unstructured":"Kaisanlahti A, Turunen J, Hekkala J, Mishra S, Karikka S, Amatya SB, et al. Gut microbiota-derived extracellular vesicles form a distinct entity from gut microbiota. mSystems. 2025. https:\/\/doi.org\/10.1128\/msystems.00311-25. e00311-25.","journal-title":"mSystems"},{"key":"4960_CR4","doi-asserted-by":"publisher","first-page":"27066","DOI":"10.3402\/jev.v4.27066","volume":"4","author":"M Y\u00e1\u00f1ez-M\u00f3","year":"2015","unstructured":"Y\u00e1\u00f1ez-M\u00f3 M, Siljander PR, \u2010M., Andreu Z, Bedina Zavec A, Borr\u00e0s FE, Buzas EI, et al. Biological properties of extracellular vesicles and their physiological functions. J Extracell Vesicles. 2015;4:27066. https:\/\/doi.org\/10.3402\/jev.v4.27066.","journal-title":"J Extracell Vesicles"},{"key":"4960_CR5","doi-asserted-by":"publisher","first-page":"8899","DOI":"10.3390\/ijms21238899","volume":"21","author":"S Haas-Neill","year":"2020","unstructured":"Haas-Neill S, Forsythe PA, Budding Relationship. Bacterial Extracellular Vesicles in the Microbiota-Gut-Brain Axis. Int J Mol Sci. 2020;21:8899. https:\/\/doi.org\/10.3390\/ijms21238899.","journal-title":"Int J Mol Sci"},{"key":"4960_CR6","doi-asserted-by":"publisher","first-page":"13787","DOI":"10.3390\/ijms232213787","volume":"23","author":"B Schaack","year":"2022","unstructured":"Schaack B, Hindr\u00e9 T, Quansah N, Hannani D, Mercier C, Laurin D. Microbiota-Derived Extracellular Vesicles Detected in Human Blood from Healthy Donors. Int J Mol Sci. 2022;23:13787. https:\/\/doi.org\/10.3390\/ijms232213787.","journal-title":"Int J Mol Sci"},{"key":"4960_CR7","doi-asserted-by":"publisher","first-page":"249","DOI":"10.1186\/s40168-023-01694-9","volume":"11","author":"A Kaisanlahti","year":"2023","unstructured":"Kaisanlahti A, Turunen J, Byts N, Samoylenko A, Bart G, Virtanen N, et al. Maternal microbiota communicates with the fetus through microbiota-derived extracellular vesicles. Microbiome. 2023;11:249. https:\/\/doi.org\/10.1186\/s40168-023-01694-9.","journal-title":"Microbiome"},{"key":"4960_CR8","doi-asserted-by":"publisher","first-page":"665","DOI":"10.1038\/ijo.2014.180","volume":"39","author":"NT Mueller","year":"2015","unstructured":"Mueller NT, Whyatt R, Hoepner L, Oberfield S, Dominguez-Bello MG, Widen EM, et al. Prenatal exposure to antibiotics, cesarean section and risk of childhood obesity. Int J Obes. 2015;39:665\u201370. https:\/\/doi.org\/10.1038\/ijo.2014.180.","journal-title":"Int J Obes"},{"key":"4960_CR9","doi-asserted-by":"publisher","first-page":"18","DOI":"10.1186\/s12941-019-0318-9","volume":"18","author":"M Zhang","year":"2019","unstructured":"Zhang M, Differding MK, Benjamin-Neelon SE, \u00d8stbye T, Hoyo C, Mueller NT. Association of prenatal antibiotics with measures of infant adiposity and the gut microbiome. Ann Clin Microbiol Antimicrob. 2019;18:18. https:\/\/doi.org\/10.1186\/s12941-019-0318-9.","journal-title":"Ann Clin Microbiol Antimicrob"},{"key":"4960_CR10","doi-asserted-by":"publisher","first-page":"328","DOI":"10.1016\/j.intimp.2018.10.021","volume":"65","author":"A Strz\u0119pa","year":"2018","unstructured":"Strz\u0119pa A, Lobo FM, Majewska-Szczepanik M, Szczepanik M. Antibiotics and autoimmune and allergy diseases: Causative factor or treatment? Int Immunopharmacol. 2018;65:328\u201341. https:\/\/doi.org\/10.1016\/j.intimp.2018.10.021.","journal-title":"Int Immunopharmacol"},{"key":"4960_CR11","doi-asserted-by":"publisher","unstructured":"Catalano PM, Shankar K. Obesity and pregnancy: mechanisms of short term and long term adverse consequences for mother and child. BMJ. 2017;356. https:\/\/doi.org\/10.1136\/bmj.j1.","DOI":"10.1136\/bmj.j1"},{"key":"4960_CR12","doi-asserted-by":"publisher","first-page":"361","DOI":"10.1146\/annurev-med-012510-175505","volume":"62","author":"G Musso","year":"2011","unstructured":"Musso G, Gambino R, Cassader M. Interactions Between Gut Microbiota and Host Metabolism Predisposing to Obesity and Diabetes. Annu Rev Med. 2011;62:361\u201380. https:\/\/doi.org\/10.1146\/annurev-med-012510-175505.","journal-title":"Annu Rev Med"},{"key":"4960_CR13","doi-asserted-by":"publisher","first-page":"40","DOI":"10.1038\/s41596-019-0236-5","volume":"15","author":"J Tulkens","year":"2020","unstructured":"Tulkens J, De Wever O, Hendrix A. Analyzing bacterial extracellular vesicles in human body fluids by orthogonal biophysical separation and biochemical characterization. Nat Protoc. 2020;15:40\u201367. https:\/\/doi.org\/10.1038\/s41596-019-0236-5.","journal-title":"Nat Protoc"},{"key":"4960_CR14","doi-asserted-by":"publisher","first-page":"228","DOI":"10.1038\/nmeth.4185","volume":"14","author":"EV-TRACK Consortium","year":"2017","unstructured":"Consortium EV-TRACK, Van Deun J, Mestdagh P, Agostinis P, Akay \u00d6, Anand S, et al. EV-TRACK: transparent reporting and centralizing knowledge in extracellular vesicle research. Nat Methods. 2017;14:228\u201332. https:\/\/doi.org\/10.1038\/nmeth.4185.","journal-title":"Nat Methods"},{"key":"4960_CR15","unstructured":"EV-TRACK, ID EV250013. https:\/\/www.evtrack.org\/search_results.php?evtraqid=EV250013&submit=1&metric_year=2025. Accessed 15 May 2026."},{"key":"4960_CR16","doi-asserted-by":"publisher","first-page":"211","DOI":"10.1007\/978-1-0716-3203-1_15","volume-title":"Cell-Secreted Vesicles","author":"N Byts","year":"2023","unstructured":"Byts N, Makieieva O, Zhyvolozhnyi A, Bart G, Korvala J, Hekkala J, et al. Purification of Bacterial-Enriched Extracellular Vesicle Samples from Feces by Density Gradient Ultracentrifugation. In: Vainio S, editor. Cell-Secreted Vesicles. New York, NY: Springer US; 2023. pp. 211\u201326. https:\/\/doi.org\/10.1007\/978-1-0716-3203-1_15."},{"key":"4960_CR17","unstructured":"Bacterial extracellular vesicles from Amniotic and\u2026 ID 1235131). - BioProject - NCBI. https:\/\/www.ncbi.nlm.nih.gov\/bioproject\/PRJNA1235131. Accessed 12 May 2026."},{"key":"4960_CR18","doi-asserted-by":"publisher","first-page":"852","DOI":"10.1038\/s41587-019-0209-9","volume":"37","author":"E Bolyen","year":"2019","unstructured":"Bolyen E, Rideout JR, Dillon MR, Bokulich NA, Abnet CC, Al-Ghalith GA, et al. Reproducible, interactive, scalable and extensible microbiome data science using QIIME 2. Nat Biotechnol. 2019;37:852\u20137. https:\/\/doi.org\/10.1038\/s41587-019-0209-9.","journal-title":"Nat Biotechnol"},{"key":"4960_CR19","doi-asserted-by":"publisher","first-page":"10","DOI":"10.14806\/ej.17.1.200","volume":"17","author":"M Martin","year":"2011","unstructured":"Martin M. Cutadapt removes adapter sequences from high-throughput sequencing reads. EMBnet J. 2011;17:10\u20132. https:\/\/doi.org\/10.14806\/ej.17.1.200.","journal-title":"EMBnet J"},{"key":"4960_CR20","doi-asserted-by":"publisher","first-page":"e2584","DOI":"10.7717\/peerj.2584","volume":"4","author":"T Rognes","year":"2016","unstructured":"Rognes T, Flouri T, Nichols B, Quince C, Mah\u00e9 F. VSEARCH: a versatile open source tool for metagenomics. PeerJ. 2016;4:e2584. https:\/\/doi.org\/10.7717\/peerj.2584.","journal-title":"PeerJ"},{"key":"4960_CR21","doi-asserted-by":"publisher","unstructured":"Quast C, Pruesse E, Yilmaz P, Gerken J, Schweer T, Yarza P, et al. The SILVA ribosomal RNA gene database project: improved data processing and web-based tools. Nucleic Acids Res. 2013;41. https:\/\/doi.org\/10.1093\/nar\/gks1219. Database issue: D590\u20136.","DOI":"10.1093\/nar\/gks1219"},{"key":"4960_CR22","doi-asserted-by":"publisher","unstructured":"Robeson MS, O\u2019Rourke DR, Kaehler BD, Ziemski M, Dillon MR, Foster JT et al. RESCRIPt: Reproducible sequence taxonomy reference database management for the masses. 2020. https:\/\/doi.org\/10.1101\/2020.10.05.326504","DOI":"10.1101\/2020.10.05.326504"},{"key":"4960_CR23","doi-asserted-by":"publisher","first-page":"226","DOI":"10.1186\/s40168-018-0605-2","volume":"6","author":"NM Davis","year":"2018","unstructured":"Davis NM, Proctor DM, Holmes SP, Relman DA, Callahan BJ. Simple statistical identification and removal of contaminant sequences in marker-gene and metagenomics data. Microbiome. 2018;6:226. https:\/\/doi.org\/10.1186\/s40168-018-0605-2.","journal-title":"Microbiome"},{"key":"4960_CR24","unstructured":"Team RC. R: A language and environment for statistical computing. R Foundation for Statistical Computing. https:\/\/www.R-project.org\/. 2020."},{"key":"4960_CR25","doi-asserted-by":"crossref","unstructured":"Wickham H. ggplot2: Elegant Graphics for Data Analysis. https:\/\/ggplot2.tidyverse.org .2016.","DOI":"10.1007\/978-3-319-24277-4_9"},{"key":"4960_CR26","unstructured":"Bisanz JE. qiime2R: Importing QIIME2 artifacts and associated data into R sessions. 2018."},{"key":"4960_CR27","doi-asserted-by":"publisher","unstructured":"Turunen J. The code used for analysis in a manuscript entitled Maternal Factors Associate with Microbiota-Derived Extracellular Vesicle profiles in Pregnancy: A Clinical Cohort Study. 2026. https:\/\/doi.org\/10.5281\/ZENODO.15119368","DOI":"10.5281\/ZENODO.15119368"},{"key":"4960_CR28","doi-asserted-by":"publisher","first-page":"3514","DOI":"10.1038\/s41467-020-17041-7","volume":"11","author":"H Lin","year":"2020","unstructured":"Lin H, Peddada SD. Analysis of compositions of microbiomes with bias correction. Nat Commun. 2020;11:3514. https:\/\/doi.org\/10.1038\/s41467-020-17041-7.","journal-title":"Nat Commun"},{"key":"4960_CR29","unstructured":"Weir CB, Jan A. BMI Classification Percentile And Cut Off Points. StatPearls. 2025. https:\/\/www.ncbi.nlm.nih.gov\/books\/NBK541070\/ Accessed 30 Jan 2025."},{"key":"4960_CR30","doi-asserted-by":"publisher","first-page":"425","DOI":"10.1007\/s00281-018-0680-2","volume":"40","author":"S Nair","year":"2018","unstructured":"Nair S, Salomon C. Extracellular vesicles and their immunomodulatory functions in pregnancy. Semin Immunopathol. 2018;40:425\u201337. https:\/\/doi.org\/10.1007\/s00281-018-0680-2.","journal-title":"Semin Immunopathol"},{"key":"4960_CR31","doi-asserted-by":"publisher","first-page":"297","DOI":"10.20517\/evcna.2024.21","volume":"5","author":"E Amabebe","year":"2024","unstructured":"Amabebe E, Kumar A, Tatiparthy M, Kammala AK, Taylor BD, Menon R. Cargo exchange between human and bacterial extracellular vesicles in gestational tissues: a new paradigm in communication and immune development. Extracell Vesicles Circ Nucleic Acids. 2024;5:297\u2013328. https:\/\/doi.org\/10.20517\/evcna.2024.21.","journal-title":"Extracell Vesicles Circ Nucleic Acids"},{"key":"4960_CR32","doi-asserted-by":"publisher","first-page":"2527","DOI":"10.3390\/ijms24032527","volume":"24","author":"E Nunzi","year":"2023","unstructured":"Nunzi E, Mezzasoma L, Bellezza I, Zelante T, Orvietani P, Coata G, et al. Microbiota-Associated HAF-EVs Regulate Monocytes by Triggering or Inhibiting Inflammasome Activation. Int J Mol Sci. 2023;24:2527. https:\/\/doi.org\/10.3390\/ijms24032527.","journal-title":"Int J Mol Sci"},{"key":"4960_CR33","doi-asserted-by":"publisher","first-page":"1213234","DOI":"10.3389\/fmicb.2023.1213234","volume":"14","author":"R Menon","year":"2023","unstructured":"Menon R, Khanipov K, Radnaa E, Ganguly E, Bento GFC, Urrabaz-Garza R, et al. Amplification of microbial DNA from bacterial extracellular vesicles from human placenta. Front Microbiol. 2023;14:1213234. https:\/\/doi.org\/10.3389\/fmicb.2023.1213234.","journal-title":"Front Microbiol"},{"key":"4960_CR34","doi-asserted-by":"publisher","first-page":"528202","DOI":"10.3389\/fimmu.2020.528202","volume":"11","author":"N Di Simone","year":"2020","unstructured":"Di Simone N, Santamaria Ortiz A, Specchia M, Tersigni C, Villa P, Gasbarrini A, et al. Recent Insights on the Maternal Microbiota: Impact on Pregnancy Outcomes. Front Immunol. 2020;11:528202. https:\/\/doi.org\/10.3389\/fimmu.2020.528202.","journal-title":"Front Immunol"},{"key":"4960_CR35","doi-asserted-by":"publisher","first-page":"894","DOI":"10.1093\/ajcn\/88.4.894","volume":"88","author":"MC Collado","year":"2008","unstructured":"Collado MC, Isolauri E, Laitinen K, Salminen S. Distinct composition of gut microbiota during pregnancy in overweight and normal-weight women. Am J Clin Nutr. 2008;88:894\u20139. https:\/\/doi.org\/10.1093\/ajcn\/88.4.894.","journal-title":"Am J Clin Nutr"},{"key":"4960_CR36","doi-asserted-by":"publisher","first-page":"2214","DOI":"10.2337\/db16-0278","volume":"65","author":"LF Gomez-Arango","year":"2016","unstructured":"Gomez-Arango LF, Barrett HL, McIntyre HD, Callaway LK, Morrison M, Dekker Nitert M. Connections Between the Gut Microbiome and Metabolic Hormones in Early Pregnancy in Overweight and Obese Women. Diabetes. 2016;65:2214\u201323. https:\/\/doi.org\/10.2337\/db16-0278.","journal-title":"Diabetes"},{"key":"4960_CR37","doi-asserted-by":"publisher","first-page":"470","DOI":"10.1016\/j.cell.2012.07.008","volume":"150","author":"O Koren","year":"2012","unstructured":"Koren O, Goodrich JK, Cullender TC, Spor A, Laitinen K, Kling B\u00e4ckhed H, et al. Host Remodeling of the Gut Microbiome and Metabolic Changes during Pregnancy. Cell. 2012;150:470\u201380. https:\/\/doi.org\/10.1016\/j.cell.2012.07.008.","journal-title":"Cell"},{"key":"4960_CR38","doi-asserted-by":"publisher","DOI":"10.1186\/s12916-016-0636-0","author":"AA Kuperman","year":"2016","unstructured":"Kuperman AA, Koren O. Antibiotic use during pregnancy: how bad is it? BMC Med. 2016. https:\/\/doi.org\/10.1186\/s12916-016-0636-0. 14:91, s12916-016-0636\u20130."},{"key":"4960_CR39","doi-asserted-by":"publisher","first-page":"629","DOI":"10.1111\/1469-0691.12411","volume":"20","author":"J Stokholm","year":"2014","unstructured":"Stokholm J, Schj\u00f8rring S, Eskildsen CE, Pedersen L, Bischoff AL, F\u00f8lsgaard N, et al. Antibiotic use during pregnancy alters the commensal vaginal microbiota. Clin Microbiol Infect. 2014;20:629\u201335. https:\/\/doi.org\/10.1111\/1469-0691.12411.","journal-title":"Clin Microbiol Infect"},{"key":"4960_CR40","doi-asserted-by":"publisher","first-page":"112678","DOI":"10.1016\/j.biopha.2022.112678","volume":"147","author":"J Geng","year":"2022","unstructured":"Geng J, Ni Q, Sun W, Li L, Feng X. The links between gut microbiota and obesity and obesity related diseases. Biomed Pharmacother. 2022;147:112678. https:\/\/doi.org\/10.1016\/j.biopha.2022.112678.","journal-title":"Biomed Pharmacother"},{"key":"4960_CR41","doi-asserted-by":"publisher","first-page":"174","DOI":"10.1038\/nature09944","volume":"473","author":"M Arumugam","year":"2011","unstructured":"MetaHIT Consortium (additional members), Arumugam M, Raes J, Pelletier E, Le Paslier D, Yamada T, et al. Enterotypes of the human gut microbiome. Nature. 2011;473:174\u201380. https:\/\/doi.org\/10.1038\/nature09944.","journal-title":"Nature"},{"key":"4960_CR42","doi-asserted-by":"publisher","first-page":"e00111","DOI":"10.1128\/Spectrum.00111-21","volume":"9","author":"L Liu","year":"2021","unstructured":"Liu L, Feng Y, Wei L, Zong Z. Genome-Based Taxonomy of Brevundimonas with Reporting Brevundimonas huaxiensis sp. nov. Microbiol Spectr. 2021;9:e00111\u201321. https:\/\/doi.org\/10.1128\/Spectrum.00111-21.","journal-title":"Microbiol Spectr"},{"key":"4960_CR43","doi-asserted-by":"publisher","first-page":"e97876","DOI":"10.1371\/journal.pone.0097876","volume":"9","author":"M Laurence","year":"2014","unstructured":"Laurence M, Hatzis C, Brash DE. Common Contaminants in Next-Generation Sequencing That Hinder Discovery of Low-Abundance Microbes. PLoS ONE. 2014;9:e97876. https:\/\/doi.org\/10.1371\/journal.pone.0097876.","journal-title":"PLoS ONE"},{"key":"4960_CR44","doi-asserted-by":"publisher","first-page":"e12404","DOI":"10.1002\/jev2.12404","volume":"13","author":"JA Welsh","year":"2024","unstructured":"Welsh JA, Goberdhan DCI, O\u2019Driscoll L, Buzas EI, Blenkiron C, Bussolati B, et al. Minimal information for studies of extracellular vesicles (MISEV2023): From basic to advanced approaches. J Extracell Vesicles. 2024;13:e12404. https:\/\/doi.org\/10.1002\/jev2.12404.","journal-title":"J Extracell Vesicles"},{"key":"4960_CR45","doi-asserted-by":"publisher","first-page":"87","DOI":"10.1186\/s12915-014-0087-z","volume":"12","author":"SJ Salter","year":"2014","unstructured":"Salter SJ, Cox MJ, Turek EM, Calus ST, Cookson WO, Moffatt MF, et al. Reagent and laboratory contamination can critically impact sequence-based microbiome analyses. BMC Biol. 2014;12:87. https:\/\/doi.org\/10.1186\/s12915-014-0087-z.","journal-title":"BMC Biol"},{"key":"4960_CR46","doi-asserted-by":"publisher","first-page":"572912","DOI":"10.3389\/fcimb.2020.572912","volume":"10","author":"J Ramirez","year":"2020","unstructured":"Ramirez J, Guarner F, Bustos Fernandez L, Maruy A, Sdepanian VL, Cohen H. Antibiotics as Major Disruptors of Gut Microbiota. Front Cell Infect Microbiol. 2020;10:572912. https:\/\/doi.org\/10.3389\/fcimb.2020.572912.","journal-title":"Front Cell Infect Microbiol"},{"key":"4960_CR47","doi-asserted-by":"publisher","first-page":"23129","DOI":"10.1038\/srep23129","volume":"6","author":"MC Collado","year":"2016","unstructured":"Collado MC, Rautava S, Aakko J, Isolauri E, Salminen S. Human gut colonisation may be initiated in utero by distinct microbial communities in the placenta and amniotic fluid. Sci Rep. 2016;6:23129. https:\/\/doi.org\/10.1038\/srep23129.","journal-title":"Sci Rep"},{"key":"4960_CR48","doi-asserted-by":"publisher","first-page":"87","DOI":"10.1186\/s40168-018-0475-7","volume":"6","author":"ES Lim","year":"2018","unstructured":"Lim ES, Rodriguez C, Holtz LR. Amniotic fluid from healthy term pregnancies does not harbor a detectable microbial community. Microbiome. 2018;6:87\u20138. https:\/\/doi.org\/10.1186\/s40168-018-0475-7.","journal-title":"Microbiome"},{"key":"4960_CR49","doi-asserted-by":"publisher","first-page":"289","DOI":"10.1016\/j.ajog.2018.05.028","volume":"219","author":"EM Rehbinder","year":"2018","unstructured":"Rehbinder EM, Carlsen KCL, Staff AC, Angell IL, Landr\u00f8 L, Hilde K, et al. Is amniotic fluid of women with uncomplicated term pregnancies free of bacteria? Am J Obstet Gynecol. 2018;219:289. https:\/\/doi.org\/10.1016\/j.ajog.2018.05.028. .e1-289.e12.","journal-title":"Am J Obstet Gynecol"},{"key":"4960_CR50","doi-asserted-by":"publisher","first-page":"1124","DOI":"10.3389\/fmicb.2019.01124","volume":"10","author":"LF Stinson","year":"2019","unstructured":"Stinson LF, Boyce MC, Payne MS, Keelan JA. The Not-so-Sterile Womb: Evidence That the Human Fetus Is Exposed to Bacteria Prior to Birth. Front Microbiol. 2019;10:1124. https:\/\/doi.org\/10.3389\/fmicb.2019.01124.","journal-title":"Front Microbiol"},{"key":"4960_CR51","doi-asserted-by":"publisher","first-page":"19449","DOI":"10.1038\/s41598-021-98951-4","volume":"11","author":"J Turunen","year":"2021","unstructured":"Turunen J, Tejesvi MV, Paalanne N, Hekkala J, Lindgren O, Kaakinen M, et al. Presence of distinctive microbiome in the first-pass meconium of newborn infants. Sci Rep. 2021;11:19449. https:\/\/doi.org\/10.1038\/s41598-021-98951-4.","journal-title":"Sci Rep"},{"key":"4960_CR52","doi-asserted-by":"publisher","first-page":"1794266","DOI":"10.1080\/19490976.2020.1794266","volume":"12","author":"Q He","year":"2020","unstructured":"He Q, Kwok L-Y, Xi X, Zhong Z, Ma T, Xu H, et al. The meconium microbiota shares more features with the amniotic fluid microbiota than the maternal fecal and vaginal microbiota. Gut Microbes. 2020;12:1794266. https:\/\/doi.org\/10.1080\/19490976.2020.1794266.","journal-title":"Gut Microbes"},{"key":"4960_CR53","doi-asserted-by":"publisher","first-page":"887","DOI":"10.1038\/s41390-022-02242-1","volume":"93","author":"J Turunen","year":"2022","unstructured":"Turunen J, Tejesvi MV, Suokas M, Virtanen N, Paalanne N, Kaisanlahti A, et al. Bacterial extracellular vesicles in the microbiome of first-pass meconium in newborn infants. Pediatr Res. 2022;93:887\u201396. https:\/\/doi.org\/10.1038\/s41390-022-02242-1.","journal-title":"Pediatr Res"},{"key":"4960_CR54","doi-asserted-by":"publisher","first-page":"895","DOI":"10.1007\/s00125-016-3880-0","volume":"59","author":"TK Soderborg","year":"2016","unstructured":"Soderborg TK, Borengasser SJ, Barbour LA, Friedman JE. Microbial transmission from mothers with obesity or diabetes to infants: an innovative opportunity to interrupt a vicious cycle. Diabetologia. 2016;59:895\u2013906. https:\/\/doi.org\/10.1007\/s00125-016-3880-0.","journal-title":"Diabetologia"},{"key":"4960_CR55","doi-asserted-by":"publisher","first-page":"9410","DOI":"10.1038\/s41467-024-53279-1","volume":"15","author":"N De Langhe","year":"2024","unstructured":"De Langhe N, Van Dorpe S, Guilbert N, Vander Cruyssen A, Roux Q, Deville S, et al. Mapping bacterial extracellular vesicle research: insights, best practices and knowledge gaps. Nat Commun. 2024;15:9410. https:\/\/doi.org\/10.1038\/s41467-024-53279-1.","journal-title":"Nat Commun"},{"key":"4960_CR56","doi-asserted-by":"publisher","first-page":"e330","DOI":"10.1038\/emm.2017.47","volume":"49","author":"H-I Choi","year":"2017","unstructured":"Choi H-I, Choi J-P, Seo J, Kim BJ, Rho M, Han JK, et al. Helicobacter pylori-derived extracellular vesicles increased in the gastric juices of gastric adenocarcinoma patients and induced inflammation mainly via specific targeting of gastric epithelial cells. Exp Mol Med. 2017;49:e330\u2013330. https:\/\/doi.org\/10.1038\/emm.2017.47.","journal-title":"Exp Mol Med"},{"key":"4960_CR57","doi-asserted-by":"publisher","first-page":"1377","DOI":"10.1007\/s10068-018-0372-7","volume":"27","author":"YJ Choi","year":"2018","unstructured":"Choi YJ, Lee DH, Kim HS, Kim Y-K. An exploratory study on the effect of daily fruits and vegetable juice on human gut microbiota. Food Sci Biotechnol. 2018;27:1377\u201386. https:\/\/doi.org\/10.1007\/s10068-018-0372-7.","journal-title":"Food Sci Biotechnol"}],"container-title":["BMC Medicine"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s12916-026-04960-3","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12916-026-04960-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12916-026-04960-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,11]],"date-time":"2026-06-11T08:33:26Z","timestamp":1781166806000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1186\/s12916-026-04960-3"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,5,28]]},"references-count":57,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["4960"],"URL":"https:\/\/doi.org\/10.1186\/s12916-026-04960-3","relation":{"references":[{"id-type":"doi","id":"10.1186\/s12916-016-0636-0","asserted-by":"subject"}]},"ISSN":["1741-7015"],"issn-type":[{"value":"1741-7015","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,5,28]]},"assertion":[{"value":"26 February 2026","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"22 May 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"28 May 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":"The research plan was reviewed by the Ethics Committee of Oulu University Hospital, Finland, before the study, who issued decision number ETTMK:3\/2016. The participants provided written informed consent before the study.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval and consent to participate"}},{"value":"Not applicable.","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":"352"}}