{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"institution":[{"name":"bioRxiv"}],"indexed":{"date-parts":[[2026,1,16]],"date-time":"2026-01-16T00:28:30Z","timestamp":1768523310212,"version":"3.49.0"},"posted":{"date-parts":[[2025,5,24]]},"group-title":"Microbiology","reference-count":42,"publisher":"openRxiv","license":[{"start":{"date-parts":[[2025,5,24]],"date-time":"2025-05-24T00:00:00Z","timestamp":1748044800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Cambridge Africa ALBORADA Research Fund","award":["G115009"],"award-info":[{"award-number":["G115009"]}]},{"name":"Gates Foundation","award":["OPP1017641"],"award-info":[{"award-number":["OPP1017641"]}]},{"name":"Gates Foundation","award":["OPP1017579"],"award-info":[{"award-number":["OPP1017579"]}]},{"name":"NIH H3Africa","award":["1U01AI110466-01A1"],"award-info":[{"award-number":["1U01AI110466-01A1"]}]},{"DOI":"10.13039\/100010269","name":"Wellcome Trust","doi-asserted-by":"crossref","award":["221372\/Z\/20\/Z"],"award-info":[{"award-number":["221372\/Z\/20\/Z"]}],"id":[{"id":"10.13039\/100010269","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"accepted":{"date-parts":[[2025,5,24]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                <jats:p>\n                  <jats:italic>Ornithobacterium hominis<\/jats:italic>\n                  is a recently described Gram-negative bacterium that colonises the human nasopharynx and may be associated with poor upper respiratory tract health. Here, we describe the isolation of\n                  <jats:italic>O. hominis<\/jats:italic>\n                  from samples collected from a South African birth cohort, creating the first archive of cultured strains of the species from Africa. Sequenced genomes from this archive reveal that South African\n                  <jats:italic>O. hominis<\/jats:italic>\n                  is more similar to Australian strains than those from Southeast Asia, and that it may share genes with other members of the microbiome that are relevant for virulence, colonisation, and antibiotic resistance. Leveraging existing microbiome data from the cohort,\n                  <jats:italic>O. hominis<\/jats:italic>\n                  was found to be closely associated with bacterial co-colonisers that are rare in non-carrier individuals, including\n                  <jats:italic>Suttonella<\/jats:italic>\n                  ,\n                  <jats:italic>Helcococcus<\/jats:italic>\n                  ,\n                  <jats:italic>Moraxella<\/jats:italic>\n                  spp., and Gracilibacteria. Their collective acquisition has a significant impact on the diversity of nasopharyngeal communities that contain\n                  <jats:italic>O. hominis<\/jats:italic>\n                  . Individuals who have not yet acquired\n                  <jats:italic>O. hominis<\/jats:italic>\n                  have a higher abundance of\n                  <jats:italic>Moraxella<\/jats:italic>\n                  (particularly\n                  <jats:italic>M. lincolnii<\/jats:italic>\n                  ) than individuals who never acquire\n                  <jats:italic>O. hominis<\/jats:italic>\n                  , suggesting that this could be a precursor state for successful colonisation. Finally, a novel co-coloniser species,\n                  <jats:italic>Helcococcus ekapensis<\/jats:italic>\n                  , was successfully isolated and sequenced.\n                <\/jats:p>\n                <jats:sec>\n                  <jats:title>Data Summary<\/jats:title>\n                  <jats:p>\n                    <jats:italic>Ornithobacterium hominis<\/jats:italic>\n                    data have been deposited under project accession ERP149886. This comprises genome sequences for isolates SA-OH-C1 (ERR13967269), SA-OH-C2 (ERR13967270), SA-OH-C3 (ERR13967271), SA-OH-C4 (ERR13967272, ERR13967275), SA-OH-C5 (ERR13967273), SA-OH-C6 (ERR13967274, ERR13967276). Previously published 16S rRNA gene data are deposited under project accessions PRJNA790843 and PRJNA548658.\n                    <jats:italic>Helcococcus ekapensis<\/jats:italic>\n                    genome data are deposited under project accession PRJEB85661.\n                  <\/jats:p>\n                <\/jats:sec>\n                <jats:sec>\n                  <jats:title>Software used<\/jats:title>\n                  <jats:list list-type=\"bullet\">\n                    <jats:list-item>\n                      <jats:p>\n                        AMRFinderPlus v3.12.8:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/github.com\/ncbi\/amr\">https:\/\/github.com\/ncbi\/amr<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        AssembleBAC-ONT v1.1.1:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/github.com\/avantonder\/assembleBAC-ONT\">https:\/\/github.com\/avantonder\/assembleBAC-ONT<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        BAKTA v1.8.1:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/bakta.computational.bio\/\">https:\/\/bakta.computational.bio\/<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        BLAST v2.16.0:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/blast.ncbi.nlm.nih.gov\/Blast.cgi\">https:\/\/blast.ncbi.nlm.nih.gov\/Blast.cgi<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        Comprehensive Antibiotic Resistance Database (CARD) Resistance Gene Identifier (RGI) tool v1.2.1:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/card.mcmaster.ca\/analyze\/rgi\">https:\/\/card.mcmaster.ca\/analyze\/rgi<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        Decontam v1.12 (R package):\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/github.com\/benjjneb\/decontam\">https:\/\/github.com\/benjjneb\/decontam<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        Eggnog-mapper v2.0.1:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"http:\/\/eggnog-mapper.embl.de\/\">http:\/\/eggnog-mapper.embl.de\/<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        FastANI v1.1.0:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/github.com\/ParBLiSS\/FastANI\">https:\/\/github.com\/ParBLiSS\/FastANI<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        Flye v2.9.2:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/github.com\/fenderglass\/Flye\">https:\/\/github.com\/fenderglass\/Flye<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        Guppy v6.5.7:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/community.nanoporetech.com\/downloads\/guppy\/release_notes\">https:\/\/community.nanoporetech.com\/downloads\/guppy\/release_notes<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        ISEScan v1.7.2.3:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/usegalaxy.eu\/root?tool_id=toolshed.g2.bx.psu.edu\/repos\/iuc\/isescan\/isescan\/1.7.2.3+galaxy1\">https:\/\/usegalaxy.eu\/root?tool_id=toolshed.g2.bx.psu.edu\/repos\/iuc\/isescan\/isescan\/1.7.2.3+galaxy1<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        Medaka v1.9.1:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/github.com\/nanoporetech\/medaka\">https:\/\/github.com\/nanoporetech\/medaka<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        MEGA11:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/www.megasoftware.net\/\">https:\/\/www.megasoftware.net\/<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        MMseqs2 v17:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/github.com\/soedinglab\/MMseqs2\">https:\/\/github.com\/soedinglab\/MMseqs2<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        Mothur v1.44.3:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/github.com\/mothur\/mothur\">https:\/\/github.com\/mothur\/mothur<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        NetCoMi v1.2.0 (R package):\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/github.com\/stefpeschel\/NetCoMi\">https:\/\/github.com\/stefpeschel\/NetCoMi<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        Panaroo v1.4.3:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/github.com\/gtonkinhill\/panaroo\">https:\/\/github.com\/gtonkinhill\/panaroo<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        PHASTEST:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/phastest.ca\/submissions\/new\">https:\/\/phastest.ca\/submissions\/new<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        Prowler (commit ID c3041ba):\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/github.com\/ProwlerForNanopore\/ProwlerTrimmer\">https:\/\/github.com\/ProwlerForNanopore\/ProwlerTrimmer<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        R v4.4.3:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/www.r-project.org\/\">https:\/\/www.r-project.org\/<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                  <\/jats:list>\n                <\/jats:sec>\n                <jats:sec>\n                  <jats:title>Databases used<\/jats:title>\n                  <jats:list list-type=\"bullet\">\n                    <jats:list-item>\n                      <jats:p>\n                        Comprehensive Antibiotic Resistance Database (CARD):\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/card.mcmaster.ca\/\">https:\/\/card.mcmaster.ca\/<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        European Nucleotide Archive:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/www.ebi.ac.uk\/ena\/\">https:\/\/www.ebi.ac.uk\/ena\/<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        Genome Taxonomy Database (GTDB) release 09-RS220:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/gtdb.ecogenomic.org\/\">https:\/\/gtdb.ecogenomic.org\/<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        RefSeq release 228:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/www.ncbi.nlm.nih.gov\/refseq\/about\/prokaryotes\/\">https:\/\/www.ncbi.nlm.nih.gov\/refseq\/about\/prokaryotes\/<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\n                        SILVA v132:\n                        <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/www.arb-silva.de\/\">https:\/\/www.arb-silva.de\/<\/jats:ext-link>\n                      <\/jats:p>\n                    <\/jats:list-item>\n                  <\/jats:list>\n                <\/jats:sec>\n                <jats:sec>\n                  <jats:title>Impact statement<\/jats:title>\n                  <jats:p>\n                    First described in 2019,\n                    <jats:italic>Ornithobacterium hominis<\/jats:italic>\n                    is an understudied bacterium that may be associated with poor respiratory health in children. The study builds upon existing knowledge of\n                    <jats:italic>O. hominis<\/jats:italic>\n                    by describing the first African isolates of the species, its potential as a reservoir of virulence and antibiotic resistance genes in the upper respiratory tract, and the unique microbiome profile of\n                    <jats:italic>O. hominis<\/jats:italic>\n                    carriers.\n                  <\/jats:p>\n                <\/jats:sec>","DOI":"10.1101\/2025.05.24.655922","type":"posted-content","created":{"date-parts":[[2025,5,25]],"date-time":"2025-05-25T00:35:12Z","timestamp":1748133312000},"source":"Crossref","is-referenced-by-count":0,"title":["Characterisation of\n                  <i>Ornithobacterium hominis<\/i>\n                  colonisation dynamics and interaction with the nasopharyngeal microbiome in a South African birth cohort"],"prefix":"10.64898","author":[{"given":"Celine C","family":"De Allende","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Susannah J","family":"Salter","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Siobhan E","family":"Brigg","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Shantelle","family":"Claassen-Weitz","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Kilaza S","family":"Mwaikono","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Lesley","family":"Workman","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Heather J","family":"Zar","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Mark P","family":"Nicol","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Julian","family":"Parkhill","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Felix S","family":"Dube","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"54368","reference":[{"key":"2025052800551002000_2025.05.24.655922v1.1","doi-asserted-by":"publisher","DOI":"10.1001\/jamanetworkopen.2023.22494"},{"key":"2025052800551002000_2025.05.24.655922v1.2","doi-asserted-by":"crossref","unstructured":"Nair H , Nokes DJ , Gessner BD , Dherani M , Madhi SA , Singleton RJ , et al. Global burden of acute lower respiratory infections due to respiratory syncytial virus in young children: a systematic review and meta-analysis. Lancet. 2010;375(9725):1545\u201355.","DOI":"10.1016\/S0140-6736(10)60206-1"},{"key":"2025052800551002000_2025.05.24.655922v1.3","unstructured":"GBD 2016 Lower Respiratory Infections Collaborators. Estimates of the global, regional, and national morbidity, mortality, and aetiologies of lower respiratory infections in 195 countries, 1990\u20132016: a systematic analysis for the Global Burden of Disease Study 2016. The Lancet Infectious Diseases. 2018;18(11):1191\u2013210."},{"key":"2025052800551002000_2025.05.24.655922v1.4","doi-asserted-by":"crossref","unstructured":"L\u00f8vlie A , Vestrheim DF , Aaberge IS , Steens A . Changes in pneumococcal carriage prevalence and factors associated with carriage in Norwegian children, four years after introduction of PCV13. BMC Infectious Diseases. 2020;20:29.","DOI":"10.1186\/s12879-019-4754-0"},{"key":"2025052800551002000_2025.05.24.655922v1.5","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1006665"},{"issue":"21","key":"2025052800551002000_2025.05.24.655922v1.6","first-page":"e00474","article-title":"Interaction between Streptococcus pneumoniae and Staphylococcus aureus Generates \u00b7OH Radicals That Rapidly Kill Staphylococcus aureus Strains","volume":"201","year":"2019","journal-title":"Journal of Bacteriology"},{"key":"2025052800551002000_2025.05.24.655922v1.7","doi-asserted-by":"publisher","DOI":"10.1042\/bj0160499"},{"key":"2025052800551002000_2025.05.24.655922v1.8","doi-asserted-by":"publisher","DOI":"10.1128\/mSphere.00852-20"},{"key":"2025052800551002000_2025.05.24.655922v1.9","doi-asserted-by":"crossref","unstructured":"Margolis E , Yates A , Levin BR . The ecology of nasal colonization of Streptococcus pneumoniae, Haemophilus influenzae and Staphylococcus aureus: the role of competition and interactions with host\u2019s immune response. BMC Microbiology. 2010;10:59.","DOI":"10.1186\/1471-2180-10-59"},{"key":"2025052800551002000_2025.05.24.655922v1.10","doi-asserted-by":"crossref","unstructured":"Salter SJ , Turner C , Watthanaworawit W , Goffau MC de , Wagner J , Parkhill J , et al. A longitudinal study of the infant nasopharyngeal microbiota: The effects of age, illness and antibiotic use in a cohort of South East Asian children. PLOS Neglected Tropical Diseases [Internet]. 2017 Oct 2 [cited 2021 May 13];11(10):e0005975. Available from: https:\/\/journals.plos.org\/plosntds\/article?id=10.1371\/journal.pntd.0005975","DOI":"10.1371\/journal.pntd.0005975"},{"key":"2025052800551002000_2025.05.24.655922v1.11","doi-asserted-by":"crossref","unstructured":"Salter SJ , Scott P , Page AJ , Tracey A , de Goffau MC , Cormie C , et al. \u2018Candidatus Ornithobacterium hominis\u2019: insights gained from draft genomes obtained from nasopharyngeal swabs. Microb Genom [Internet]. 2019 Feb 5 [cited 2021 May 13];5(2). Available from: https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC6421346\/","DOI":"10.1099\/mgen.0.000247"},{"key":"2025052800551002000_2025.05.24.655922v1.12","doi-asserted-by":"crossref","unstructured":"Lawrence KA , Harris TM , Salter SJ , Hall RW , Smith-Vaughan HC , Chang AB , et al. Method for culturing Candidatus Ornithobacterium hominis. Journal of Microbiological Methods [Internet]. 2019 Apr 1 [cited 2021 May 13];159:157\u201360. Available from: https:\/\/www.sciencedirect.com\/science\/article\/pii\/S0167701219300326","DOI":"10.1016\/j.mimet.2019.03.006"},{"key":"2025052800551002000_2025.05.24.655922v1.13","doi-asserted-by":"crossref","unstructured":"Claassen-Weitz S , Gardner-Lubbe S , Xia Y , Mwaikono KS , Harris Mounaud S , Nierman WC , et al. Succession and determinants of the early life nasopharyngeal microbiota in a South African birth cohort. Microbiome. 2023;11:127.","DOI":"10.1186\/s40168-023-01563-5"},{"issue":"2","key":"2025052800551002000_2025.05.24.655922v1.14","doi-asserted-by":"crossref","first-page":"e0036721","DOI":"10.1128\/Spectrum.00367-21","article-title":"Upper Respiratory Tract Microbiome of Australian Aboriginal and Torres Strait Islander Children in Ear and Nose Health and Disease","volume":"9","year":"2021","journal-title":"Microbiology Spectrum"},{"key":"2025052800551002000_2025.05.24.655922v1.15","doi-asserted-by":"publisher","DOI":"10.1136\/thoraxjnl-2014-206242"},{"key":"2025052800551002000_2025.05.24.655922v1.16","doi-asserted-by":"crossref","unstructured":"Claassen-Weitz S , Gardner-Lubbe S , Mwaikono KS , du Toit E , Zar HJ , Nicol MP . Optimizing 16S rRNA gene profile analysis from low biomass nasopharyngeal and induced sputum specimens. 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