{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,10,24]],"date-time":"2025-10-24T13:18:58Z","timestamp":1761311938557,"version":"3.41.2"},"reference-count":41,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2022,12,16]],"date-time":"2022-12-16T00:00:00Z","timestamp":1671148800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100013996","name":"Global Health Innovative Technology Fund","doi-asserted-by":"publisher","award":["GHIT 2013-105, G2014-109, G2016-106, G2019-208"],"award-info":[{"award-number":["GHIT 2013-105, G2014-109, G2016-106, G2019-208"]}],"id":[{"id":"10.13039\/501100013996","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001700","name":"Ministry of Education, Culture, Sports, Science and Technology","doi-asserted-by":"publisher","award":["15651988, 24249024, 38201103-01"],"award-info":[{"award-number":["15651988, 24249024, 38201103-01"]}],"id":[{"id":"10.13039\/501100001700","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100009619","name":"Japan Agency for Medical Research and Development","doi-asserted-by":"publisher","award":["JP20lm0203135, 17nk0101206j0003"],"award-info":[{"award-number":["JP20lm0203135, 17nk0101206j0003"]}],"id":[{"id":"10.13039\/100009619","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100009099","name":"Irish Aid","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100009099","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Cell. Infect. Microbiol."],"abstract":"<jats:p>BK-SE36, based on <jats:italic>Plasmodium falciparum<\/jats:italic> serine repeat antigen 5 (SERA5), is a blood-stage malaria vaccine candidate currently being evaluated in clinical trials. Phase 1 trials in Uganda and Burkina Faso have demonstrated promising safety and immunogenicity profiles. However, the genetic diversity of <jats:italic>sera5<\/jats:italic> in Africa and the role of allele\/variant-specific immunity remain a major concern. Here, sequence analyses were done on 226 strains collected from the two clinical trial\/follow-up studies and 88 strains from two cross-sectional studies in Africa. Compared to other highly polymorphic vaccine candidate antigens, polymorphisms in <jats:italic>sera5<\/jats:italic> were largely confined to the repeat regions of the gene. Results also confirmed a SERA5 consensus sequence with African-specific polymorphisms. Mismatches with the vaccine-type SE36 (BK-SE36) in the octamer repeat, serine repeat, and flanking regions, and single-nucleotide polymorphisms in non-repeat regions could compromise vaccine response and efficacy. However, the haplotype diversity of SERA5 was similar between vaccinated and control participants. There was no marked bias or difference in the patterns of distribution of the SE36 haplotype and no statistically significant genetic differentiation among parasites infecting BK-SE36 vaccinees and controls. Results indicate that BK-SE36 does not elicit an allele-specific immune response.<\/jats:p>","DOI":"10.3389\/fcimb.2022.1058081","type":"journal-article","created":{"date-parts":[[2022,12,16]],"date-time":"2022-12-16T17:00:19Z","timestamp":1671210019000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":2,"title":["African-specific polymorphisms in Plasmodium falciparum serine repeat antigen 5 in Uganda and Burkina Faso clinical samples do not interfere with antibody response to BK-SE36 vaccination"],"prefix":"10.3389","volume":"12","author":[{"given":"Nobuko","family":"Arisue","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Nirianne Marie Q.","family":"Palacpac","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Edward H.","family":"Ntege","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Adoke","family":"Yeka","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Betty","family":"Balikagala","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Bernard N.","family":"Kanoi","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Edith Christiane","family":"Bougouma","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Alfred B.","family":"Tiono","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Issa","family":"Nebie","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Amidou","family":"Diarra","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Sophie","family":"Houard","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Flavia","family":"D\u2019Alessio","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Odile","family":"Leroy","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Sodiomon B.","family":"Sirima","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Thomas G.","family":"Egwang","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Toshihiro","family":"Horii","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1965","published-online":{"date-parts":[[2022,12,16]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"47533","DOI":"10.1074\/jbc.M207145200","article-title":"Serine repeat antigen (SERA5) is predominantly expressed among the SERA multigene family of Plasmodium falciparum, and the acquired antibody titers correlate with serum inhibition of the parasite growth","volume":"277","author":"Aoki","year":"2002","journal-title":"J. Biol. Chem."},{"key":"B2","doi-asserted-by":"publisher","first-page":"170","DOI":"10.1186\/s13071-020-04044-y","article-title":"Characteristic features of the SERA multigene family in the malaria parasite","volume":"13","author":"Arisue","year":"2020","journal-title":"Parasitol. Vectors."},{"key":"B3","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2022.978591","article-title":"Safety and immunogenicity of BK-SE36 in a blinded, randomized, controlled, age de-escalating phase ib clinical trial in Burkinabe children","volume":"13","author":"Bougouma","year":"2022","journal-title":"Front. Immunol."},{"key":"B4","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1016\/0166-6851(92)90010-h","article-title":"Intramolecular mapping of Plasmodium falciparum p126 proteolytic fragments by n-terminal amino acid sequencing","volume":"53","author":"Debrabant","year":"1992","journal-title":"Mol. Biochem. Parasitol."},{"key":"B5","doi-asserted-by":"publisher","first-page":"564","DOI":"10.1111\/j.1755-0998.2010.02847.x","article-title":"Arlequin suite ver 3.5: a new series of programs to perform population genetics analyses under Linux and windows","volume":"10","author":"Excoffier","year":"2010","journal-title":"Mol. Ecol. Resour."},{"key":"B6","doi-asserted-by":"publisher","first-page":"7246","DOI":"10.1016\/j.vaccine.2020.09.056","article-title":"First-in-human randomised trial and follow-up study of Plasmodium falciparum blood-stage malaria vaccine BK-SE36 with CpG-ODN(K3)","volume":"38","author":"Ezoe","year":"2020","journal-title":"Vaccine."},{"key":"B7","doi-asserted-by":"publisher","first-page":"121","DOI":"10.1006\/expr.1996.4118","article-title":"Plasmodium falciparum: An epitope within a highly conserved region of the 47-kDa amino-terminal domain of the serine repeat antigen is a target of parasite-inhibitory antibodies","volume":"85","author":"Fox","year":"1997","journal-title":"Exp. Parasitol."},{"key":"B8","doi-asserted-by":"publisher","first-page":"380","DOI":"10.1016\/j.parint.2010.05.002","article-title":"Evidences of protection against blood-stage infection of Plasmodium falciparum by the novel protein vaccine SE36","volume":"59","author":"Horii","year":"2010","journal-title":"Parasitol. Int."},{"key":"B9","doi-asserted-by":"publisher","first-page":"1547","DOI":"10.1093\/molbev\/msy096","article-title":"MEGA X: Molecular evolutionary genetics analysis across computing platforms","volume":"35","author":"Kumar","year":"2018","journal-title":"Mol. Biol. Evol."},{"key":"B10","doi-asserted-by":"publisher","first-page":"480","DOI":"10.1080\/21645515.2019.1669415","article-title":"RTS,S\/AS01 vaccine (Mosquirix\u2122): An overview","volume":"16","author":"Laurens","year":"2020","journal-title":"Hum. Vaccin. Immunother."},{"key":"B11","doi-asserted-by":"publisher","first-page":"1451","DOI":"10.1093\/bioinformatics\/btp187","article-title":"DnaSP v5: a software for comprehensive analysis of DNA polymorphism data","volume":"25","author":"Librado","year":"2009","journal-title":"Bioinformatics."},{"key":"B12","doi-asserted-by":"publisher","first-page":"177","DOI":"10.1016\/s0166-6851(01)00452-2","article-title":"Characterization of proteases involved in the processing of Plasmodium falciparum serine repeat antigen (SERA)","volume":"120","author":"Li","year":"2002","journal-title":"Mol. Biochem. Parasitol."},{"key":"B13","doi-asserted-by":"publisher","DOI":"10.1038\/d41586-021-02755-5","article-title":"Scientists hail historic malaria vaccine approval - but point to challenges ahead","author":"Maxmen","year":"2021","journal-title":"Nature"},{"key":"B14","doi-asserted-by":"publisher","first-page":"249","DOI":"10.1016\/s0166-6851(97)00038-8","article-title":"Sequence diversity in the amino-terminal 47 kDa fragment of the Plasmodium falciparum serine repeat antigen","volume":"86","author":"Morimatsu","year":"1997","journal-title":"Mol. Biochem. Parasitol."},{"key":"B15","doi-asserted-by":"publisher","first-page":"2025","DOI":"10.1056\/NEJMoa1505819","article-title":"Genetic diversity and protective efficacy of RTS,S\/AS01  malaria vaccine","volume":"373","author":"Neafsey","year":"2015","journal-title":"N. Engl. J. Med."},{"key":"B16","doi-asserted-by":"publisher","first-page":"418","DOI":"10.1093\/oxfordjournals.molbev.a040410","article-title":"Simple methods for estimating the numbers of synonymous and nonsynonymous nucleotide substitutions","volume":"3","author":"Nei","year":"1986","journal-title":"Mol. Biol. Evol."},{"key":"B17","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0004708","article-title":"Blood stage malaria vaccine eliciting high antigen-specific antibody concentrations confers no protection to young children in Western Kenya","volume":"4","author":"Ogutu","year":"2009","journal-title":"PloS One"},{"key":"B18","doi-asserted-by":"publisher","first-page":"191","DOI":"10.4269\/ajtmh.2006.74.191","article-title":"High titers of IgG antibodies against Plasmodium falciparum serine repeat antigen 5 (SERA5) are associated with protection against severe malaria in Ugandan children","volume":"74","author":"Okech","year":"2006","journal-title":"Am. J. Trop. Med. Hyg."},{"key":"B19","doi-asserted-by":"publisher","first-page":"912","DOI":"10.4269\/ajtmh.2001.65.912","article-title":"Natural human immunoglobulin G subclass responses to Plasmodium falciparum serine repeat antigen in Uganda","volume":"65","author":"Okech","year":"2001","journal-title":"Am. J. Trop. Med. Hyg."},{"key":"B20","doi-asserted-by":"publisher","first-page":"511","DOI":"10.1093\/infdis\/jis709","article-title":"Molecular basis of allele-specific efficacy of a blood-stage malaria vaccine: vaccine development implications","volume":"207","author":"Ouattara","year":"2013","journal-title":"J. Infect. Dis."},{"key":"B21","doi-asserted-by":"publisher","first-page":"237","DOI":"10.1016\/j.parint.2013.01.006","article-title":"Association of naturally acquired IgG antibodies against Plasmodium falciparum serine repeat antigen-5 with reduced placental parasitemia and normal birth weight in pregnant Ugandan women: a pilot study","volume":"62","author":"Owalla","year":"2013","journal-title":"Parasitol. Int."},{"key":"B22","doi-asserted-by":"publisher","first-page":"5837","DOI":"10.1016\/j.vaccine.2011.06.052","article-title":"Plasmodium falciparum serine repeat antigen 5 (SE36) as a malaria vaccine candidate","volume":"29","author":"Palacpac","year":"2011","journal-title":"Vaccine."},{"key":"B23","doi-asserted-by":"publisher","first-page":"F1000 Faculty Rev","DOI":"10.12688\/f1000research.22143.1","article-title":"Malaria vaccines: Facing unknowns","volume":"9","author":"Palacpac","year":"2020","journal-title":"F1000Res."},{"key":"B24","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0064073","article-title":"Phase 1b randomized trial and follow-up study in Uganda of the blood-stage malaria vaccine candidate BK-SE36","volume":"8","author":"Palacpac","year":"2013","journal-title":"PloS One"},{"key":"B25","doi-asserted-by":"publisher","first-page":"1299","DOI":"10.1016\/s0264-410x(98)00057-7","article-title":"Complement-mediated killing of Plasmodium falciparum erythrocytic schizont with antibodies to the recombinant serine repeat antigen (SERA)","volume":"16","author":"Pang","year":"1998","journal-title":"Vaccine."},{"key":"B26","doi-asserted-by":"publisher","first-page":"1821","DOI":"10.1128\/IAI.67.4.1821-1827.1999","article-title":"Antibodies reactive with the N-terminal domain of Plasmodium falciparum serine repeat antigen inhibit cell proliferation by agglutinating merozoites and schizonts","volume":"67","author":"Pang","year":"1999","journal-title":"Infect. Immun."},{"key":"B27","doi-asserted-by":"publisher","first-page":"830","DOI":"10.4269\/ajtmh.2011.10-0498","article-title":"Continuing intense malaria transmission in northern Uganda","volume":"84","author":"Proietti","year":"2011","journal-title":"Am. J. Trop. Med. Hyg."},{"key":"B28","doi-asserted-by":"publisher","first-page":"368","DOI":"10.1111\/mmi.12941","article-title":"Plasmodium falciparum SERA5 plays a non-enzymatic role in the malarial asexual blood-stage lifecycle","volume":"96","author":"Stallmach","year":"2015","journal-title":"Mol. Microbiol."},{"key":"B29","doi-asserted-by":"publisher","first-page":"1069","DOI":"10.1016\/0264-410x(95)00238-v","article-title":"Production of recombinant SERA proteins of Plasmodium falciparum in Escherichia coli by using synthetic genes","volume":"14","author":"Sugiyama","year":"1996","journal-title":"Vaccine."},{"key":"B30","doi-asserted-by":"publisher","first-page":"2ra5","DOI":"10.1126\/scitranslmed.3000257","article-title":"Extreme polymorphism in a vaccine antigen and risk of clinical malaria: implications for vaccine development","volume":"1","author":"Takala","year":"2009","journal-title":"Sci. Transl. Med."},{"key":"B31","doi-asserted-by":"publisher","first-page":"1583","DOI":"10.1016\/j.vaccine.2011.12.124","article-title":"Geographic differentiation of polymorphism in the Plasmodium falciparum malaria vaccine candidate gene SERA5","volume":"30","author":"Tanabe","year":"2012","journal-title":"Vaccine."},{"key":"B32","doi-asserted-by":"publisher","first-page":"1283","DOI":"10.1016\/j.cub.2010.05.053","article-title":"Plasmodium falciparum accompanied the human expansion out of Africa","volume":"20","author":"Tanabe","year":"2010","journal-title":"Curr. Biol."},{"key":"B33","doi-asserted-by":"publisher","first-page":"1004","DOI":"10.1056\/NEJMoa1008115","article-title":"A field trial to assess a blood-stage malaria vaccine","volume":"365","author":"Thera","year":"2011","journal-title":"N. Engl. J. Med."},{"key":"B34","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0086936","article-title":"Malaria incidence in children in south-West Burkina Faso: comparison of active and passive case detection methods","volume":"9","author":"Tiono","year":"2014","journal-title":"PloS One"},{"key":"B35","doi-asserted-by":"publisher","first-page":"5052","DOI":"10.1038\/s41598-018-23194-9","article-title":"Molecular camouflage of Plasmodium falciparum merozoites by binding of host vitronectin to P47 fragment of SERA5","volume":"8","author":"Tougan","year":"2018","journal-title":"Sci. Rep."},{"key":"B36","doi-asserted-by":"publisher","first-page":"322","DOI":"10.1016\/S0140-6736(19)31097-9","article-title":"Mapping the global prevalence, incidence, and mortality of Plasmodium falciparum, 2000-17: A spatial and temporal modelling study","volume":"394","author":"Weiss","year":"2019","journal-title":"Lancet"},{"key":"B37","unstructured":"World malaria report2021"},{"key":"B38","doi-asserted-by":"publisher","first-page":"395","DOI":"10.1111\/j.1558-5646.1965.tb01731.x","article-title":"The interpretation of population structure by F-statistics with special regard to systems of mating","volume":"19","author":"Wright","year":"1965","journal-title":"Evolution."},{"key":"B39","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0098460","article-title":"Protective epitopes of the Plasmodium falciparum SERA5 malaria vaccine reside in intrinsically unstructured n-terminal repetitive sequences","volume":"9","author":"Yagi","year":"2014","journal-title":"PloS One"},{"key":"B40","doi-asserted-by":"publisher","DOI":"10.1038\/srep34363","article-title":"Antibody titres and boosting after natural malaria infection in BK-SE36 vaccine responders during a follow-up study in Uganda","volume":"6","author":"Yagi","year":"2016","journal-title":"Sci. Rep."},{"key":"B41","doi-asserted-by":"publisher","first-page":"1072","DOI":"10.1016\/j.cell.2007.10.049","article-title":"Subcellular discharge of a serine protease mediates release of invasive malaria parasites from host erythrocytes","volume":"131","author":"Yeoh","year":"2007","journal-title":"Cell."}],"container-title":["Frontiers in Cellular and Infection Microbiology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fcimb.2022.1058081\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,12,16]],"date-time":"2022-12-16T17:00:26Z","timestamp":1671210026000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fcimb.2022.1058081\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,12,16]]},"references-count":41,"alternative-id":["10.3389\/fcimb.2022.1058081"],"URL":"https:\/\/doi.org\/10.3389\/fcimb.2022.1058081","relation":{},"ISSN":["2235-2988"],"issn-type":[{"type":"electronic","value":"2235-2988"}],"subject":[],"published":{"date-parts":[[2022,12,16]]},"article-number":"1058081"}}