{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T12:11:29Z","timestamp":1790856689976,"version":"4.1.0"},"reference-count":76,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2020,8,24]],"date-time":"2020-08-24T00:00:00Z","timestamp":1598227200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2020,8,24]],"date-time":"2020-08-24T00:00:00Z","timestamp":1598227200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Malar J"],"published-print":{"date-parts":[[2020,12]]},"abstract":"<jats:title>Abstract<\/jats:title><jats:p><jats:italic>Plasmodium vivax<\/jats:italic>has been largely neglected over the past century, despite a widespread recognition of its burden across region where it is endemic. The parasite invades reticulocytes, employing the interaction between<jats:italic>Plasmodium vivax<\/jats:italic>Duffy binding protein (PvDBP) and human Duffy antigen receptor for chemokines (DARC). However,<jats:italic>P. vivax<\/jats:italic>has now been observed in Duffy-negative individuals, presenting a potentially serious public health problem as the majority of African populations are Duffy-negative. Invasion of Duffy-negative reticulocytes is suggested to be through duplication of the PvDBP and a novel protein encoded by<jats:italic>P. vivax<\/jats:italic>erythrocyte binding protein (EBP) genes. The emergence and spread of specific<jats:italic>P. vivax<\/jats:italic>strains with ability to invade Duffy-negative reticulocytes has, therefore, drawn substantial attention and further complicated the epidemiology and public health implication of vivax malaria. Given the right environment and vectorial capacity for transmission coupled with the parasite\u2019s ability to invade Duffy-negative individuals,<jats:italic>P. vivax<\/jats:italic>could increase its epidemiological significance in Africa. In this review, authors present accruing knowledge on the paradigm shift in<jats:italic>P. vivax<\/jats:italic>invasion of Duffy-negative reticulocytes against the established mechanism of invading only Duffy-positive individuals and offer a perspective on the epidemiological diagnostic and public health implication in Africa.<\/jats:p>","DOI":"10.1186\/s12936-020-03372-9","type":"journal-article","created":{"date-parts":[[2020,8,24]],"date-time":"2020-08-24T10:03:45Z","timestamp":1598263425000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":31,"title":["The biology of unconventional invasion of Duffy-negative reticulocytes by Plasmodium vivax and its implication in malaria epidemiology and public health"],"prefix":"10.1186","volume":"19","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1216-8711","authenticated-orcid":false,"given":"Lemu","family":"Golassa","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lucas","family":"Amenga-Etego","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Eugenia","family":"Lo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alfred","family":"Amambua-Ngwa","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2020,8,24]]},"reference":[{"key":"3372_CR1","doi-asserted-by":"crossref","first-page":"e1007943","DOI":"10.1371\/journal.ppat.1007943","volume":"15","author":"H Kumar","year":"2019","unstructured":"Kumar H, Tolia NH. Getting in the structural biology of malaria invasion. PLoS Pathog. 2019;15:e1007943.","journal-title":"PLoS Pathog"},{"key":"3372_CR2","doi-asserted-by":"crossref","first-page":"332","DOI":"10.1016\/S0140-6736(19)31096-7","volume":"394","author":"KE Battle","year":"2019","unstructured":"Battle KE, Lucas TCD, Nguyen M, Howes RE, Nandi AK, Twohig KA, et al. Mapping the global endemicity and clinical burden of Plasmodium vivax, 2000\u201317: a spatial and temporal modelling study. Lancet. 2019;394:332\u201343.","journal-title":"Lancet"},{"key":"3372_CR3","doi-asserted-by":"crossref","first-page":"266","DOI":"10.1038\/ncomms1265","volume":"2","author":"RE Howes","year":"2011","unstructured":"Howes RE, Patil AP, Piel FB, Nyangiri OA, Kabaria CW, Gething PW, et al. The global distribution of the Duffy blood group. Nat Commun. 2011;2:266.","journal-title":"Nat Commun"},{"key":"3372_CR4","doi-asserted-by":"crossref","first-page":"a0255585","DOI":"10.1101\/cshperspect.a025585","volume":"7","author":"JH Adams","year":"2017","unstructured":"Adams JH, Mueller I. The biology of Plasmodium vivax. Cold Spring Harb Perspect Med. 2017;7:a0255585.","journal-title":"Cold Spring Harb Perspect Med"},{"key":"3372_CR5","doi-asserted-by":"crossref","first-page":"555","DOI":"10.1016\/S1473-3099(09)70177-X","volume":"9","author":"I Mueller","year":"2009","unstructured":"Mueller I, Galinski MR, Baird JK, Carlton JM, Kochar DK, Alonso PL, et al. Key gaps in the knowledge of Plasmodium vivax, a neglected human malaria parasite. Lancet Infect Dis. 2009;9:555\u201366.","journal-title":"Lancet Infect Dis"},{"key":"3372_CR6","doi-asserted-by":"crossref","first-page":"e774","DOI":"10.1371\/journal.pntd.0000774","volume":"4","author":"CA Guerra","year":"2010","unstructured":"Guerra CA, Howes RE, Patil AP, Gething PW, van Boeckel TP, Temperley WH, et al. The international limits and population at risk of Plasmodium vivax transmission in 2009. PLoS Negl Trop Dis. 2010;4:e774.","journal-title":"PLoS Negl Trop Dis"},{"key":"3372_CR7","doi-asserted-by":"crossref","first-page":"e103262","DOI":"10.1371\/journal.pone.0103262","volume":"9","author":"HGN Mbenda","year":"2014","unstructured":"Mbenda HGN, Das A. Molecular evidence of Plasmodium vivax mono and mixed malaria parasite infections in duffy-negative native Cameroonians. PLoS ONE. 2014;9:e103262.","journal-title":"PLoS ONE"},{"key":"3372_CR8","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1016\/B978-0-12-407826-0.00002-3","volume":"81","author":"PA Zimmerman","year":"2013","unstructured":"Zimmerman PA, Ferreira MU, Howes RE, Mercereau-Puijalon O. Red blood cell polymorphism and susceptibility to Plasmodium vivax. Adv Parasitol. 2013;81:27\u201376.","journal-title":"Adv Parasitol"},{"key":"3372_CR9","doi-asserted-by":"crossref","first-page":"420","DOI":"10.1016\/j.pt.2018.02.006","volume":"34","author":"K Gunalan","year":"2018","unstructured":"Gunalan K, Niangaly A, Thera MA, Doumbo OK, Miller LH. Plasmodium vivax infections of Duffy-negative erythrocytes: historically undetected or a recent adaptation? Trends Parasitol. 2018;34:420\u20139.","journal-title":"Trends Parasitol"},{"key":"3372_CR10","doi-asserted-by":"crossref","first-page":"97","DOI":"10.4269\/ajtmh.2001.64.97","volume":"64","author":"K Mendis","year":"2001","unstructured":"Mendis K, Sina BJB, Marchesini P, Carter R. The neglected burden of Plasmodium vivax malaria. Am J Trop Med Hyg. 2001;64:97\u2013106.","journal-title":"Am J Trop Med Hyg"},{"key":"3372_CR11","doi-asserted-by":"crossref","first-page":"636","DOI":"10.4269\/ajtmh.17-0461","volume":"97","author":"PA Zimmerman","year":"2017","unstructured":"Zimmerman PA. Plasmodium vivax infection in Duffy-negative people in Africa. Am J Trop Med Hyg. 2017;97:636\u20138.","journal-title":"Am J Trop Med Hyg"},{"key":"3372_CR12","doi-asserted-by":"crossref","first-page":"84","DOI":"10.1186\/s12936-015-0596-4","volume":"14","author":"E Lo","year":"2015","unstructured":"Lo E, Yewhalaw D, Zhong D, Zemene E, Degefa T, Tushune K, et al. Molecular epidemiology of Plasmodium vivax and Plasmodium falciparum malaria among Duffy-positive and Duffy-negative populations in Ethiopia. Malar J. 2015;14:84.","journal-title":"Malar J"},{"key":"3372_CR13","doi-asserted-by":"crossref","first-page":"959","DOI":"10.1038\/ng.3599","volume":"48","author":"RD Pearson","year":"2016","unstructured":"Pearson RD, Amato R, Auburn S, Miotto O, Almagro-Garcia J, Amaratunga C, et al. Genomic analysis of local variation and recent evolution in Plasmodium vivax. Nat Genet. 2016;48:959\u201364.","journal-title":"Nat Genet"},{"key":"3372_CR14","first-page":"413","volume":"56","author":"FB Livingstone","year":"1984","unstructured":"Livingstone FB. The Duffy blood groups, vivax malaria, and malaria selection in human populations: a review. Hum Biol. 1984;56:413\u201325.","journal-title":"Hum Biol"},{"key":"3372_CR15","doi-asserted-by":"crossref","first-page":"5967","DOI":"10.1073\/pnas.0912496107","volume":"107","author":"D Menard","year":"2010","unstructured":"Menard D, Barnadas C, Bouchier C, Henry-Halldin C, Gray LR, Ratsimbasoa A, et al. Plasmodium vivax clinical malaria is commonly observed in Duffy-negative Malagasy people. Proc Natl Acad Sci USA. 2010;107:5967\u201371.","journal-title":"Proc Natl Acad Sci USA"},{"key":"3372_CR16","doi-asserted-by":"crossref","first-page":"6271","DOI":"10.1073\/pnas.1606113113","volume":"113","author":"K Gunalan","year":"2016","unstructured":"Gunalan K, Lo E, Hostetler JB, Yewhalaw D, Mu J, Neafsey DE, et al. Role of Plasmodium vivax Duffy-binding protein 1 in invasion of Duffy-null Africans. Proc Natl Acad Sci USA. 2016;113:6271\u20136.","journal-title":"Proc Natl Acad Sci USA"},{"key":"3372_CR17","doi-asserted-by":"crossref","first-page":"744","DOI":"10.4269\/ajtmh.17-0254","volume":"97","author":"A Niangaly","year":"2017","unstructured":"Niangaly A, Gunalan K, Ouattara A, Coulibaly D, S\u00e1 JM, Adams M, et al. Plasmodium vivax infections over 3\u00a0years in Duffy blood group negative Malians in Bandiagara, Mali. Am J Trop Med Hyg. 2017;97:744\u201352.","journal-title":"Am J Trop Med Hyg"},{"key":"3372_CR18","doi-asserted-by":"crossref","first-page":"74","DOI":"10.1186\/s12936-017-1722-2","volume":"16","author":"G Russo","year":"2017","unstructured":"Russo G, Faggioni G, Paganotti GM, Djeunang Dongho GB, Pomponi A, De Santis R, et al. Molecular evidence of Plasmodium vivax infection in Duffy negative symptomatic individuals from Dschang, West Cameroon. Malar J. 2017;16:74.","journal-title":"Malar J"},{"key":"3372_CR19","doi-asserted-by":"crossref","first-page":"220","DOI":"10.1016\/j.pt.2009.02.003","volume":"25","author":"NM Anstey","year":"2009","unstructured":"Anstey NM, Russell B, Yeo TW, Price RN. The pathophysiology of vivax malaria. Trends Parasitol. 2009;25:220\u20137.","journal-title":"Trends Parasitol"},{"key":"3372_CR20","volume-title":"Current topics in malaria","author":"WE Villamil-G\u00f3mez","year":"2016","unstructured":"Villamil-G\u00f3mez WE. Severe and complicated malaria due to Plasmodium vivax. In: Rodriguez-Morales A, editor. Current topics in malaria. Rijeka: InTech Open Publ; 2016."},{"key":"3372_CR21","volume-title":"World malaria report 2016","author":"WHO","year":"2016","unstructured":"WHO. World malaria report 2016. Geneva: World Health Organization; 2016."},{"key":"3372_CR22","doi-asserted-by":"crossref","first-page":"401","DOI":"10.1093\/femsre\/fuz010","volume":"43","author":"P Ngotho","year":"2019","unstructured":"Ngotho P, Soares AB, Hentzschel F, Achcar F, Bertuccini L, Marti M. Revisiting gametocyte biology in malaria parasites. FEMS Microbiol. 2019;43:401\u201314.","journal-title":"FEMS Microbiol"},{"key":"3372_CR23","doi-asserted-by":"crossref","first-page":"4","DOI":"10.4269\/ajtmh.16-0160","volume":"95","author":"PL Olliaro","year":"2016","unstructured":"Olliaro PL, Barnwell JW, Barry A, Mendis K, Mueller I, Reeder JC, et al. Implications of Plasmodium vivax biology for control, elimination, and research. Am J Trop Med Hyg. 2016;95:4\u201314.","journal-title":"Am J Trop Med Hyg"},{"key":"3372_CR24","doi-asserted-by":"crossref","first-page":"297","DOI":"10.1186\/1475-2875-10-297","volume":"10","author":"NJ White","year":"2011","unstructured":"White NJ. Determinants of relapse periodicity in Plasmodium vivax malaria. Malar J. 2011;10:297.","journal-title":"Malar J"},{"key":"3372_CR25","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1186\/s12916-015-0302-y","volume":"13","author":"AM Siqueira","year":"2015","unstructured":"Siqueira AM, Lacerda MVG, Magalh\u00e3es BML, Mour\u00e3o MPG, Melo GC, Alexandre MAA, et al. Characterization of Plasmodium vivax-associated admissions to reference hospitals in Brazil and India. BMC Med. 2015;13:57.","journal-title":"BMC Med"},{"key":"3372_CR26","doi-asserted-by":"crossref","first-page":"1314","DOI":"10.1182\/blood-2014-08-596015","volume":"125","author":"B Malleret","year":"2015","unstructured":"Malleret B, Li A, Zhang R, Tan KSW, Suwanarusk R, Claser C, et al. Plasmodium vivax: restricted tropism and rapid remodeling of CD71-positive reticulocytes. Blood. 2015;125:1314\u201324.","journal-title":"Blood"},{"key":"3372_CR27","doi-asserted-by":"crossref","first-page":"1941","DOI":"10.1126\/science.8009226","volume":"264","author":"BKL Sim","year":"1994","unstructured":"Sim BKL, Chitnis CE, Wasniowska K, Hadley TJ, Miller LH. Receptor and ligand domains for invasion of erythrocytes by Plasmodium falciparum. Science. 1994;264:1941\u20134.","journal-title":"Science"},{"key":"3372_CR28","doi-asserted-by":"crossref","first-page":"e1003869","DOI":"10.1371\/journal.ppat.1003869","volume":"10","author":"JJD Batchelor","year":"2014","unstructured":"Batchelor JJD, Malpede BMB, Omattage NNS, DeKoster GT, Henzler-Wildman KA, Tolia NH. Red blood cell invasion by Plasmodium vivax: structural basis for DBP engagement of DARC. PLoS Pathog. 2014;10:e1003869.","journal-title":"PLoS Pathog"},{"key":"3372_CR29","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1126\/science.aan1078","volume":"359","author":"J Gruszczyk","year":"2018","unstructured":"Gruszczyk J, Kanjee U, Chan LJ, Menant S, Malleret B, Lim NTY, et al. Transferrin receptor 1 is a reticulocyte-specific receptor for Plasmodium vivax. Science. 2018;359:48\u201355.","journal-title":"Science"},{"key":"3372_CR30","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1097\/MOH.0000000000000219","volume":"23","author":"ND Salinas","year":"2016","unstructured":"Salinas ND, Tolia NH. Red cell receptors as access points for malaria infection. Curr Opin Hematol. 2016;23:215\u201323.","journal-title":"Curr Opin Hematol"},{"key":"3372_CR31","doi-asserted-by":"crossref","first-page":"1213","DOI":"10.1016\/0092-8674(92)90642-P","volume":"69","author":"MR Galinski","year":"1992","unstructured":"Galinski MR, Medina CC, Ingravallo P, Barnwell JW. A reticulocyte-binding protein complex of Plasmodium vivax merozoites. Cell. 1992;69:1213\u201326.","journal-title":"Cell"},{"key":"3372_CR32","doi-asserted-by":"crossref","first-page":"1441","DOI":"10.1182\/blood-2017-03-774364","volume":"130","author":"E Ovchynnikova","year":"2017","unstructured":"Ovchynnikova E, Aglialoro F, Bentlage AEH, Vidarsson G, Salinas ND, Von Lindern M, et al. DARC extracellular domain remodeling in maturating reticulocytes explains Plasmodium vivax tropism. Blood. 2017;130:1441\u20134.","journal-title":"Blood"},{"key":"3372_CR33","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.bbrc.2012.08.055","volume":"425","author":"J Li","year":"2012","unstructured":"Li J, Han ET. Dissection of the Plasmodium vivax reticulocyte binding-like proteins (PvRBPs). Biochem Biophys Res Commun. 2012;425:1\u20136.","journal-title":"Biochem Biophys Res Commun"},{"key":"3372_CR34","doi-asserted-by":"crossref","first-page":"302","DOI":"10.1056\/NEJM197608052950602","volume":"295","author":"L Miller","year":"1976","unstructured":"Miller L, Mason S, Clyde D, McGinniss M. The resistance factor to Plasmodium vivax in blacks. N Engl J Med. 1976;295:302\u20134.","journal-title":"N Engl J Med"},{"key":"3372_CR35","first-page":"93","volume":"16","author":"G H\u00f6her","year":"2018","unstructured":"H\u00f6her G, Fiegenbaum M, Almeida S. Molecular basis of the Duffy blood group system. Blood Transfus. 2018;16:93\u2013100.","journal-title":"Blood Transfus"},{"key":"3372_CR36","doi-asserted-by":"crossref","first-page":"2237","DOI":"10.1182\/blood.V92.7.2237","volume":"92","author":"N Parasol","year":"1998","unstructured":"Parasol N, Reid M, Rios M, Castilho L, Harari I, Kosower NS. A novel mutation in the coding sequence of the FY*B allele of the Duffy chemokine receptor gene is associated with an altered erythrocyte phenotype. Blood. 1998;92:2237\u201343.","journal-title":"Blood"},{"key":"3372_CR37","doi-asserted-by":"crossref","first-page":"224","DOI":"10.1038\/ng0695-224","volume":"10","author":"C Tournamille","year":"1995","unstructured":"Tournamille C, Colin Y, Cartron JP, Van Kim CL. Disruption of a GATA motif in the Duffy gene promoter abolishes erythroid gene expression in Duffy-negative individuals. Nat Genet. 1995;10:224\u20138.","journal-title":"Nat Genet"},{"key":"3372_CR38","doi-asserted-by":"crossref","first-page":"1311","DOI":"10.1084\/jem.181.4.1311","volume":"181","author":"SC Peiper","year":"1995","unstructured":"Peiper SC, Wang Z, Neote K, Martin AW, Showell HJ, Conklyn MJ, et al. The Duffy antigen\/receptor for chemoklnes (DARC) is expressed in endothelial cells of duffy negative individuals who lack the erythrocyte receptor. J Exp Med. 1995;181:1311\u20137.","journal-title":"J Exp Med"},{"key":"3372_CR39","doi-asserted-by":"crossref","first-page":"575","DOI":"10.4269\/ajtmh.2006.75.575","volume":"75","author":"JR Ryan","year":"2006","unstructured":"Ryan JR, Stoute JA, Amon J, Dunton RF, Mtalib R, Koros J, et al. Evidence for transmission of Plasmodium vivax among a Duffy antigen negative population in Western Kenya. Am J Trop Med Hyg. 2006;75:575\u201381.","journal-title":"Am J Trop Med Hyg"},{"key":"3372_CR40","doi-asserted-by":"crossref","first-page":"296","DOI":"10.1111\/vox.12273","volume":"109","author":"GH Lopez","year":"2015","unstructured":"Lopez GH, Morrison J, Condon JA, Wilson B, Martin JR, Liew YW, et al. Duffy blood group phenotype-genotype correlations using high-resolution melting analysis PCR and microarray reveal complex cases including a new null FY*A allele: the role for sequencing in genotyping algorithms. Vox Sang. 2015;109:296\u2013303.","journal-title":"Vox Sang"},{"key":"3372_CR41","doi-asserted-by":"crossref","first-page":"52","DOI":"10.1111\/vox.12185","volume":"108","author":"GH Lopez","year":"2015","unstructured":"Lopez GH, Condon JA, Wilson B, Martin JR, Liew YW, Flower RL, et al. A novel FY*A allele with the 265T and 298A SNPs formerly associated exclusively with the FY*B allele and weak Fyb antigen expression: implication for genotyping interpretative algorithms. Vox Sang. 2015;108:52\u20137.","journal-title":"Vox Sang"},{"key":"3372_CR42","doi-asserted-by":"crossref","first-page":"190","DOI":"10.1111\/j.1423-0410.2004.00554.x","volume":"87","author":"L Castilho","year":"2004","unstructured":"Castilho L, Rios M, Pellegrino J, Saad STO, Costa FF, Reid ME. A novel FY allele in Brazilians. Vox Sang. 2004;87:190\u20135.","journal-title":"Vox Sang"},{"key":"3372_CR43","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1016\/0166-6851(91)90228-X","volume":"44","author":"XX Fang","year":"1991","unstructured":"Fang XX, Kaslow DDC, Adams JJH, Miller LHL. Cloning of the Plasmodium vivax Duffy receptor. Mol Biochem Parasitol. 1991;44:125\u201332.","journal-title":"Mol Biochem Parasitol"},{"key":"3372_CR44","doi-asserted-by":"crossref","first-page":"e0005091","DOI":"10.1371\/journal.pntd.0005091","volume":"10","author":"JB Hostetler","year":"2016","unstructured":"Hostetler JB, Lo E, Kanjee U, Amaratunga C, Suon S, Sreng S, et al. Independent origin and global distribution of distinct Plasmodium vivax Duffy binding protein gene duplications. PLoS Negl Trop Dis. 2016;10:e0005091.","journal-title":"PLoS Negl Trop Dis"},{"key":"3372_CR45","doi-asserted-by":"crossref","first-page":"953","DOI":"10.1038\/ng.3588","volume":"48","author":"DN Hupalo","year":"2016","unstructured":"Hupalo DN, Luo Z, Melnikov A, Sutton PL, Rogov P, Escalante A, et al. Population genomics studies identify signatures of global dispersal and drug resistance in Plasmodium vivax. Nat Genet. 2016;48:953\u20138.","journal-title":"Nat Genet"},{"key":"3372_CR46","doi-asserted-by":"crossref","first-page":"6270","DOI":"10.1128\/AAC.03026-14","volume":"58","author":"C Amaratunga","year":"2014","unstructured":"Amaratunga C, Sreng S, Mao S, Tullo GS, Anderson JM, Chuor CM, et al. Chloroquine remains effective for treating Plasmodium vivax malaria in Pursat Province, western Cambodia. Antimicrob Agents Chemother. 2014;58:6270\u20132.","journal-title":"Antimicrob Agents Chemother"},{"key":"3372_CR47","doi-asserted-by":"crossref","first-page":"e2489","DOI":"10.1371\/journal.pntd.0002489","volume":"7","author":"D Menard","year":"2013","unstructured":"Menard D, Chan ER, Benedet C, Ratsimbasoa A, Kim S, Chim P, et al. Whole genome fequencing of field isolates reveals a common duplication of the Duffy binding protein gene in Malagasy Plasmodium vivax strains. PLoS Negl Trop Dis. 2013;7:e2489.","journal-title":"PLoS Negl Trop Dis"},{"key":"3372_CR48","doi-asserted-by":"crossref","first-page":"467","DOI":"10.1016\/j.ijpara.2014.03.002","volume":"44","author":"HM Abkallo","year":"2014","unstructured":"Abkallo HM, Liu W, Hokama S, Ferreira PE, Nakazawa S, Maeno Y, et al. DNA from pre-erythrocytic stage malaria parasites is detectable by PCR in the faeces and blood of hosts. Int J Parasitol. 2014;44:467\u201373.","journal-title":"Int J Parasitol"},{"key":"3372_CR49","doi-asserted-by":"crossref","first-page":"e8096","DOI":"10.1073\/pnas.1608828113","volume":"113","author":"CM Parobek","year":"2016","unstructured":"Parobek CM, Lin JT, Saunders DL, Barnett EJ, Lon C, Lanteri CA, et al. Selective sweep suggests transcriptional regulation may underlie Plasmodium vivax resilience to malaria control measures in Cambodia. Proc Natl Acad Sci USA. 2016;113:e8096\u2013105.","journal-title":"Proc Natl Acad Sci USA"},{"key":"3372_CR50","doi-asserted-by":"crossref","first-page":"953","DOI":"10.1038\/s41467-020-14574-9","volume":"11","author":"J Popovici","year":"2020","unstructured":"Popovici J, Roesch C, Carias LL, Khim N, Kim S, Vantaux A, et al. Amplification of Duffy binding protein-encoding gene allows Plasmodium vivax to evade host anti-DBP humoral immunity. Nat Commun. 2020;11:953.","journal-title":"Nat Commun"},{"key":"3372_CR51","doi-asserted-by":"crossref","first-page":"e0007222","DOI":"10.1371\/journal.pntd.0007222","volume":"13","author":"E Lo","year":"2019","unstructured":"Lo E, Hostetler JB, Yewhalaw D, Pearson RD, Hamid MMA, Gunalan K, et al. Frequent expansion of Plasmodium vivax Duffy binding protein in Ethiopia and its epidemiological significance. PLoS Negl Trop Dis. 2019;13:e0007222.","journal-title":"PLoS Negl Trop Dis"},{"key":"3372_CR52","doi-asserted-by":"crossref","first-page":"e0006555","DOI":"10.1371\/journal.pntd.0006555","volume":"12","author":"C Roesch","year":"2018","unstructured":"Roesch C, Popovici J, Bin S, Run V, Kim S, Ramboarina S, et al. Genetic diversity in two Plasmodium vivax protein ligands for reticulocyte invasion. PLoS Negl Trop Dis. 2018;12:e0006555.","journal-title":"PLoS Negl Trop Dis"},{"key":"3372_CR53","doi-asserted-by":"crossref","first-page":"e0004222","DOI":"10.1371\/journal.pntd.0004222","volume":"9","author":"RE Howes","year":"2015","unstructured":"Howes RE, Reiner RC, Battle KE, Longbottom J, Mappin B, Ordanovich D, et al. Plasmodium vivax transmission in Africa. PLoS Negl Trop Dis. 2015;9:e0004222.","journal-title":"PLoS Negl Trop Dis"},{"key":"3372_CR54","doi-asserted-by":"crossref","first-page":"e0007140","DOI":"10.1371\/journal.pntd.0007140","volume":"13","author":"KA Twohig","year":"2019","unstructured":"Twohig KA, Pfeffer DA, Baird JK, Price RN, Zimmerman PA, Hay SI, et al. Growing evidence of Plasmodium vivax across malaria-endemic Africa. PLoS Negl Trop Dis. 2019;13:e0007140.","journal-title":"PLoS Negl Trop Dis"},{"key":"3372_CR55","doi-asserted-by":"crossref","first-page":"371","DOI":"10.2307\/3274317","volume":"44","author":"RS Bray","year":"1958","unstructured":"Bray RS. The susceptibility of Liberians to the Madagascar strain of Plasmodium vivax. J Parasitol. 1958;44:371\u20133.","journal-title":"J Parasitol"},{"key":"3372_CR56","doi-asserted-by":"crossref","first-page":"13851","DOI":"10.1038\/s41598-018-32254-z","volume":"8","author":"FS Kano","year":"2018","unstructured":"Kano FS, de Souza AM, de Menezes Torres L, Costa MA, Souza-Silva FA, Sanchez BAM, et al. Susceptibility to Plasmodium vivax malaria associated with DARC (Duffy antigen) polymorphisms is influenced by the time of exposure to malaria. Sci Rep. 2018;8:13851.","journal-title":"Sci Rep"},{"key":"3372_CR57","doi-asserted-by":"crossref","first-page":"430","DOI":"10.1186\/1475-2875-11-430","volume":"11","author":"TAA Carvalho","year":"2012","unstructured":"Carvalho TAA, Queiroz MG, Cardoso GL, Diniz IG, Silva ANLM, Pinto AYN, et al. Plasmodium vivax infection in Anaj\u00e1s, State of Par\u00e1: no differential resistance profile among Duffy-negative and Duffy-positive individuals. Malar J. 2012;11:430.","journal-title":"Malar J"},{"key":"3372_CR58","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1186\/s12936-019-2918-4","volume":"18","author":"RK Abou-Ali","year":"2019","unstructured":"Abou-Ali RK, Dhyani A, Ter\u00e7o AL, Toro DM, Gomes KS, Tezza LC, et al. Impact of Duffy polymorphisms on parasite density in Brazilian Amazonian patients infected by Plasmodium vivax. Malar J. 2019;18:289.","journal-title":"Malar J"},{"key":"3372_CR59","doi-asserted-by":"crossref","first-page":"20113","DOI":"10.1073\/pnas.1109621108","volume":"108","author":"CL King","year":"2011","unstructured":"King CL, Adams JH, Xianli J, Grimberg BT, McHenry AM, Greenberg LJ, et al. Fy a\/Fy b antigen polymorphism in human erythrocyte duffy antigen affects susceptibility to Plasmodium vivax malaria. Proc Natl Acad Sci USA. 2011;108:20113\u20138.","journal-title":"Proc Natl Acad Sci USA"},{"key":"3372_CR60","doi-asserted-by":"crossref","first-page":"15","DOI":"10.21307\/immunohematology-2019-387","volume":"21","author":"IJ Woolley","year":"2005","unstructured":"Woolley IJ, Wood EM, Sramkoski RM, Zimmerman PA, Miller PA, Kazura JW. Expression of Duffy antigen receptor for chemokines during reticulocyte maturation: using a CD71 flow cytometric technique to identify reticulocytes. Immunohematology. 2005;21:15\u201320.","journal-title":"Immunohematology"},{"key":"3372_CR61","doi-asserted-by":"crossref","first-page":"2147","DOI":"10.1182\/blood.V92.6.2147","volume":"92","author":"C Tournamille","year":"1998","unstructured":"Tournamille C, Le C, Kim V, Gane P, Le Pennec PY, Roubinet F, et al. Arg89Cys substitution results in very low membrane expression of the Duffy antigen\/receptor for chemokines in Fy(x) individuals. Blood. 1998;92:2147\u201356.","journal-title":"Blood"},{"key":"3372_CR62","doi-asserted-by":"crossref","first-page":"301","DOI":"10.1186\/s12936-018-2456-5","volume":"17","author":"M Berm\u00fadez","year":"2018","unstructured":"Berm\u00fadez M, Moreno-P\u00e9rez DA, Ar\u00e9valo-Pinz\u00f3n G, Curtidor H, Patarroyo MA. Plasmodium vivax in vitro continuous culture: the spoke in the wheel. Malar J. 2018;17:301.","journal-title":"Malar J"},{"key":"3372_CR63","doi-asserted-by":"crossref","first-page":"1089","DOI":"10.1128\/IAI.01036-09","volume":"78","author":"P Chootong","year":"2010","unstructured":"Chootong P, Ntumngia FB, VanBuskirk KM, Xainli J, Cole-Tobian JLJ, Campbell CO, et al. Mapping epitopes of the Plasmodium vivax Duffy binding protein with naturally acquired inhibitory antibodies. Infect Immun. 2010;78:1089\u201395.","journal-title":"Infect Immun"},{"key":"3372_CR64","doi-asserted-by":"crossref","first-page":"e2569","DOI":"10.1371\/journal.pntd.0002569","volume":"7","author":"J Hester","year":"2013","unstructured":"Hester J, Chan ER, Menard D, Mercereau-Puijalon O, Barnwell J, Zimmerman PA, et al. De Novo assembly of a field isolate genome reveals novel Plasmodium vivax erythrocyte invasion genes. PLoS Negl Trop Dis. 2013;7:e2569.","journal-title":"PLoS Negl Trop Dis"},{"key":"3372_CR65","doi-asserted-by":"crossref","first-page":"e28673","DOI":"10.7554\/eLife.28673","volume":"6","author":"CT Fran\u00e7a","year":"2017","unstructured":"Fran\u00e7a CT, White MT, He WQ, Hostetler JB, Brewster J, Frato G, et al. Identification of highly-protective combinations of Plasmodium vivax recombinant proteins for vaccine development. Elife. 2017;6:e28673.","journal-title":"Elife"},{"key":"3372_CR66","doi-asserted-by":"crossref","first-page":"1558","DOI":"10.1093\/infdis\/jix170","volume":"215","author":"ED Gupta","year":"2017","unstructured":"Gupta ED, Anand G, Singh H, Chaddha K, Bharti PK, Singh N, et al. Naturally acquired human antibodies against reticulocyte-binding domains of Plasmodium vivax proteins, PvRBP2c and PvRBP1a, exhibit binding-inhibitory activity. J Infect Dis. 2017;215:1558\u201368.","journal-title":"J Infect Dis"},{"key":"3372_CR67","doi-asserted-by":"crossref","first-page":"e76062","DOI":"10.1371\/journal.pone.0076062","volume":"8","author":"B Malleret","year":"2013","unstructured":"Malleret B, Xu F, Mohandas N, Suwanarusk R, Chu C, Leite JA, et al. Significant biochemical, biophysical and metabolic diversity in circulating human cord blood reticulocytes. PLoS ONE. 2013;8:e76062.","journal-title":"PLoS ONE"},{"key":"3372_CR68","doi-asserted-by":"crossref","first-page":"3906","DOI":"10.1038\/s41598-019-40321-2","volume":"9","author":"JH Han","year":"2019","unstructured":"Han JH, Cheng Y, Muh F, Ahmed MA, Cho JS, Nyunt MH, et al. Inhibition of parasite invasion by monoclonal antibody against epidermal growth factor-like domain of Plasmodium vivax merozoite surface protein 1 paralog. Sci Rep. 2019;9:3906.","journal-title":"Sci Rep"},{"key":"3372_CR69","doi-asserted-by":"crossref","first-page":"e00239-18","DOI":"10.1128\/IAI.00239-18","volume":"86","author":"JH Han","year":"2018","unstructured":"Han JH, Cho JS, Cheng Y, Muh F, Yoo WG, Russell B, et al. Plasmodium vivax merozoite surface protein 1 paralog as a mediator of parasite adherence to reticulocytes. Infect Immun. 2018;86:e00239-18.","journal-title":"Infect Immun"},{"key":"3372_CR70","doi-asserted-by":"crossref","first-page":"35581","DOI":"10.1038\/srep35581","volume":"6","author":"Y Cheng","year":"2016","unstructured":"Cheng Y, Lu F, Wang B, Li J, Han JH, Ito D, et al. Plasmodium vivax GPI-anchored micronemal antigen (PvGAMA) binds human erythrocytes independent of Duffy antigen status. Sci Rep. 2016;6:35581.","journal-title":"Sci Rep"},{"key":"3372_CR71","doi-asserted-by":"crossref","first-page":"e1261-16","DOI":"10.1128\/mBio.01261-16","volume":"7","author":"FB Ntumngia","year":"2016","unstructured":"Ntumngia FB, Thomson-Luque R, de Menezes Torres L, Gunalan K, Carvalho LH, Adams JH. A novel erythrocyte binding protein of Plasmodium vivax suggests an alternate invasion pathway into duffy-positive reticulocytes. MBio. 2016;7:e1261-16.","journal-title":"MBio"},{"key":"3372_CR72","doi-asserted-by":"crossref","first-page":"1091","DOI":"10.1016\/j.ijpara.2012.08.005","volume":"42","author":"R Culleton","year":"2012","unstructured":"Culleton R, Carter R. African Plasmodium vivax: distribution and origins. Int J Parasitol. 2012;42:1091\u20137.","journal-title":"Int J Parasitol"},{"key":"3372_CR73","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1016\/j.ijpara.2016.05.008","volume":"47","author":"DE Loy","year":"2017","unstructured":"Loy DE, Liu W, Li Y, Learn GH, Plenderleith LJ, Sundararaman SA, et al. Out of Africa: origins and evolution of the human malaria parasites Plasmodium falciparum and Plasmodium vivax. Int J Parasitol. 2017;47:87\u201397.","journal-title":"Int J Parasitol"},{"key":"3372_CR74","doi-asserted-by":"crossref","first-page":"405","DOI":"10.1186\/1475-2875-12-405","volume":"12","author":"L Canier","year":"2013","unstructured":"Canier L, Khim N, Kim S, Sluydts V, Heng S, Dourng D, et al. An innovative tool for moving malaria PCR detection of parasite reservoir into the field. Malar J. 2013;12:405.","journal-title":"Malar J"},{"key":"3372_CR75","doi-asserted-by":"crossref","first-page":"8123","DOI":"10.1073\/pnas.1306004110","volume":"110","author":"F Prugnolle","year":"2013","unstructured":"Prugnolle F, Rougeron V, Becquart P, Berry A, Makanga B, Rahola N, et al. Diversity, host switching and evolution of Plasmodium vivax infecting african great apes. Proc Natl Acad Sci USA. 2013;110:8123\u20138.","journal-title":"Proc Natl Acad Sci USA"},{"key":"3372_CR76","doi-asserted-by":"crossref","first-page":"e1001642","DOI":"10.1371\/journal.pmed.1001642","volume":"11","author":"J Cao","year":"2014","unstructured":"Cao J, Sturrock HJW, Cotter C, Zhou S, Zhou H, Liu Y, et al. Communicating and monitoring surveillance and response activities for malaria elimination: china\u2019s \u201c1-3-7\u201d strategy. PLoS Med. 2014;11:e1001642.","journal-title":"PLoS Med"}],"container-title":["Malaria Journal"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12936-020-03372-9.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s12936-020-03372-9\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12936-020-03372-9.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,11,9]],"date-time":"2022-11-09T16:38:25Z","timestamp":1668011905000},"score":1,"resource":{"primary":{"URL":"https:\/\/malariajournal.biomedcentral.com\/articles\/10.1186\/s12936-020-03372-9"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,8,24]]},"references-count":76,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2020,12]]}},"alternative-id":["3372"],"URL":"https:\/\/doi.org\/10.1186\/s12936-020-03372-9","relation":{},"ISSN":["1475-2875"],"issn-type":[{"value":"1475-2875","type":"electronic"}],"subject":[],"published":{"date-parts":[[2020,8,24]]},"assertion":[{"value":"27 May 2020","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"10 August 2020","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"24 August 2020","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Not applicable.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval and consent to participate"}},{"value":"Not applicable.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"The authors declare that they have no competing interests.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"299"}}