{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,16]],"date-time":"2026-09-16T07:21:28Z","timestamp":1789543288974,"version":"build-2803163510"},"reference-count":36,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2019,10,2]],"date-time":"2019-10-02T00:00:00Z","timestamp":1569974400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2019,10,2]],"date-time":"2019-10-02T00:00:00Z","timestamp":1569974400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"publisher","award":["RO3055\/2-1"],"award-info":[{"award-number":["RO3055\/2-1"]}],"id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"publisher","award":["KN1097\/3-1"],"award-info":[{"award-number":["KN1097\/3-1"]}],"id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"publisher","award":["KN1097\/4-1"],"award-info":[{"award-number":["KN1097\/4-1"]}],"id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001824","name":"Grantov\u00e1 Agentura \u010cesk\u00e9 Republiky","doi-asserted-by":"publisher","award":["GA17-25455S"],"award-info":[{"award-number":["GA17-25455S"]}],"id":[{"id":"10.13039\/501100001824","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001824","name":"Grantov\u00e1 Agentura \u010cesk\u00e9 Republiky","doi-asserted-by":"publisher","award":["GC18-23521J"],"award-info":[{"award-number":["GC18-23521J"]}],"id":[{"id":"10.13039\/501100001824","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title><jats:p>In our most recent study, we found that in Tanzania infection with <jats:italic>Treponema pallidum<\/jats:italic> (<jats:italic>TP<\/jats:italic>) subsp. <jats:italic>pertenue<\/jats:italic> (<jats:italic>TPE<\/jats:italic>) is present in four different monkey species. In order to gain information on the diversity and epidemiological spread of the infection in Tanzanian nonhuman primates (NHP), we identified two suitable candidate genes for multi-locus sequence typing (MLST). We demonstrate the functionality of the MLST system in invasively and non-invasively collected samples. While we were not able to demonstrate frequent interspecies transmission of <jats:italic>TPE<\/jats:italic> in Tanzanian monkeys, our results show a clustering of <jats:italic>TPE<\/jats:italic> strains according to geography and not host species, which is suggestive for rare transmission events between different NHP species. In addition to the geographic stability, we describe the relative temporal stability of the strains infecting NHPs and identified multi-strain infection. Differences between <jats:italic>TPE<\/jats:italic> strains of NHP and human origin are highlighted. Our results show that antibiotic resistance does not occur in Tanzanian <jats:italic>TPE<\/jats:italic> strains of NHP origin.<\/jats:p>","DOI":"10.1038\/s41598-019-50779-9","type":"journal-article","created":{"date-parts":[[2019,10,2]],"date-time":"2019-10-02T10:06:02Z","timestamp":1570010762000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":21,"title":["Strain diversity of Treponema pallidum subsp. pertenue suggests rare interspecies transmission in African nonhuman primates"],"prefix":"10.1038","volume":"9","author":[{"given":"Idrissa S.","family":"Chuma","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0190-4266","authenticated-orcid":false,"given":"Christian","family":"Roos","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anagaw","family":"Atickem","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Torsten","family":"Bohm","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"D.","family":"Anthony Collins","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Linda","family":"Grillov\u00e1","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Luisa K.","family":"Hallmaier-Wacker","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rudovick R.","family":"Kazwala","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Julius D.","family":"Keyyu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Simone","family":"L\u00fcert","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ulrich","family":"Maloueki","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3076-4840","authenticated-orcid":false,"given":"Jan","family":"Oppelt","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kl\u00e1ra J.","family":"Petr\u017eelkov\u00e1","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alexander","family":"Piel","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fiona A.","family":"Stewart","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"David","family":"\u0160majs","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5744-4946","authenticated-orcid":false,"given":"Sascha","family":"Knauf","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2019,10,2]]},"reference":[{"key":"50779_CR1","doi-asserted-by":"publisher","DOI":"10.1038\/s41426-018-0156-4","volume":"7","author":"S Knauf","year":"2018","unstructured":"Knauf, S. et al. Nonhuman primates across sub-Saharan Africa are infected with the yaws bacterium Treponema pallidum subsp. pertenue. Emerg Microbes Infect 7, 157 (2018).","journal-title":"Emerg Microbes Infect"},{"key":"50779_CR2","doi-asserted-by":"publisher","first-page":"1002","DOI":"10.3201\/eid2406.180037","volume":"24","author":"IS Chuma","year":"2018","unstructured":"Chuma, I. S. et al. Widespread Treponema pallidum infection in nonhuman primates, Tanzania. Emerg Infect Dis 24, 1002\u20131009 (2018).","journal-title":"Emerg Infect Dis"},{"key":"50779_CR3","doi-asserted-by":"publisher","DOI":"10.1038\/emi.2017.65","volume":"6","author":"LK Hallmaier-Wacker","year":"2017","unstructured":"Hallmaier-Wacker, L. K., Munster, V. J. & Knauf, S. Disease reservoirs: from conceptual frameworks to applicable criteria. Emerg Microbes Infect 6, e79 (2017).","journal-title":"Emerg Microbes Infect"},{"key":"50779_CR4","doi-asserted-by":"publisher","first-page":"2058","DOI":"10.3201\/eid1912.130863","volume":"19","author":"S Knauf","year":"2013","unstructured":"Knauf, S., Liu, H. & Harper, K. N. Treponemal infection in nonhuman primates as possible reservoir for human yaws. Emerg Infect Dis 19, 2058\u201360 (2013).","journal-title":"Emerg Infect Dis"},{"key":"50779_CR5","doi-asserted-by":"publisher","first-page":"e1273","DOI":"10.1371\/journal.pntd.0001273","volume":"5","author":"R Peng","year":"2011","unstructured":"Peng, R. et al. Molecular typing of Treponema pallidum: a systematic review and meta-analysis. PLoS Negl Trop Dis 5, e1273 (2011).","journal-title":"PLoS Negl Trop Dis"},{"key":"50779_CR6","doi-asserted-by":"publisher","first-page":"202","DOI":"10.2340\/00015555-2190","volume":"96","author":"K Salado-Rasmussen","year":"2016","unstructured":"Salado-Rasmussen, K. et al. Molecular typing of Treponema pallidum in Denmark: A nationwide study of syphilis. Acta Derm Venereol 96, 202\u20136 (2016).","journal-title":"Acta Derm Venereol"},{"key":"50779_CR7","doi-asserted-by":"publisher","first-page":"641","DOI":"10.1097\/OLQ.0000000000000006","volume":"40","author":"P Grange","year":"2013","unstructured":"Grange, P. et al. Molecular subtyping of Treponema pallidum in Paris, France. Sex Transm Dis 40, 641\u20134 (2013).","journal-title":"Sex Transm Dis"},{"key":"50779_CR8","doi-asserted-by":"publisher","first-page":"3743","DOI":"10.1128\/JCM.43.8.3743-3746.2005","volume":"43","author":"V Pope","year":"2005","unstructured":"Pope, V. et al. Molecular subtyping of Treponema pallidum from North and South Carolina. J Clin Microbiol 43, 3743\u20133746 (2005).","journal-title":"J Clin Microbiol"},{"key":"50779_CR9","doi-asserted-by":"publisher","first-page":"408","DOI":"10.1097\/00007435-199809000-00004","volume":"25","author":"A Pillay","year":"1998","unstructured":"Pillay, A. et al. Molecular subtyping of Treponema pallidum subspecies pallidum. Sex Transm Dis 25, 408\u2013414 (1998).","journal-title":"Sex Transm Dis"},{"key":"50779_CR10","doi-asserted-by":"publisher","first-page":"3693","DOI":"10.1128\/JCM.01292-14","volume":"52","author":"L Grillov\u00e1","year":"2014","unstructured":"Grillov\u00e1, L. et al. Molecular typing of Treponema pallidum in the Czech Republic during 2011 to 2013: increased prevalence of identified genotypes and of isolates with macrolide resistance. J Clin Microbiol 52, 3693\u20133700 (2014).","journal-title":"J Clin Microbiol"},{"key":"50779_CR11","doi-asserted-by":"publisher","first-page":"e0006113","DOI":"10.1371\/journal.pntd.0006113","volume":"11","author":"C Godornes","year":"2017","unstructured":"Godornes, C., Giacani, L., Barry, A., Mitj\u00e0, O. & Lukehart, S. Development of a Multilocus sequence typing (MLST) scheme for Treponema pallidum subsp. pertenue: Application to yaws in Lihir Island, Papua New Guinea. PLoS Neglect Trop Dis 11, e0006113 (2017).","journal-title":"PLoS Neglect Trop Dis"},{"key":"50779_CR12","doi-asserted-by":"publisher","first-page":"e0203632","DOI":"10.1371\/journal.pone.0203632","volume":"13","author":"SS Katz","year":"2018","unstructured":"Katz, S. S. et al. Molecular strain typing of the yaws pathogen, Treponema pallidum subspecies pertenue. PLoS ONE 13, e0203632 (2018).","journal-title":"PLoS ONE"},{"key":"50779_CR13","doi-asserted-by":"publisher","first-page":"292","DOI":"10.1177\/0300985811402839","volume":"49","author":"S Knauf","year":"2011","unstructured":"Knauf, S. et al. Treponema infection associated with genital ulceration in wild baboons. Vet Pathol 49, 292\u2013303 (2011).","journal-title":"Vet Pathol"},{"key":"50779_CR14","doi-asserted-by":"publisher","first-page":"832","DOI":"10.1099\/jmm.0.007542-0","volume":"58","author":"P Mat\u011bjkov\u00e1","year":"2009","unstructured":"Mat\u011bjkov\u00e1, P. et al. Macrolide treatment failure in a case of secondary syphilis: a novel A2059G mutation in the 23S rRNA gene of Treponema pallidum subsp. pallidum. J Med Microbiol 58, 832\u2013836 (2009).","journal-title":"J Med Microbiol"},{"key":"50779_CR15","doi-asserted-by":"publisher","first-page":"908","DOI":"10.1128\/JCM.02770-12","volume":"51","author":"C Chen","year":"2013","unstructured":"Chen, C. et al. Detection of the A2058G and A2059G 23S rRNA gene point mutations associated with azithromycin resistance in Treponema pallidum by use of a TaqMan real-time multiplex PCR assay. J Clin Microbiol 51, 908\u2013913 (2013).","journal-title":"J Clin Microbiol"},{"key":"50779_CR16","doi-asserted-by":"publisher","first-page":"806","DOI":"10.1128\/AAC.44.3.806-807.2000","volume":"44","author":"L Stamm","year":"2000","unstructured":"Stamm, L. & Bergen, H. A point mutation associated with bacterial macrolide resistance is present in both 23S rRNA genes of an erythromycin-resistant Treponema pallidum clinical isolate. Antimicrob Agents and Chemother 44, 806\u2013807 (2000).","journal-title":"Antimicrob Agents and Chemother"},{"key":"50779_CR17","doi-asserted-by":"publisher","first-page":"154","DOI":"10.1056\/NEJMoa040216","volume":"351","author":"S Lukehart","year":"2014","unstructured":"Lukehart, S. et al. Macrolide resistance in Treponema pallidum in the United States and Ireland. N Engl J Med 351, 154\u2013158 (2014).","journal-title":"N Engl J Med"},{"key":"50779_CR18","doi-asserted-by":"publisher","first-page":"37","DOI":"10.1093\/oxfordjournals.molbev.a026036","volume":"16","author":"H Bandelt","year":"1999","unstructured":"Bandelt, H., Forster, P. & Rohl, A. Median-joining networks for inferring intraspecific phylogenies. Mol Biol and Evol 16, 37\u201348 (1999).","journal-title":"Mol Biol and Evol"},{"key":"50779_CR19","doi-asserted-by":"publisher","first-page":"221","DOI":"10.1093\/molbev\/msp259","volume":"27","author":"M Gouy","year":"2010","unstructured":"Gouy, M., Guindon, S. & Gascuel, O. SeaView version 4: A multiplatform graphical user interface for sequence alignment and phylogenetic tree building. Mol Biol Evol 27, 221\u2013224 (2010).","journal-title":"Mol Biol Evol"},{"key":"50779_CR20","doi-asserted-by":"publisher","first-page":"268","DOI":"10.1093\/molbev\/msu300","volume":"32","author":"L-TT Nguyen","year":"2015","unstructured":"Nguyen, L.-T. T., Schmidt, H. A., Haeseler, A. & von Minh, B. Q. IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Mol Biol and Evol 32, 268\u2013274 (2015).","journal-title":"Mol Biol and Evol"},{"key":"50779_CR21","doi-asserted-by":"publisher","first-page":"e0005894","DOI":"10.1371\/journal.pntd.0005894","volume":"11","author":"M Strouhal","year":"2017","unstructured":"Strouhal, M. et al. Complete genome sequences of two strains of Treponema pallidum subsp. pertenue from Ghana, Africa: Identical genome sequences in samples isolated more than 7 years apart. PLoS Negl Trop Dis 11, e0005894 (2017).","journal-title":"PLoS Negl Trop Dis"},{"key":"50779_CR22","doi-asserted-by":"publisher","first-page":"e0200773","DOI":"10.1371\/journal.pone.0200773","volume":"13","author":"L Grillov\u00e1","year":"2018","unstructured":"Grillov\u00e1, L. et al. Molecular characterization of Treponema pallidum subsp. pallidum in Switzerland and France with a new multilocus sequence typing scheme. PloS ONE 13, e0200773 (2018).","journal-title":"PloS ONE"},{"key":"50779_CR23","doi-asserted-by":"publisher","first-page":"e0202619","DOI":"10.1371\/journal.pone.0202619","volume":"13","author":"L Grillov\u00e1","year":"2018","unstructured":"Grillov\u00e1, L. et al. Sequencing of Treponema pallidum subsp. pallidum from isolate UZ1974 using anti-treponemal antibodies enrichment: First complete whole genome sequence obtained directly from human clinical material. PLoS ONE 13, e0202619 (2018).","journal-title":"PLoS ONE"},{"key":"50779_CR24","doi-asserted-by":"publisher","first-page":"85","DOI":"10.1016\/j.ebiom.2016.07.033","volume":"11","author":"S Knauf","year":"2016","unstructured":"Knauf, S. et al. Isolation of Treponema DNA from necrophagous flies in a natural ecosystem. EBioMed 11, 85\u201390 (2016).","journal-title":"EBioMed"},{"key":"50779_CR25","doi-asserted-by":"crossref","unstructured":"Zinner, D., Chuma, I. S., Knauf, S. & Roos, C. Inverted intergeneric introgression between critically endangered kipunjis and yellow baboons in two disjunct populations. Biol Lett, 14 (2018).","DOI":"10.1098\/rsbl.2017.0729"},{"key":"50779_CR26","unstructured":"Collins, D., Sindimwo, A. & Wallis, P. Reproductive disease of olive baboons (Papio anubis) of Gombe National Park: outbreak, time-course, and attempts to limit recurrence. In: Climate change: challenges and opportunities towards sustainable wildlife conservation and development. 236\u2013248 (Tanzania Wildlife Research Institute, 2011)."},{"key":"50779_CR27","doi-asserted-by":"publisher","first-page":"1599","DOI":"10.1016\/S0140-6736(18)30204-6","volume":"391","author":"O Mitj\u00e0","year":"2018","unstructured":"Mitj\u00e0, O. et al. Re-emergence of yaws after single mass azithromycin treatment followed by targeted treatment: a longitudinal study. Lancet 391, 1599\u20131607 (2018).","journal-title":"Lancet"},{"key":"50779_CR28","doi-asserted-by":"publisher","first-page":"1555","DOI":"10.1016\/S0140-6736(18)30205-8","volume":"391","author":"D \u0160majs","year":"2018","unstructured":"\u0160majs, D. & Posp\u00ed\u0161ilov\u00e1, P. Macrolide resistance in yaws. Lancet 391, 1555\u20131556 (2018).","journal-title":"Lancet"},{"key":"50779_CR29","first-page":"3","volume":"35","author":"T Turner","year":"1957","unstructured":"Turner, T. & Hollander, D. Biology of the treponematoses based on studies carried out at the International Treponematosis Laboratory Center of the Johns Hopkins University under the auspices of the World Health Organization. Monogr Ser World Health Organ 35, 3\u2013266 (1957).","journal-title":"Monogr Ser World Health Organ"},{"key":"50779_CR30","doi-asserted-by":"publisher","first-page":"e1832","DOI":"10.1371\/journal.pntd.0001832","volume":"6","author":"H P\u011btro\u0161ov\u00e1","year":"2012","unstructured":"P\u011btro\u0161ov\u00e1, H. et al. Whole genome sequence of Treponema pallidum ssp. pallidum, strain Mexico A, suggests recombination between yaws and syphilis strains. PLoS Negl Trop Dis 6, e1832 (2012).","journal-title":"PLoS Negl Trop Dis"},{"key":"50779_CR31","doi-asserted-by":"publisher","first-page":"517","DOI":"10.1097\/OLQ.0000000000000474","volume":"43","author":"P Grange","year":"2016","unstructured":"Grange, P. et al. Treponema pallidum 11qj subtype may correspond to a Treponema pallidum subsp. endemicum strain. Sex Transm Dis 43, 517\u2013518 (2016).","journal-title":"Sex Transm Dis"},{"key":"50779_CR32","doi-asserted-by":"publisher","first-page":"e0005434","DOI":"10.1371\/journal.pntd.0005434","volume":"11","author":"L Mikalov\u00e1","year":"2017","unstructured":"Mikalov\u00e1, L. et al. Human Treponema pallidum 11q\/j isolate belongs to subsp. endemicum but contains two loci with a sequence in TP0548 and TP0488 similar to subsp. pertenue and subsp. pallidum, respectively. PLoS Negl Trop Dis 11, e0005434 (2017).","journal-title":"PLoS Negl Trop Dis"},{"key":"50779_CR33","doi-asserted-by":"publisher","first-page":"1380","DOI":"10.1086\/656533","volume":"202","author":"C Marra","year":"2010","unstructured":"Marra, C. et al. Enhanced molecular typing of Treponema pallidum: geographical distribution of strain types and association with neurosyphilis. J Infect Dis 202, 1380\u20131388 (2010).","journal-title":"J Infect Dis"},{"key":"50779_CR34","doi-asserted-by":"publisher","first-page":"568","DOI":"10.1002\/ajpa.20560","volume":"132","author":"F Levrero","year":"2007","unstructured":"Levrero, F., Gatti, S., Gauthier-Hion, A. & Menard, N. Yaws disease in a wild gorilla population and its impact on the reproductive status of males. Am J Phys Anthropol 132, 568\u2013575 (2007).","journal-title":"Am J Phys Anthropol"},{"key":"50779_CR35","doi-asserted-by":"publisher","first-page":"1848","DOI":"10.1128\/IAI.01424-07","volume":"76","author":"M Brinkman","year":"2008","unstructured":"Brinkman, M. et al. A novel Treponema pallidum antigen, TP0136, is an outer membrane protein that binds human fibronectin. Infect Immun 76, 1848\u20131857 (2008).","journal-title":"Infect Immun"},{"key":"50779_CR36","first-page":"1","volume":"415","author":"JD Radolf","year":"2018","unstructured":"Radolf, J. D. & Kumar, S. The Treponema pallidum outer membrane. Curr Top Microbiol Immunol 415, 1\u201338 (2018).","journal-title":"Curr Top Microbiol Immunol"}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-019-50779-9.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-019-50779-9","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-019-50779-9.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,12,17]],"date-time":"2022-12-17T14:12:41Z","timestamp":1671286361000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-019-50779-9"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2019,10,2]]},"references-count":36,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2019,12]]}},"alternative-id":["50779"],"URL":"https:\/\/doi.org\/10.1038\/s41598-019-50779-9","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2019,10,2]]},"assertion":[{"value":"11 February 2019","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"19 September 2019","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"2 October 2019","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing Interests"}}],"article-number":"14243"}}