{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,5]],"date-time":"2026-08-05T09:33:49Z","timestamp":1785922429618,"version":"3.56.0"},"update-to":[{"DOI":"10.1371\/journal.ppat.1011572","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2023,9,1]],"date-time":"2023-09-01T00:00:00Z","timestamp":1693526400000}}],"reference-count":99,"publisher":"Public Library of Science (PLoS)","issue":"8","license":[{"start":{"date-parts":[[2023,8,22]],"date-time":"2023-08-22T00:00:00Z","timestamp":1692662400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100000038","name":"Natural Sciences and Engineering Research Council of Canada","doi-asserted-by":"publisher","award":["RGPIN-2019-04483"],"award-info":[{"award-number":["RGPIN-2019-04483"]}],"id":[{"id":"10.13039\/501100000038","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000106","name":"Saskatchewan Health Research Foundation","doi-asserted-by":"publisher","award":["4583"],"award-info":[{"award-number":["4583"]}],"id":[{"id":"10.13039\/501100000106","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100011094","name":"Public Health Agency of Canada","doi-asserted-by":"crossref","award":["4500428376"],"award-info":[{"award-number":["4500428376"]}],"id":[{"id":"10.13039\/100011094","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100023859","name":"Canadian Lyme Disease Foundation","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100023859","id-type":"DOI","asserted-by":"crossref"}]},{"name":"WCVM Wildlife Health Research Foundation"},{"DOI":"10.13039\/100008920","name":"University of Saskatchewan","doi-asserted-by":"crossref","award":["Dean's Fellowship"],"award-info":[{"award-number":["Dean's Fellowship"]}],"id":[{"id":"10.13039\/100008920","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["www.plospathogens.org"],"crossmark-restriction":false},"short-container-title":["PLoS Pathog"],"abstract":"<jats:p>Pathogen life history theory assumes a positive relationship between pathogen load in host tissues and pathogen transmission. Empirical evidence for this relationship is surprisingly rare due to the difficulty of measuring transmission for many pathogens. The comparative method, where a common host is experimentally infected with a set of pathogen strains, is a powerful approach for investigating the relationships between pathogen load and transmission. The validity of such experimental estimates of strain-specific transmission is greatly enhanced if they can predict the pathogen population strain structure in nature.<jats:italic>Borrelia burgdorferi<\/jats:italic>is a multi-strain, tick-borne spirochete that causes Lyme disease in North America. This study used 11 field-collected strains of<jats:italic>B<\/jats:italic>.<jats:italic>burgdorferi<\/jats:italic>, a rodent host (<jats:italic>Mus musculus<\/jats:italic>, C3H\/HeJ) and its tick vector (<jats:italic>Ixodes scapularis<\/jats:italic>) to determine the relationship between pathogen load in host tissues and lifetime host-to-tick transmission (HTT). Mice were experimentally infected via tick bite with 1 of 11 strains. Lifetime HTT was measured by infesting mice with<jats:italic>I<\/jats:italic>.<jats:italic>scapularis<\/jats:italic>larval ticks on 3 separate occasions. The prevalence and abundance of the strains in the mouse tissues and the ticks were determined by qPCR. We used published databases to obtain estimates of the frequencies of these strains in wild<jats:italic>I<\/jats:italic>.<jats:italic>scapularis<\/jats:italic>tick populations. Spirochete loads in ticks and lifetime HTT varied significantly among the 11 strains of<jats:italic>B<\/jats:italic>.<jats:italic>burgdorferi<\/jats:italic>. Strains with higher spirochete loads in the host tissues were more likely to infect feeding larval ticks, which molted into nymphal ticks that had a higher probability of<jats:italic>B<\/jats:italic>.<jats:italic>burgdorferi<\/jats:italic>infection (<jats:italic>i<\/jats:italic>.<jats:italic>e<\/jats:italic>., higher HTT). Our laboratory-based estimates of lifetime HTT were predictive of the frequencies of these strains in wild<jats:italic>I<\/jats:italic>.<jats:italic>scapularis<\/jats:italic>populations. For<jats:italic>B<\/jats:italic>.<jats:italic>burgdorferi<\/jats:italic>, the strains that establish high abundance in host tissues and that have high lifetime transmission are the strains that are most common in nature.<\/jats:p>","DOI":"10.1371\/journal.ppat.1011572","type":"journal-article","created":{"date-parts":[[2023,8,22]],"date-time":"2023-08-22T17:53:59Z","timestamp":1692726839000},"page":"e1011572","update-policy":"https:\/\/doi.org\/10.1371\/journal.ppat.corrections_policy","source":"Crossref","is-referenced-by-count":16,"title":["Variation among strains of Borrelia burgdorferi in host tissue abundance and lifetime transmission determine the population strain structure in nature"],"prefix":"10.1371","volume":"19","author":[{"given":"Christopher B.","family":"Zinck","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Prasobh","family":"Raveendram Thampy","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Eva-Maria E.","family":"Uhlemann","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hesham","family":"Adam","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jenny","family":"Wachter","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Danae","family":"Suchan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andrew D. S.","family":"Cameron","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ryan O. M.","family":"Rego","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dustin","family":"Brisson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Catherine","family":"Bouchard","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nicholas H.","family":"Ogden","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9384-0436","authenticated-orcid":true,"given":"Maarten J.","family":"Voordouw","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2023,8,22]]},"reference":[{"issue":"2","key":"ppat.1011572.ref001","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1111\/j.1420-9101.2008.01658.x","article-title":"Virulence evolution and the trade-off hypothesis: history, current state of affairs and the future","volume":"22","author":"S Alizon","year":"2009","journal-title":"J Evol Biol"},{"key":"ppat.1011572.ref002","doi-asserted-by":"crossref","first-page":"C42","DOI":"10.1016\/j.vaccine.2008.04.012","article-title":"Virulence evolution in response to vaccination: The case of malaria","volume":"26","author":"MJ Mackinnon","year":"2008","journal-title":"Vaccine"},{"issue":"1","key":"ppat.1011572.ref003","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1086\/419267","article-title":"Models of parasite virulence","volume":"71","author":"SA Frank","year":"1996","journal-title":"Q Rev Biol"},{"issue":"7","key":"ppat.1011572.ref004","doi-asserted-by":"crossref","first-page":"915","DOI":"10.1017\/S003118201500092X","article-title":"The adaptive evolution of virulence: a review of theoretical predictions and empirical tests","volume":"143","author":"CE Cressler","year":"2016","journal-title":"Parasitology"},{"issue":"OCT","key":"ppat.1011572.ref005","doi-asserted-by":"crossref","first-page":"411","DOI":"10.1017\/S0031182000055360","article-title":"Coevolution of hosts and parasites","volume":"85","author":"RM Anderson","year":"1982","journal-title":"Parasitology"},{"key":"ppat.1011572.ref006","first-page":"288","article-title":"The three Ts of virulence evolution during zoonotic emergence","author":"E Visher","year":"2021","journal-title":"P Roy Soc B-Biol Sci"},{"issue":"44","key":"ppat.1011572.ref007","doi-asserted-by":"crossref","first-page":"17441","DOI":"10.1073\/pnas.0708559104","article-title":"Variation in HIV-1 set-point viral load: Epidemiological analysis and an evolutionary hypothesis","volume":"104","author":"C Fraser","year":"2007","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"1446","key":"ppat.1011572.ref008","doi-asserted-by":"crossref","first-page":"965","DOI":"10.1098\/rstb.2003.1414","article-title":"Virulence in malaria: an evolutionary viewpoint","volume":"359","author":"MJ Mackinnon","year":"2004","journal-title":"Philos T Roy Soc B"},{"issue":"21","key":"ppat.1011572.ref009","doi-asserted-by":"crossref","first-page":"7489","DOI":"10.1073\/pnas.0710909105","article-title":"Virulence-transmission trade-offs and population divergence in virulence in a naturally occuring butterfly parasite","volume":"105","author":"JC de Roode","year":"2008","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"7","key":"ppat.1011572.ref010","doi-asserted-by":"crossref","first-page":"1265","DOI":"10.1371\/journal.pbio.0040197","article-title":"Empirical support for optimal virulence in a castrating parasite","volume":"4","author":"KH Jensen","year":"2006","journal-title":"PLOS Biol"},{"key":"ppat.1011572.ref011","article-title":"The evolution of life histories","author":"SC Stearns","year":"1992","journal-title":"Oxford: Oxford University Press"},{"issue":"3","key":"ppat.1011572.ref012","doi-asserted-by":"crossref","first-page":"689","DOI":"10.2307\/2640710","article-title":"Genetic relationships between parasite virulence and transmission in the rodent malaria Plasmodium chabaudi","volume":"53","author":"MJ Mackinnon","year":"1999","journal-title":"Evolution"},{"key":"ppat.1011572.ref013","doi-asserted-by":"crossref","first-page":"1448","DOI":"10.1111\/j.1420-9101.2012.02515.x","article-title":"Infection intensity and infectivity of the tick-borne pathogen Borrelia afzelii","volume":"25","author":"L. Raberg","year":"2012","journal-title":"J Evol Biol"},{"issue":"1747","key":"ppat.1011572.ref014","first-page":"4677","article-title":"Virulence, drug sensitivity and transmission success in the rodent malaria, Plasmodium chabaudi.","volume":"279","author":"P Schneider","year":"2012","journal-title":"P Roy Soc B-Biol Sci"},{"issue":"4","key":"ppat.1011572.ref015","doi-asserted-by":"crossref","first-page":"636","DOI":"10.1111\/evo.13692","article-title":"Virulence-driven trade-offs in disease transmission: A meta-analysis","volume":"73","author":"MA Acevedo","year":"2019","journal-title":"Evolution"},{"issue":"5","key":"ppat.1011572.ref016","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0065043","article-title":"An analysis of the sequence variability of meningococcal fHbp, NadA and NHBA over a 50-year period in the Netherlands","volume":"8","author":"S Bambini","year":"2013","journal-title":"PLOS ONE"},{"issue":"1688","key":"ppat.1011572.ref017","first-page":"1635","article-title":"Long-term evolution of antigen repertoires among carried meningococci","volume":"277","author":"CO Buckee","year":"2010","journal-title":"P Roy Soc B-Biol Sci"},{"issue":"1851","key":"ppat.1011572.ref018","article-title":"Fitness estimates from experimental infections predict the long-term strain structure of a vector-borne pathogen in the field","volume":"7","author":"J Durand","year":"2017","journal-title":"Scientific Reports"},{"issue":"5","key":"ppat.1011572.ref019","doi-asserted-by":"crossref","first-page":"1034","DOI":"10.1111\/jeb.13075","article-title":"Evolution of antigenic diversity in the tick-transmitted bacterium Borrelia afzelii: a role for host specialization","volume":"30","author":"L Raberg","year":"2017","journal-title":"J Evol Biol"},{"issue":"6","key":"ppat.1011572.ref020","doi-asserted-by":"crossref","DOI":"10.1371\/journal.ppat.1000476","article-title":"Pneumococcal capsular polysaccharide structure predicts serotype prevalence","volume":"5","author":"DM Weinberger","year":"2009","journal-title":"PLOS Pathog"},{"key":"ppat.1011572.ref021","first-page":"2","article-title":"Lyme borreliosis.","author":"AC Steere","year":"2016","journal-title":"Nature Reviews Disease Primers"},{"issue":"4","key":"ppat.1011572.ref022","doi-asserted-by":"crossref","first-page":"487","DOI":"10.1111\/j.1469-0691.2011.03492.x","article-title":"The expanding Lyme Borrelia complex-clinical significance of genomic species?","volume":"17","author":"G Stanek","year":"2011","journal-title":"Clin Microbiol Infec"},{"key":"ppat.1011572.ref023","doi-asserted-by":"crossref","first-page":"660","DOI":"10.1038\/nrmicro1475","article-title":"Fundamental processes in the evolutionary ecology of Lyme borreliosis","volume":"4","author":"K Kurtenbach","year":"2006","journal-title":"Nat Rev Microbiol"},{"key":"ppat.1011572.ref024","doi-asserted-by":"crossref","first-page":"S191","DOI":"10.1017\/S0031182003004694","article-title":"Lyme borreliosis in Europe and North America","volume":"129","author":"J Piesman","year":"2004","journal-title":"Parasitology"},{"issue":"6","key":"ppat.1011572.ref025","doi-asserted-by":"crossref","first-page":"e00434","DOI":"10.1128\/mBio.00434-12","article-title":"Experimental infections of the reservoir species Peromyscus leucopus with diverse strains of Borrelia burgdorferi, a Lyme disease agent.","volume":"3","author":"E Baum","year":"2012","journal-title":"mBio"},{"key":"ppat.1011572.ref026","article-title":"Competition between strains of Borrelia afzelii in the host tissues and consequences for transmission to ticks","author":"D Genn\u00e9","year":"2021","journal-title":"The ISME Journal"},{"key":"ppat.1011572.ref027","doi-asserted-by":"crossref","first-page":"131","DOI":"10.1016\/j.meegid.2015.09.012","article-title":"Cross-reactive acquired immunity influences transmission success of the Lyme disease pathogen, Borrelia afzelii","volume":"36","author":"M Jacquet","year":"2015","journal-title":"Infection Genetics and Evolution"},{"issue":"7","key":"ppat.1011572.ref028","doi-asserted-by":"crossref","first-page":"4303","DOI":"10.1128\/IAI.69.7.4303-4312.2001","article-title":"Impact of genotypic variation of Borrelia burgdorferi sensu stricto on kinetics of dissemination and severity of disease in C3H\/HeJ mice","volume":"69","author":"G Wang","year":"2001","journal-title":"Infect Immun"},{"issue":"6","key":"ppat.1011572.ref029","doi-asserted-by":"crossref","first-page":"782","DOI":"10.1086\/343043","article-title":"Disease severity in a murine model of Lyme borreliosis is associated with the genotype of the infecting Borrelia burgdorferi sensu stricto strain","volume":"186","author":"GQ Wang","year":"2002","journal-title":"J Infect Dis"},{"key":"ppat.1011572.ref030","doi-asserted-by":"crossref","first-page":"713","DOI":"10.1534\/genetics.104.028738","article-title":"ospC diversity in Borrelia burgdorferi: different hosts are different niches","volume":"168","author":"D Brisson","year":"2004","journal-title":"Genetics"},{"issue":"11","key":"ppat.1011572.ref031","doi-asserted-by":"crossref","first-page":"6783","DOI":"10.1128\/AEM.70.11.6783-6788.2004","article-title":"Interaction and transmission of two Borrelia burgdorferi sensu stricto strains in a tick-rodent maintenance system","volume":"70","author":"M Derdakova","year":"2004","journal-title":"Appl Environ Microbiol"},{"issue":"1890","key":"ppat.1011572.ref032","article-title":"Competition between strains of Borrelia afzelii inside the rodent host and the tick vector.","volume":"285","author":"D Genn\u00e9","year":"2018","journal-title":"P Roy Soc B-Biol Sci"},{"issue":"431","key":"ppat.1011572.ref033","first-page":"1","article-title":"Competition between strains of Borrelia afzelii in immature Ixodes ricinus ticks is not affected by season","volume":"9","author":"D Genn\u00e9","year":"2019","journal-title":"Front Cell Infect Microbiol"},{"issue":"1","key":"ppat.1011572.ref034","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1128\/AEM.01567-07","article-title":"Fitness variation of Borrelia burgdorferi sensu stricto strains in mice","volume":"74","author":"K Hanincova","year":"2008","journal-title":"Appl Environ Microbiol"},{"key":"ppat.1011572.ref035","first-page":"9","article-title":"Comparison of the lifetime host-to-tick transmission between two strains of the Lyme disease pathogen Borrelia afzelii.","author":"M Jacquet","year":"2016","journal-title":"Parasit Vectors"},{"issue":"64","key":"ppat.1011572.ref036","first-page":"1","article-title":"Closely-related Borrelia burgdorferi (sensu stricto) strains exhibit similar fitness in single infections and asymmetric competition in multiple infections","volume":"10","author":"EC Rynkiewicz","year":"2017"},{"issue":"3","key":"ppat.1011572.ref037","doi-asserted-by":"crossref","first-page":"334","DOI":"10.1016\/j.ttbdis.2015.02.007","article-title":"Genetic variation in transmission success of the Lyme borreliosis pathogen Borrelia afzelii.","volume":"6","author":"N Tonetti","year":"2015","journal-title":"Ticks Tick Borne Dis"},{"issue":"2","key":"ppat.1011572.ref038","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0118265","article-title":"The Lyme disease pathogen has no effect on the survival of its rodent reservoir host.","volume":"10","author":"MJ Voordouw","year":"2015","journal-title":"PLOS ONE"},{"issue":"2","key":"ppat.1011572.ref039","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1089\/vbz.2009.0215","article-title":"Borrelia burgdorferi has minimal impact on the Lyme disease reservoir host Peromyscus leucopus","volume":"11","author":"LE Schwanz","year":"2011","journal-title":"Vector-Borne Zoonot"},{"key":"ppat.1011572.ref040","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1016\/j.semcdb.2016.07.002","article-title":"Infection resistance and tolerance in Peromyscus spp., natural reservoirs of microbes that are virulent for humans","volume":"61","author":"AG Barbour","year":"2017","journal-title":"Semin Cell Dev Biol"},{"issue":"2","key":"ppat.1011572.ref041","doi-asserted-by":"crossref","first-page":"155","DOI":"10.7589\/0090-3558-30.2.155","article-title":"Experimental Borrelia burgdorferi infection in Peromyscus leucopus","volume":"30","author":"KD Moody","year":"1994","journal-title":"J Wildl Dis"},{"key":"ppat.1011572.ref042","article-title":"Borrelia burgdorferi strain and host sex influence pathogen prevalence and abundance in the tissues of a laboratory rodent host","author":"CB Zinck","year":"2022","journal-title":"Mol Ecol"},{"key":"ppat.1011572.ref043","first-page":"8","article-title":"Whole genome sequencing and phylogenetic analysis of strains of the agent of Lyme disease Borrelia burgdorferi from Canadian emergence zones","author":"S Tyler","year":"2018","journal-title":"Scientific Reports"},{"issue":"17","key":"ppat.1011572.ref044","doi-asserted-by":"crossref","first-page":"6059","DOI":"10.1128\/AEM.00231-12","article-title":"Two boundaries separate Borrelia burgdorferi populations in North America","volume":"78","author":"G Margos","year":"2012","journal-title":"Appl Environ Microbiol"},{"issue":"4","key":"ppat.1011572.ref045","doi-asserted-by":"crossref","first-page":"1309","DOI":"10.1128\/AEM.03730-14","article-title":"Complex population structure of Borrelia burgdorferi in southeastern and south central Canada as revealed by phylogeographic analysis","volume":"81","author":"S Mechai","year":"2015","journal-title":"Appl Environ Microbiol"},{"issue":"10","key":"ppat.1011572.ref046","doi-asserted-by":"crossref","first-page":"3244","DOI":"10.1128\/AEM.02636-10","article-title":"Investigation of genotypes of Borrelia burgdorferi in Ixodes scapularis ticks collected during surveillance in Canada","volume":"77","author":"NH Ogden","year":"2011","journal-title":"Appl Environ Microbiol"},{"issue":"9","key":"ppat.1011572.ref047","doi-asserted-by":"crossref","first-page":"5262","DOI":"10.1128\/IAI.72.9.5262-5266.2004","article-title":"Comparison of disseminated and nondisseminated strains of Borrelia burgdorferi sensu stricto in mice naturally infected by tick bite","volume":"72","author":"MC Dolan","year":"2004","journal-title":"Infect Immun"},{"issue":"1","key":"ppat.1011572.ref048","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1128\/IAI.66.1.161-168.1998","article-title":"Distinct characteristics of resistance to Borrelia burgdorferi induced arthritis in C57BL\/6N mice","volume":"66","author":"Y Ma","year":"1998","journal-title":"Infect Immun"},{"issue":"2","key":"ppat.1011572.ref049","first-page":"263","article-title":"Kinetics of Borrelia burgdorferi dissemination and evolution of disease after intradermal inoculation of mice","volume":"139","author":"SW Barthold","year":"1991","journal-title":"Am J Pathol"},{"issue":"7","key":"ppat.1011572.ref050","doi-asserted-by":"crossref","first-page":"3164","DOI":"10.1128\/JCM.42.7.3164-3168.2004","article-title":"Multiplex real-time PCR for detection of Anaplasma phagocytophilum and Borrelia burgdorferi","volume":"42","author":"JW Courtney","year":"2004","journal-title":"J Clin Microbiol"},{"issue":"5","key":"ppat.1011572.ref051","doi-asserted-by":"crossref","first-page":"1014","DOI":"10.1111\/j.1365-2672.2012.05384.x","article-title":"PCR inhibitors\u2014occurrence, properties and removal","volume":"113","author":"C Schrader","year":"2012","journal-title":"J Appl Microbiol"},{"issue":"10","key":"ppat.1011572.ref052","doi-asserted-by":"crossref","first-page":"2569","DOI":"10.1007\/s00216-018-0931-z","article-title":"Inhibition mechanisms of hemoglobin, immunoglobulin G, and whole blood in digital and real-time PCR","volume":"410","author":"M Sidstedt","year":"2018","journal-title":"Anal Bioanal Chem"},{"issue":"7","key":"ppat.1011572.ref053","doi-asserted-by":"crossref","first-page":"1147","DOI":"10.3201\/eid1607.091452","article-title":"Geographic differences in genetic locus linkages for Borrelia burgdorferi","volume":"16","author":"B Travinsky","year":"2010","journal-title":"Emerg Infect Dis"},{"key":"ppat.1011572.ref054","article-title":"Variation among strains of Borrelia burgdorferi in host tissue abundance and lifetime transmission determine the population strain structure in nature","author":"CB Zinck","year":"2023","journal-title":"In: Dryad"},{"key":"ppat.1011572.ref055","author":"R Development Core Team","year":"2021"},{"issue":"2","key":"ppat.1011572.ref056","doi-asserted-by":"crossref","first-page":"567","DOI":"10.1073\/pnas.0233733100","article-title":"The ecology of infectious disease: Effects of host diversity and community composition on Lyme disease risk","volume":"100","author":"K LoGiudice","year":"2003","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"1884","key":"ppat.1011572.ref057","article-title":"Borrelia afzelii alters reproductive success in a rodent host.","volume":"285","author":"C Cayol","year":"2018","journal-title":"P Roy Soc B-Biol Sci"},{"key":"ppat.1011572.ref058","doi-asserted-by":"crossref","first-page":"1515","DOI":"10.1086\/382594","article-title":"Borrelia burgdorferi infection in a natural population of Peromyscus leucopus mice: a longitudinal study in an area where Lyme borreliosis is highly endemic","volume":"189","author":"J Bunikis","year":"2004","journal-title":"J Infect Dis"},{"issue":"12","key":"ppat.1011572.ref059","doi-asserted-by":"crossref","first-page":"1980","DOI":"10.2460\/ajvr.1990.51.12.1980","article-title":"Experimental infection of the white-footed mouse with Borrelia burgdorferi","volume":"51","author":"SD Wright","year":"1990","journal-title":"Am J Vet Res"},{"issue":"5","key":"ppat.1011572.ref060","doi-asserted-by":"crossref","first-page":"186","DOI":"10.1016\/S0169-4758(98)01224-1","article-title":"Ticks are not insects: consequences of contrasting vector biology for transmission potential","volume":"14","author":"SE Randolph","year":"1998","journal-title":"Parasitol Today"},{"issue":"21","key":"ppat.1011572.ref061","doi-asserted-by":"crossref","first-page":"7624","DOI":"10.1073\/pnas.0500078102","article-title":"Virulence and competitive ability in genetically diverse malaria infections","volume":"102","author":"JC de Roode","year":"2005","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"3","key":"ppat.1011572.ref062","doi-asserted-by":"crossref","first-page":"311","DOI":"10.3201\/eid0903.020628","article-title":"Experimental infection of north American birds with the New York 1999 strain of West Nile virus","volume":"9","author":"N Komar","year":"2003","journal-title":"Emerg Infect Dis"},{"issue":"4","key":"ppat.1011572.ref063","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1038\/ni.1855","article-title":"An intravascular immune response to Borrelia burgdorferi involves Kupffer cells and iNKT cells","volume":"11","author":"WY Lee","year":"2010","journal-title":"Nat Immunol"},{"issue":"2","key":"ppat.1011572.ref064","article-title":"Reviewing molecular adaptations of Lyme borreliosis spirochetes in the context of reproductive fitness in natural transmission cycles","volume":"40","author":"J. Tsao","year":"2009","journal-title":"Vet Res (Paris)."},{"issue":"5","key":"ppat.1011572.ref065","doi-asserted-by":"crossref","first-page":"616","DOI":"10.4269\/ajtmh.2002.66.616","article-title":"Chemotactic migration of the Lyme disease spirochete (Borrelia burgdorferi) to salivary gland extracts of vector ticks","volume":"66","author":"CM Shih","year":"2002","journal-title":"Am J Trop Med Hyg"},{"issue":"5","key":"ppat.1011572.ref066","doi-asserted-by":"crossref","DOI":"10.1016\/j.ttbdis.2021.101782","article-title":"A transwell assay method to evaluate Borrelia burgdorferi sensu stricto migratory chemoattraction toward tick saliva proteins","volume":"12","author":"TJ Van Gundy","year":"2021","journal-title":"Ticks Tick Borne Dis"},{"issue":"3","key":"ppat.1011572.ref067","doi-asserted-by":"crossref","first-page":"600","DOI":"10.1093\/jmedent\/43.3.600","article-title":"Ixodes scapularis ticks collected by passive surveillance in Canada: Analysis of geographic distribution and infection with Lyme Borreliosis agent Borrelia burgdorferi","volume":"43","author":"NH Ogden","year":"2006","journal-title":"J Med Entomol"},{"issue":"6","key":"ppat.1011572.ref068","doi-asserted-by":"crossref","first-page":"1120","DOI":"10.4269\/ajtmh.2009.09-0208","article-title":"Niche partitioning of Borrelia burgdorferi and Borrelia miyamotoi in the same tick vector and mammalian reservoir species","volume":"81","author":"AG Barbour","year":"2009","journal-title":"Am J Trop Med Hyg"},{"issue":"2","key":"ppat.1011572.ref069","doi-asserted-by":"crossref","first-page":"166","DOI":"10.1093\/jmedent\/43.2.166","article-title":"Spatiotemporal patterns of host-seeking Ixodes scapularis nymphs (Acari: Iodidae) in the United States","volume":"43","author":"MA Diuk-Wasser","year":"2006","journal-title":"J Med Entomol"},{"issue":"4","key":"ppat.1011572.ref070","doi-asserted-by":"crossref","first-page":"604","DOI":"10.3201\/eid1204.051016","article-title":"Epidemic spread of Lyme borreliosis, Northeastern United States","volume":"12","author":"K Hanincova","year":"2006","journal-title":"Emerg Infect Dis"},{"issue":"2","key":"ppat.1011572.ref071","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0149345","article-title":"Evidence for host-genotype associations of Borrelia burgdorferi sensu stricto.","volume":"11","author":"S Mechai","year":"2016","journal-title":"PLOS ONE"},{"key":"ppat.1011572.ref072","doi-asserted-by":"crossref","first-page":"594","DOI":"10.1016\/j.meegid.2013.12.011","article-title":"Occurrence and transmission efficiencies of Borrelia burgdorferi ospC types in avian and mammalian wildlife","volume":"27","author":"HB Vuong","year":"2014","journal-title":"Infect Genet Evol"},{"issue":"1","key":"ppat.1011572.ref073","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1093\/jmedent\/45.1.139","article-title":"Estimating reservoir competence of Borrelia burgdorferi hosts: prevalence and infectivity, sensitivity, and specificity","volume":"45","author":"J Brunner","year":"2008","journal-title":"J Med Entomol"},{"issue":"1","key":"ppat.1011572.ref074","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1093\/oxfordjournals.aje.a115306","article-title":"Comparing the relative potential of rodents as reservoirs of the Lyme disease spirochete (Borrelia burgdorferi).","volume":"130","author":"TN Mather","year":"1989","journal-title":"Am J Epidemiol"},{"issue":"6","key":"ppat.1011572.ref075","doi-asserted-by":"crossref","first-page":"1058","DOI":"10.1371\/journal.pbio.0040145","article-title":"Climate, deer, rodents, and acorns as determinants of variation in Lyme-disease risk","volume":"4","author":"RS Ostfeld","year":"2006","journal-title":"PLOS Biol"},{"issue":"1","key":"ppat.1011572.ref076","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1089\/153036601750137688","article-title":"Effects of acorn production and mouse abundance on abundance and Borrelia burgdorferi infection prevalence of nymphal Ixodes scapularis ticks.","volume":"1","author":"RS Ostfeld","year":"2001","journal-title":"Vector-Borne Zoonot."},{"issue":"5353","key":"ppat.1011572.ref077","doi-asserted-by":"crossref","first-page":"1023","DOI":"10.1126\/science.279.5353.1023","article-title":"Chain reactions linking acorns to gypsy moth outbreaks and Lyme disease risk","volume":"279","author":"CG Jones","year":"1998","journal-title":"Science"},{"issue":"6","key":"ppat.1011572.ref078","doi-asserted-by":"crossref","DOI":"10.1007\/s11538-021-00896-2","article-title":"Host Specialisation, Immune Cross-Reaction and the Composition of Communities of Co-circulating Borrelia Strains","volume":"83","author":"B Adams","year":"2021","journal-title":"Bull Math Biol"},{"issue":"3","key":"ppat.1011572.ref079","doi-asserted-by":"crossref","first-page":"1056","DOI":"10.1073\/pnas.0334222100","article-title":"Placental mammal diversification and the Cretaceous-Tertiary boundary","volume":"100","author":"MS Springer","year":"2003","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"3","key":"ppat.1011572.ref080","first-page":"397","article-title":"Presence of multiple variants of Borrelia burgdorferi in the natural reservoir Peromyscus leucopus throughout a transmission season.","volume":"8","author":"KI Swanson","year":"2008"},{"issue":"4","key":"ppat.1011572.ref081","doi-asserted-by":"crossref","first-page":"545","DOI":"10.1086\/669905","article-title":"Multiple-strain infections of Borrelia afzelii: a role for within-host interactions in the maintenance of antigenic diversity","volume":"181","author":"M Andersson","year":"2013","journal-title":"Am Nat"},{"issue":"1675","key":"ppat.1011572.ref082","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1098\/rstb.2014.0293","article-title":"Within-host competition between Borrelia afzelii ospC strains in wild hosts as revealed by massively parallel amplicon sequencing","volume":"370","author":"M Strandh","year":"2015","journal-title":"Philos T Roy Soc B"},{"issue":"3","key":"ppat.1011572.ref083","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1016\/j.ttbdis.2011.06.003","article-title":"Borrelia afzelii ospC genotype diversity in Ixodes ricinus questing ticks and ticks from rodents in two Lyme borreliosis endemic areas: Contribution of co-feeding ticks","volume":"2","author":"D P\u00e9rez","year":"2011","journal-title":"Ticks Tick Borne Dis"},{"issue":"6","key":"ppat.1011572.ref084","doi-asserted-by":"crossref","first-page":"102058","DOI":"10.1016\/j.ttbdis.2022.102058","article-title":"Clobetasol increases the abundance of Borrelia burgdorferi in the skin 70 times more in male mice compared to female mice","volume":"13","author":"B Bourgeois","year":"2022","journal-title":"Ticks Tick Borne Dis"},{"issue":"10","key":"ppat.1011572.ref085","doi-asserted-by":"crossref","first-page":"626","DOI":"10.1038\/nri.2016.90","article-title":"Sex differences in immune responses","volume":"16","author":"SL Klein","year":"2016","journal-title":"Nat Rev Immunol"},{"issue":"2","key":"ppat.1011572.ref086","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1016\/j.cellimm.2015.02.004","article-title":"Suppressive effects of androgens on the immune system","volume":"294","author":"A Trigunaite","year":"2015","journal-title":"Cell Immunol"},{"issue":"2","key":"ppat.1011572.ref087","doi-asserted-by":"crossref","DOI":"10.1371\/journal.ppat.1005374","article-title":"SeXX Matters in Infectious Disease Pathogenesis.","volume":"12","author":"LG vom Steeg","year":"2016","journal-title":"PLOS Pathog"},{"issue":"1759","key":"ppat.1011572.ref088","first-page":"20130364","article-title":"Polymorphisms at the innate immune receptor TLR2 are associated with Borrelia infection in a wild rodent population.","volume":"280","author":"B Tschirren","year":"2013","journal-title":"P Roy Soc B-Biol Sci"},{"issue":"1","key":"ppat.1011572.ref089","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0226798","article-title":"Optimization of tissue sampling for Borrelia burgdorferi in white-footed mice (Peromyscus leucopus).","volume":"15","author":"SG Zawada","year":"2020","journal-title":"PLOS ONE."},{"issue":"2","key":"ppat.1011572.ref090","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1016\/j.ttbdis.2011.11.004","article-title":"Synchronous phenology of juvenile Ixodes scapularis, vertebrate host relationships, and associated patterns of Borrelia burgdorferi ribotypes in the midwestern United States.","volume":"3","author":"SA Hamer","year":"2012","journal-title":"Ticks Tick Borne Dis"},{"key":"ppat.1011572.ref091","doi-asserted-by":"crossref","first-page":"915","DOI":"10.1017\/S003118201200008X","article-title":"The effect of spatial heterogeneity on the aggregation of ticks on white-footed mice","volume":"139","author":"G Devevey","year":"2012","journal-title":"Parasitology"},{"issue":"8","key":"ppat.1011572.ref092","doi-asserted-by":"crossref","first-page":"4074","DOI":"10.1002\/ece3.3961","article-title":"Effects of a zoonotic pathogen, Borrelia burgdorferi, on the behavior of a key reservoir host.","volume":"8","author":"RS Ostfeld","year":"2018","journal-title":"Ecol Evol"},{"key":"ppat.1011572.ref093","doi-asserted-by":"crossref","first-page":"909","DOI":"10.1016\/S0020-7519(03)00128-0","article-title":"Empirical evidence for key hosts in persistence of a tick-borne disease","volume":"33","author":"SE Perkins","year":"2003","journal-title":"Int J Parasitol"},{"issue":"4","key":"ppat.1011572.ref094","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1016\/j.ttbdis.2011.03.005","article-title":"Associations between Ixodes scapularis ticks and small mammal hosts in a newly endemic zone in southeastern Canada: Implications for Borrelia burgdorferi transmission.","volume":"2","author":"C Bouchard","year":"2011","journal-title":"Ticks Tick Borne Dis"},{"issue":"4","key":"ppat.1011572.ref095","doi-asserted-by":"crossref","first-page":"589","DOI":"10.4269\/ajtmh.1999.60.598","article-title":"Longitudinal study of infection with Borrelia burgdorferi in a population of Peromyscus leucopus at a Lyme disease-enzootic site in Maryland","volume":"60","author":"EK Hofmeister","year":"1999","journal-title":"Am J Trop Med Hyg"},{"issue":"11","key":"ppat.1011572.ref096","doi-asserted-by":"crossref","first-page":"e1003766","DOI":"10.1371\/journal.ppat.1003766","article-title":"Natural selection promotes antigenic evolvability","volume":"9","author":"C Graves","year":"2013","journal-title":"PLOS Pathog"},{"key":"ppat.1011572.ref097","doi-asserted-by":"crossref","first-page":"92","DOI":"10.4269\/ajtmh.1987.36.92","article-title":"Reservoir competence of white-footed mice for Lyme disease spirochetes","volume":"36","author":"JG Donahue","year":"1987","journal-title":"Am J Trop Med Hyg"},{"issue":"4","key":"ppat.1011572.ref098","doi-asserted-by":"crossref","first-page":"2442","DOI":"10.1128\/IAI.72.4.2442-2444.2004","article-title":"Adaptation of diverse Lyme disease spirochetes in a natural rodent reservoir host","volume":"72","author":"D Richter","year":"2004","journal-title":"Infect Immun"},{"issue":"8","key":"ppat.1011572.ref099","doi-asserted-by":"crossref","first-page":"2476","DOI":"10.1128\/AEM.02633-08","article-title":"Climate and tick seasonality are predictors of Borrelia burgdorferi genotype distribution","volume":"75","author":"AG Gatewood","year":"2009","journal-title":"Appl Environ Microbiol"}],"updated-by":[{"DOI":"10.1371\/journal.ppat.1011572","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2023,9,1]],"date-time":"2023-09-01T00:00:00Z","timestamp":1693526400000}}],"container-title":["PLOS Pathogens"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.ppat.1011572","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,10,26]],"date-time":"2024-10-26T15:53:01Z","timestamp":1729957981000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.ppat.1011572"}},"subtitle":[],"editor":[{"given":"D. Scott","family":"Samuels","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2023,8,22]]},"references-count":99,"journal-issue":{"issue":"8","published-online":{"date-parts":[[2023,8,22]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.ppat.1011572","relation":{"new_version":[{"id-type":"doi","id":"10.1371\/journal.ppat.1011572","asserted-by":"object"}]},"ISSN":["1553-7374"],"issn-type":[{"value":"1553-7374","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,8,22]]}}}