{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,17]],"date-time":"2026-09-17T18:13:48Z","timestamp":1789668828239,"version":"4.0.1"},"reference-count":37,"publisher":"The Royal Society","issue":"10","license":[{"start":{"date-parts":[[2021,10,1]],"date-time":"2021-10-01T00:00:00Z","timestamp":1633046400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/royalsociety.org\/-\/media\/journals\/author\/Licence-to-Publish-20062019-final.pdf"},{"start":{"date-parts":[[2021,10,1]],"date-time":"2021-10-01T00:00:00Z","timestamp":1633046400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/royalsociety.org\/journals\/ethics-policies\/data-sharing-mining\/"},{"start":{"date-parts":[[2021,10,1]],"date-time":"2021-10-01T00:00:00Z","timestamp":1633046400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/royalsociety.org\/journals\/ethics-policies\/data-sharing-mining\/"}],"funder":[{"name":"National Science Foundation","award":["IOS-1950307"],"award-info":[{"award-number":["IOS-1950307"]}]},{"name":"National Science Foundation","award":["IOS-1950307"],"award-info":[{"award-number":["IOS-1950307"]}]}],"content-domain":{"domain":["royalsocietypublishing.org"],"crossmark-restriction":true},"short-container-title":[],"published-print":{"date-parts":[[2021,10,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Individuals can express a range of disease phenotypes during infection, with important implications for epidemics. Tolerance, in particular, is a host response that minimizes the per-pathogen fitness costs of infection. Because tolerant hosts show milder clinical signs and higher survival, despite similar pathogen burdens, their potential for prolonged pathogen shedding may facilitate the spread of pathogens. To test this, we simulated outbreaks of mycoplasmal conjunctivitis in house finches, asking how the speed of transmission varied with tissue-specific and behavioural components of tolerance, milder conjunctivitis and anorexia for a given pathogen load, respectively. Because tissue-specific tolerance hinders pathogen deposition onto bird feeders, important transmission hubs, we predicted it would slow transmission. Because behavioural tolerance should increase interactions with bird feeders, we predicted it would speed transmission. Our findings supported these predictions, suggesting that variation in tolerance could help identify individuals most likely to transmit pathogens.<\/jats:p>","DOI":"10.1098\/rsbl.2021.0362","type":"journal-article","created":{"date-parts":[[2021,10,26]],"date-time":"2021-10-26T19:05:49Z","timestamp":1635275149000},"update-policy":"https:\/\/doi.org\/10.1098\/crossmark-policy","source":"Crossref","is-referenced-by-count":31,"title":["Disease tolerance alters host competence in a wild songbird"],"prefix":"10.1098","volume":"17","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-3764-9447","authenticated-orcid":true,"given":"Rachel M.","family":"Ruden","sequence":"first","affiliation":[{"name":"Ames, IA, USA 1","department":["Department of Natural Resource Ecology and Management, Iowa State University,"]},{"name":"Des Moines, IA, USA 2","department":["Iowa Department of Natural Resources,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5577-8798","authenticated-orcid":true,"given":"James S.","family":"Adelman","sequence":"additional","affiliation":[{"name":"Ames, IA, USA 1","department":["Department of Natural Resource Ecology and Management, Iowa State University,"]},{"name":"Memphis, TN, USA 3","department":["Department of Biological Sciences, University of Memphis,"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"175","published-online":{"date-parts":[[2021,10,27]]},"reference":[{"key":"2026010519421838300_RSBL20210362C1","doi-asserted-by":"crossref","first-page":"419","DOI":"10.1016\/j.pt.2015.05.002","article-title":"The context of host competence: a role for plasticity in host\u2013parasite dynamics","volume":"31","author":"Gervasi","year":"2015","journal-title":"Trends Parasitol."},{"key":"2026010519421838300_RSBL20210362C2","doi-asserted-by":"crossref","first-page":"1225","DOI":"10.1093\/icb\/icw064","article-title":"Host competence: an organismal trait to integrate immunology and epidemiology","volume":"56","author":"Martin","year":"2016","journal-title":"Integr. Comp. Biol."},{"key":"2026010519421838300_RSBL20210362C3","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1086\/701169","article-title":"How individual variation in host tolerance affects competence to transmit parasites","volume":"92","author":"Burgan","year":"2019","journal-title":"Physiol. Biochem. Zool."},{"key":"2026010519421838300_RSBL20210362C4","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1016\/j.tree.2018.12.009","article-title":"Extreme competence: keystone hosts of infections","volume":"34","author":"Martin","year":"2019","journal-title":"Trends Ecol. Evol."},{"key":"2026010519421838300_RSBL20210362C5","doi-asserted-by":"crossref","first-page":"355","DOI":"10.1038\/nature04153","article-title":"Superspreading and the effect of individual variation on disease emergence","volume":"438","author":"Lloyd-Smith","year":"2005","journal-title":"Nature"},{"key":"2026010519421838300_RSBL20210362C6","doi-asserted-by":"crossref","first-page":"e1001769","DOI":"10.1371\/journal.pbio.1001769","article-title":"Limiting damage during infection: lessons from infection tolerance for novel therapeutics","volume":"12","author":"Vale","year":"2014","journal-title":"PLoS Biol."},{"key":"2026010519421838300_RSBL20210362C7","doi-asserted-by":"crossref","first-page":"1253","DOI":"10.1093\/icb\/icz074","article-title":"Modeling the effects of resource-driven immune defense on parasite transmission in heterogeneous host populations","volume":"59","author":"Hall","year":"2019","journal-title":"Integr. Comp. Biol."},{"key":"2026010519421838300_RSBL20210362C8","doi-asserted-by":"crossref","first-page":"1271","DOI":"10.1111\/jeb.12379","article-title":"Evidence of trade-offs shaping virulence evolution in an emerging wildlife pathogen","volume":"27","author":"Williams","year":"2014","journal-title":"J. Evol. Biol."},{"key":"2026010519421838300_RSBL20210362C9","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1002\/evl3.203","article-title":"Experimental evidence for stabilizing selection on virulence in a bacterial pathogen","volume":"4","author":"Bonneaud","year":"2020","journal-title":"Evol. Lett."},{"key":"2026010519421838300_RSBL20210362C10","doi-asserted-by":"crossref","first-page":"e1001989","DOI":"10.1371\/journal.pbio.1001989","article-title":"How to live with the enemy: understanding tolerance to parasites","volume":"12","author":"R\u00e5berg","year":"2014","journal-title":"PLoS Biol."},{"key":"2026010519421838300_RSBL20210362C11","doi-asserted-by":"crossref","first-page":"812","DOI":"10.1126\/science.1148526","article-title":"Disentangling genetic variation for resistance and tolerance to infectious diseases in animals","volume":"318","author":"R\u00e5berg","year":"2007","journal-title":"Science"},{"key":"2026010519421838300_RSBL20210362C12","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1016\/j.yhbeh.2016.10.013","article-title":"Tolerance of infection: a role for animal behavior, potential immune mechanisms, and consequences for parasite transmission","volume":"88","author":"Adelman","year":"2017","journal-title":"Horm. Behav."},{"key":"2026010519421838300_RSBL20210362C13","doi-asserted-by":"crossref","first-page":"1220","DOI":"10.1093\/icb\/icz065","article-title":"What does tolerance mean for animal disease dynamics when pathology enhances transmission?","volume":"59","author":"Henschen","year":"2019","journal-title":"Integr. Comp. Biol."},{"key":"2026010519421838300_RSBL20210362C14","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1098\/rstb.2008.0184","article-title":"Decomposing health: tolerance and resistance to parasites in animals","volume":"364","author":"R\u00e5berg","year":"2009","journal-title":"Phil. Trans. R. Soc. B"},{"key":"2026010519421838300_RSBL20210362C15","doi-asserted-by":"crossref","first-page":"443","DOI":"10.1016\/S0165-2427(02)00069-7","article-title":"The concept of sickness behavior: a brief chronological account of four key discoveries","volume":"87","author":"Johnson","year":"2002","journal-title":"Vet. Immunol. Immunopathol."},{"key":"2026010519421838300_RSBL20210362C16","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1016\/S0149-7634(88)80004-6","article-title":"Biological basis of the behavior of sick animals","volume":"12","author":"Hart","year":"1988","journal-title":"Neurosci. Biobehav. Rev."},{"key":"2026010519421838300_RSBL20210362C17","doi-asserted-by":"crossref","first-page":"936","DOI":"10.1126\/science.1214935","article-title":"Disease tolerance as a defense strategy","volume":"335","author":"Medzhitov","year":"2012","journal-title":"Science"},{"key":"2026010519421838300_RSBL20210362C18","doi-asserted-by":"crossref","first-page":"480","DOI":"10.2307\/1592250","article-title":"Mycoplasma gallisepticum isolated from house finches (Carpodacus mexicanus) with conjunctivitis","volume":"40","author":"Ley","year":"1996","journal-title":"Avian Dis."},{"key":"2026010519421838300_RSBL20210362C19","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1080\/03079450701286277","article-title":"Experimental evidence for transmission of Mycoplasma gallisepticum in house finches by fomites","volume":"36","author":"Dhondt","year":"2007","journal-title":"Avian Pathol."},{"key":"2026010519421838300_RSBL20210362C20","doi-asserted-by":"crossref","first-page":"20151429","DOI":"10.1098\/rspb.2015.1429","article-title":"Feeder use predicts both acquisition and transmission of a contagious pathogen in a North American songbird","volume":"282","author":"Adelman","year":"2015","journal-title":"Proc. R. Soc. B"},{"key":"2026010519421838300_RSBL20210362C21","doi-asserted-by":"crossref","first-page":"7866","DOI":"10.1073\/pnas.1018580108","article-title":"Rapid evolution of disease resistance is accompanied by functional changes in gene expression in a wild bird","volume":"108","author":"Bonneaud","year":"2011","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2026010519421838300_RSBL20210362C22","doi-asserted-by":"crossref","first-page":"544","DOI":"10.1002\/evl3.133","article-title":"Evolution of both host resistance and tolerance to an emerging bacterial pathogen","volume":"3","author":"Bonneaud","year":"2019","journal-title":"Evol. Lett."},{"key":"2026010519421838300_RSBL20210362C23","doi-asserted-by":"crossref","first-page":"674","DOI":"10.1086\/670024","article-title":"House finch populations differ in early inflammatory signaling and pathogen tolerance at the peak of Mycoplasma gallisepticum infection","volume":"181","author":"Adelman","year":"2013","journal-title":"Am. Nat."},{"key":"2026010519421838300_RSBL20210362C24","doi-asserted-by":"crossref","first-page":"20130594","DOI":"10.1098\/rsbl.2013.0594","article-title":"Deposition of pathogenic Mycoplasma gallisepticum onto bird feeders: host pathology is more important than temperature-driven increases in food intake","volume":"9","author":"Adelman","year":"2013","journal-title":"Biol. Lett."},{"key":"2026010519421838300_RSBL20210362C25","doi-asserted-by":"crossref","first-page":"793","DOI":"10.1007\/s10393-017-1261-x","article-title":"Host responses to pathogen priming in a natural songbird host","volume":"14","author":"Leon","year":"2017","journal-title":"EcoHealth"},{"key":"2026010519421838300_RSBL20210362C26","volume-title":"Identification guide to north American birds, part 1: Columbidae to Ploceidae","author":"Pyle","year":"1997"},{"key":"2026010519421838300_RSBL20210362C27","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1637\/7403-062805R.1","article-title":"Characterization of experimental Mycoplasma gallisepticum infection in captive house finch flocks","volume":"50","author":"Sydenstricker","year":"2006","journal-title":"Avian Dis."},{"key":"2026010519421838300_RSBL20210362C28","doi-asserted-by":"crossref","first-page":"385","DOI":"10.1080\/03079450802216629","article-title":"Detection and quantification of Mycoplasma gallisepticum genome load in conjunctival samples of experimentally infected house finches (Carpodacus mexicanus) using real-time polymerase chain reaction","volume":"37","author":"Grodio","year":"2008","journal-title":"Avian Pathol."},{"key":"2026010519421838300_RSBL20210362C29","author":"Therneau","year":"2021"},{"key":"2026010519421838300_RSBL20210362C30","doi-asserted-by":"crossref","DOI":"10.1007\/978-1-4757-3294-8","volume-title":"Modeling survival data: extending the Cox model","author":"Therneau","year":"2000"},{"key":"2026010519421838300_RSBL20210362C31","volume-title":"R: a language and environment for statistical computing, v.4.0.2","author":"R Development Core Team","year":"2020"},{"key":"2026010519421838300_RSBL20210362C32","volume-title":"Model selection and multimodel inference: a practical information-theoretic approach","author":"Burnham","year":"2002"},{"key":"2026010519421838300_RSBL20210362C33","doi-asserted-by":"crossref","first-page":"235","DOI":"10.1111\/j.1461-0248.2011.01730.x","article-title":"Living fast and dying of infection: host life history drives interspecific variation in infection and disease risk","volume":"15","author":"Johnson","year":"2012","journal-title":"Ecol. Lett."},{"key":"2026010519421838300_RSBL20210362C34","doi-asserted-by":"crossref","first-page":"e38920","DOI":"10.1371\/journal.pone.0038920","article-title":"Frequent arousal from hibernation linked to severity of infection and mortality in bats with white-nose syndrome","volume":"7","author":"Reeder","year":"2012","journal-title":"PLoS ONE"},{"key":"2026010519421838300_RSBL20210362C35","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1007\/s00360-017-1109-2","article-title":"Energy conserving thermoregulatory patterns and lower disease severity in a bat resistant to the impacts of white-nose syndrome","volume":"188","author":"Moore","year":"2018","journal-title":"J. Comp. Physiol. B: Biochem. Syst. Environ. Physiol."},{"key":"2026010519421838300_RSBL20210362C36","doi-asserted-by":"crossref","first-page":"525","DOI":"10.7589\/JWD-D-20-00123","article-title":"Using multivariate analyses to explore disease progression of finch mycoplasmosis","volume":"57","author":"Ruden","year":"2021","journal-title":"J. Wildl. Dis."},{"key":"2026010519421838300_RSBL20210362C37","doi-asserted-by":"crossref","unstructured":"Ruden RM , AdelmanJS. 2021Disease tolerance alters host competence in a wild songbird. Figshare.","DOI":"10.1098\/rsbl.2021.0362"}],"container-title":["Biology Letters"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/royalsocietypublishing.org\/doi\/pdf\/10.1098\/rsbl.2021.0362","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/royalsocietypublishing.org\/doi\/full-xml\/10.1098\/rsbl.2021.0362","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/royalsocietypublishing.org\/rsbl\/article-pdf\/doi\/10.1098\/rsbl.2021.0362\/439502\/rsbl.2021.0362.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/royalsocietypublishing.org\/rsbl\/article-pdf\/doi\/10.1098\/rsbl.2021.0362\/439502\/rsbl.2021.0362.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,1,6]],"date-time":"2026-01-06T00:42:27Z","timestamp":1767660147000},"score":1,"resource":{"primary":{"URL":"https:\/\/royalsocietypublishing.org\/rsbl\/article\/17\/10\/20210362\/62841\/Disease-tolerance-alters-host-competence-in-a-wild"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,10,1]]},"references-count":37,"journal-issue":{"issue":"10","published-print":{"date-parts":[[2021,10,1]]}},"URL":"https:\/\/doi.org\/10.1098\/rsbl.2021.0362","relation":{},"ISSN":["1744-957X"],"issn-type":[{"value":"1744-957X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,10,1]]},"assertion":[{"value":"2021-07-02","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2021-10-04","order":1,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2021-10-27","order":2,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"20210362"}}