{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T01:30:16Z","timestamp":1790213416031,"version":"4.1.0"},"reference-count":180,"publisher":"Public Library of Science (PLoS)","issue":"9","license":[{"start":{"date-parts":[[2022,9,7]],"date-time":"2022-09-07T00:00:00Z","timestamp":1662508800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100000024","name":"Canadian Institutes of Health Research","doi-asserted-by":"crossref","award":["OV5-170349"],"award-info":[{"award-number":["OV5-170349"]}],"id":[{"id":"10.13039\/501100000024","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100000024","name":"Canadian Institutes of Health Research","doi-asserted-by":"crossref","award":["VRI-172779"],"award-info":[{"award-number":["VRI-172779"]}],"id":[{"id":"10.13039\/501100000024","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100000024","name":"Canadian Institutes of Health Research","doi-asserted-by":"crossref","award":["OV2-170357"],"award-info":[{"award-number":["OV2-170357"]}],"id":[{"id":"10.13039\/501100000024","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100008920","name":"University of Saskatchewan","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100008920","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100008920","name":"University of Saskatchewan","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100008920","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.plospathogens.org"],"crossmark-restriction":false},"short-container-title":["PLoS Pathog"],"abstract":"<jats:p>Severe Acute Respiratory Syndrome Coronavirus (SARS-CoV-2) can cause the life-threatening acute respiratory disease called COVID-19 (Coronavirus Disease 2019) as well as debilitating multiorgan dysfunction that persists after the initial viral phase has resolved. Long COVID or Post-Acute Sequelae of COVID-19 (PASC) is manifested by a variety of symptoms, including fatigue, dyspnea, arthralgia, myalgia, heart palpitations, and memory issues sometimes affecting between 30% and 75% of recovering COVID-19 patients. However, little is known about the mechanisms causing Long COVID and there are no widely accepted treatments or therapeutics. After introducing the clinical aspects of acute COVID-19 and Long COVID in humans, we summarize the work in animals (mice, Syrian hamsters, ferrets, and nonhuman primates (NHPs)) to model human COVID-19. The virology, pathology, immune responses, and multiorgan involvement are explored. Additionally, any studies investigating time points longer than 14 days post infection (pi) are highlighted for insight into possible long-term disease characteristics. Finally, we discuss how the models can be leveraged for treatment evaluation, including pharmacological agents that are currently in human clinical trials for treating Long COVID. The establishment of a recognized Long COVID preclinical model representing the human condition would allow the identification of mechanisms causing disease as well as serve as a vehicle for evaluating potential therapeutics.<\/jats:p>","DOI":"10.1371\/journal.ppat.1010741","type":"journal-article","created":{"date-parts":[[2022,9,7]],"date-time":"2022-09-07T17:33:09Z","timestamp":1662571989000},"page":"e1010741","update-policy":"https:\/\/doi.org\/10.1371\/journal.ppat.corrections_policy","source":"Crossref","is-referenced-by-count":25,"title":["After the virus has cleared\u2014Can preclinical models be employed for Long COVID research?"],"prefix":"10.1371","volume":"18","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-7212-7487","authenticated-orcid":true,"given":"Ethan B.","family":"Jansen","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Spencer N.","family":"Orvold","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cynthia L.","family":"Swan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anthony","family":"Yourkowski","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Brittany M.","family":"Thivierge","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Magen E.","family":"Francis","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anni","family":"Ge","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Melissa","family":"Rioux","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Joseph","family":"Darbellay","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"John G.","family":"Howland","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0940-0847","authenticated-orcid":true,"given":"Alyson A.","family":"Kelvin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2022,9,7]]},"reference":[{"key":"ppat.1010741.ref001","doi-asserted-by":"crossref","first-page":"104364","DOI":"10.1016\/j.jcv.2020.104364","article-title":"Clinical features and short-term outcomes of 221 patients with COVID-19 in Wuhan","volume":"127","author":"G Zhang","year":"2020","journal-title":"China J Clin Virol"},{"issue":"6","key":"ppat.1010741.ref002","doi-asserted-by":"crossref","first-page":"363","DOI":"10.1038\/s41577-020-0311-8","article-title":"The trinity of COVID-19: immunity, inflammation and intervention","volume":"20","author":"MZ Tay","year":"2020","journal-title":"Nat Rev Immunol"},{"issue":"12","key":"ppat.1010741.ref003","doi-asserted-by":"crossref","first-page":"1100","DOI":"10.1016\/j.it.2020.10.004","article-title":"Mechanisms of SARS-CoV-2 Transmission and Pathogenesis","volume":"41","author":"AG Harrison","year":"2020","journal-title":"Trends Immunol"},{"issue":"2","key":"ppat.1010741.ref004","doi-asserted-by":"crossref","first-page":"194","DOI":"10.1038\/s41590-021-01104-y","article-title":"Pathological sequelae of long-haul COVID","volume":"23","author":"S Mehandru","year":"2022","journal-title":"Nat Immunol"},{"issue":"4","key":"ppat.1010741.ref005","doi-asserted-by":"crossref","first-page":"601","DOI":"10.1038\/s41591-021-01283-z","article-title":"Post-acute COVID-19 syndrome","volume":"27","author":"A Nalbandian","year":"2021","journal-title":"Nat Med"},{"issue":"10270","key":"ppat.1010741.ref006","doi-asserted-by":"crossref","first-page":"220","DOI":"10.1016\/S0140-6736(20)32656-8","article-title":"6-month consequences of COVID-19 in patients discharged from hospital: a cohort study","volume":"397","author":"C Huang","year":"2021","journal-title":"Lancet"},{"key":"ppat.1010741.ref007","doi-asserted-by":"crossref","first-page":"n136","DOI":"10.1136\/bmj.n136","article-title":"Managing the long term effects of covid-19: summary of NICE, SIGN, and RCGP rapid guideline","volume":"372","author":"W Shah","year":"2021","journal-title":"BMJ"},{"key":"ppat.1010741.ref008","doi-asserted-by":"crossref","first-page":"686029","DOI":"10.3389\/fimmu.2021.686029","article-title":"Unraveling the Mystery Surrounding Post-Acute Sequelae of COVID-19","volume":"12","author":"RK Ramakrishnan","year":"2021","journal-title":"Front Immunol"},{"issue":"7862","key":"ppat.1010741.ref009","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1038\/s41586-021-03553-9","article-title":"High-dimensional characterization of post-acute sequelae of COVID-19","volume":"594","author":"Z Al-Aly","year":"2021","journal-title":"Nature"},{"issue":"3","key":"ppat.1010741.ref010","doi-asserted-by":"crossref","first-page":"378","DOI":"10.1016\/j.jinf.2021.01.004","article-title":"Post-acute COVID-19 syndrome. Incidence and risk factors: A Mediterranean cohort study","volume":"82","author":"O Moreno-Perez","year":"2021","journal-title":"J Infect"},{"issue":"3","key":"ppat.1010741.ref011","first-page":"697","article-title":"A comparative study of human betacoronavirus spike proteins: structure, function and therapeutics","volume":"166","author":"J Verma","year":"2021","journal-title":"Arch Virol"},{"key":"ppat.1010741.ref012","article-title":"Signs and symptoms to determine if a patient presenting in primary care or hospital outpatient settings has COVID-19","volume":"2","author":"T Struyf","year":"2021","journal-title":"Cochrane Db Syst Rev"},{"issue":"10","key":"ppat.1010741.ref013","doi-asserted-by":"crossref","first-page":"P19","DOI":"10.1164\/rccm.2020C1","article-title":"Diagnosis and Management of COVID-19 Disease","volume":"201","author":"S Jamil","year":"2020","journal-title":"Am J Respir Crit Care Med"},{"issue":"20","key":"ppat.1010741.ref014","doi-asserted-by":"crossref","first-page":"2089","DOI":"10.1001\/jama.2020.6771","article-title":"Alterations in Smell or Taste in Mildly Symptomatic Outpatients With SARS-CoV-2 Infection","volume":"323","author":"G Spinato","year":"2020","journal-title":"JAMA"},{"issue":"9","key":"ppat.1010741.ref015","doi-asserted-by":"crossref","first-page":"486","DOI":"10.12968\/jowc.2020.29.9.486","article-title":"COVID toes and other cutaneous manifestations of COVID-19","volume":"29","author":"S Rose-Sauld","year":"2020","journal-title":"J Wound Care"},{"issue":"10","key":"ppat.1010741.ref016","doi-asserted-by":"crossref","first-page":"2057","DOI":"10.1002\/acn3.51166","article-title":"Neurological manifestations of coronavirus infections\u2014a systematic review","volume":"7","author":"J Almqvist","year":"2020","journal-title":"Ann Clin Transl Neurol"},{"issue":"7","key":"ppat.1010741.ref017","doi-asserted-by":"crossref","first-page":"1017","DOI":"10.1038\/s41591-020-0968-3","article-title":"Extrapulmonary manifestations of COVID-19","volume":"26","author":"A Gupta","year":"2020","journal-title":"Nat Med"},{"issue":"7825","key":"ppat.1010741.ref018","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1038\/d41586-020-02598-6","article-title":"The lasting misery of coronavirus long-haulers","volume":"585","author":"M. Marshall","year":"2020","journal-title":"Nature"},{"issue":"3","key":"ppat.1010741.ref019","doi-asserted-by":"crossref","first-page":"1013","DOI":"10.1007\/s11357-020-00193-1","article-title":"SARS-CoV-2 and COVID-19 in older adults: what we may expect regarding pathogenesis, immune responses, and outcomes (vol 42, pg 505, 2020)","volume":"42","author":"J Nikolich-Zugich","year":"2020","journal-title":"Geroscience"},{"issue":"17","key":"ppat.1010741.ref020","doi-asserted-by":"crossref","DOI":"10.3390\/jcm10173855","article-title":"Specific Risk Factors for Fatal Outcome in Critically Ill COVID-19 Patients: Results from a European Multicenter Study","volume":"10","author":"D Meintrup","year":"2021","journal-title":"J Clin Med"},{"issue":"5","key":"ppat.1010741.ref021","doi-asserted-by":"crossref","first-page":"750","DOI":"10.1038\/s12276-021-00592-0","article-title":"Type I and III interferon responses in SARS-CoV-2 infection","volume":"53","author":"YM Kim","year":"2021","journal-title":"Exp Mol Med"},{"issue":"5","key":"ppat.1010741.ref022","doi-asserted-by":"crossref","first-page":"1036","DOI":"10.1016\/j.cell.2020.04.026","article-title":"Imbalanced Host Response to SARS-CoV-2 Drives Development of COVID-19","volume":"181","author":"D Blanco-Melo","year":"2020","journal-title":"Cell"},{"issue":"6504","key":"ppat.1010741.ref023","doi-asserted-by":"crossref","first-page":"718","DOI":"10.1126\/science.abc6027","article-title":"Impaired type I interferon activity and inflammatory responses in severe COVID-19 patients","volume":"369","author":"J Hadjadj","year":"2020","journal-title":"Science"},{"issue":"1","key":"ppat.1010741.ref024","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1038\/s41591-020-01202-8","article-title":"Immune determinants of COVID-19 disease presentation and severity","volume":"27","author":"P. Brodin","year":"2021","journal-title":"Nat Med"},{"issue":"7","key":"ppat.1010741.ref025","doi-asserted-by":"crossref","first-page":"1063","DOI":"10.1016\/j.chom.2021.06.009","article-title":"Antibody and B cell responses to SARS-CoV-2 infection and vaccination","volume":"29","author":"K Roltgen","year":"2021","journal-title":"Cell Host Microbe."},{"issue":"18","key":"ppat.1010741.ref026","doi-asserted-by":"crossref","first-page":"1724","DOI":"10.1056\/NEJMoa2026116","article-title":"Humoral Immune Response to SARS-CoV-2 in Iceland","volume":"383","author":"DF Gudbjartsson","year":"2020","journal-title":"N Engl J Med"},{"issue":"7","key":"ppat.1010741.ref027","doi-asserted-by":"crossref","first-page":"1489","DOI":"10.1016\/j.cell.2020.05.015","article-title":"Targets of T Cell Responses to SARS-CoV-2 Coronavirus in Humans with COVID-19 Disease and Unexposed Individuals","volume":"181","author":"A Grifoni","year":"2020","journal-title":"Cell"},{"issue":"6529","key":"ppat.1010741.ref028","article-title":"Immunological memory to SARS-CoV-2 assessed for up to 8 months after infection","volume":"371","author":"JM Dan","year":"2021","journal-title":"Science"},{"issue":"6521","key":"ppat.1010741.ref029","doi-asserted-by":"crossref","first-page":"1227","DOI":"10.1126\/science.abd7728","article-title":"Robust neutralizing antibodies to SARS-CoV-2 infection persist for months","volume":"370","author":"A Wajnberg","year":"2020","journal-title":"Science"},{"key":"ppat.1010741.ref030","doi-asserted-by":"crossref","DOI":"10.3389\/fmicb.2021.698169","article-title":"Long COVID or Post-acute Sequelae of COVID-19 (PASC): An Overview of Biological Factors That May Contribute to Persistent Symptoms","volume":"12","author":"AD Proal","year":"2021","journal-title":"Front Microbiol"},{"issue":"4","key":"ppat.1010741.ref031","doi-asserted-by":"crossref","first-page":"506","DOI":"10.1016\/j.cmi.2020.12.001","article-title":"Long COVID-19-it\u2019s not over until?","volume":"27","author":"D Yelin","year":"2021","journal-title":"Clin Microbiol Infect"},{"key":"ppat.1010741.ref032","author":"World Health Organization","year":"2021","journal-title":"A clinical case definition of post COVID-19 condition by a Delphi consensus"},{"key":"ppat.1010741.ref033","doi-asserted-by":"crossref","first-page":"n1648","DOI":"10.1136\/bmj.n1648","article-title":"Long covid-mechanisms, risk factors, and management","volume":"374","author":"H Crook","year":"2021","journal-title":"BMJ"},{"issue":"5","key":"ppat.1010741.ref034","doi-asserted-by":"crossref","first-page":"2555","DOI":"10.1002\/jmv.26624","article-title":"Chronic COVID syndrome: Need for an appropriate medical terminology for long-COVID and COVID long-haulers","volume":"93","author":"AM Baig","year":"2021","journal-title":"J Med Virol"},{"issue":"2","key":"ppat.1010741.ref035","doi-asserted-by":"crossref","first-page":"e1","DOI":"10.1016\/j.jinf.2020.12.002","article-title":"Clinical, virological and imaging profile in patients with prolonged forms of COVID-19: A cross-sectional study","volume":"82","author":"D Salmon-Ceron","year":"2021","journal-title":"J Infect"},{"key":"ppat.1010741.ref036","doi-asserted-by":"crossref","first-page":"371","DOI":"10.1016\/j.ijid.2020.06.014","article-title":"Multisystem Inflammatory Syndrome in Children (MIS-C) with COVID-19: Insights from simultaneous familial Kawasaki Disease cases","volume":"97","author":"R Ebina-Shibuya","year":"2020","journal-title":"Int J Infect Dis"},{"key":"ppat.1010741.ref037","first-page":"51","article-title":"Multi-system inflammatory syndrome in children & adolescents (MIS-C): A systematic review of clinical features and presentation","volume":"38","author":"T Radia","year":"2021","journal-title":"Paediatr Respir Rev"},{"issue":"8","key":"ppat.1010741.ref038","doi-asserted-by":"crossref","first-page":"453","DOI":"10.1038\/s41577-020-0367-5","article-title":"Understanding SARS-CoV-2-related multisystem inflammatory syndrome in children","volume":"20","author":"AH Rowley","year":"2020","journal-title":"Nat Rev Immunol"},{"key":"ppat.1010741.ref039","article-title":"Immunological dysfunction persists for 8 months following initial mild-to-moderate SARS-CoV-2 infection","author":"C Phetsouphanh","year":"2022","journal-title":"Nat Immunol"},{"key":"ppat.1010741.ref040","doi-asserted-by":"crossref","first-page":"746021","DOI":"10.3389\/fimmu.2021.746021","article-title":"Persistence of SARS CoV-2 S1 Protein in CD16+ Monocytes in Post-Acute Sequelae of COVID-19 (PASC) up to 15 Months Post-Infection","volume":"12","author":"BK Patterson","year":"2021","journal-title":"Front Immunol"},{"issue":"5","key":"ppat.1010741.ref041","article-title":"Patient-reported outcome measures after COVID-19: a prospective cohort study","volume":"56","author":"AW Wong","year":"2020","journal-title":"Eur Respir J"},{"key":"ppat.1010741.ref042","volume-title":"Prevalence of long COVID symptoms and COVID-19 complications","author":"ONS","year":"2020"},{"issue":"11","key":"ppat.1010741.ref043","doi-asserted-by":"crossref","first-page":"1265","DOI":"10.1001\/jamacardio.2020.3557","article-title":"Outcomes of Cardiovascular Magnetic Resonance Imaging in Patients Recently Recovered From Coronavirus Disease 2019 (COVID-19)","volume":"5","author":"VO Puntmann","year":"2020","journal-title":"JAMA Cardiol"},{"issue":"10","key":"ppat.1010741.ref044","doi-asserted-by":"crossref","first-page":"737","DOI":"10.1080\/23744235.2021.1924397","article-title":"Long COVID or post-COVID-19 syndrome: putative pathophysiology, risk factors, and treatments","volume":"53","author":"SJ Yong","year":"2021","journal-title":"Infect Dis (Lond)"},{"key":"ppat.1010741.ref045","doi-asserted-by":"crossref","first-page":"93","DOI":"10.2147\/POR.S316186","article-title":"Risk Predictors and Symptom Features of Long COVID Within a Broad Primary Care Patient Population Including Both Tested and Untested Patients","volume":"12","author":"R Jones","year":"2021","journal-title":"Pragmat Obs Res"},{"issue":"30","key":"ppat.1010741.ref046","doi-asserted-by":"crossref","first-page":"993","DOI":"10.15585\/mmwr.mm6930e1","article-title":"Symptom Duration and Risk Factors for Delayed Return to Usual Health Among Outpatients with COVID-19 in a Multistate Health Care Systems Network\u2014United States, March-June 2020","volume":"69","author":"MW Tenforde","year":"2020","journal-title":"Mmwr-Morbid Mortal W"},{"issue":"4","key":"ppat.1010741.ref047","doi-asserted-by":"crossref","first-page":"626","DOI":"10.1038\/s41591-021-01292-y","article-title":"Attributes and predictors of long COVID","volume":"27","author":"CH Sudre","year":"2021","journal-title":"Nat Med"},{"issue":"7","key":"ppat.1010741.ref048","doi-asserted-by":"crossref","first-page":"379","DOI":"10.1038\/s41574-021-00495-0","article-title":"Long COVID\u2014metabolic risk factors and novel therapeutic management","volume":"17","author":"K Khunti","year":"2021","journal-title":"Nat Rev Endocrinol"},{"issue":"8","key":"ppat.1010741.ref049","article-title":"Implications of Innate Immunity in Post-Acute Sequelae of Non-Persistent Viral Infections","volume":"10","author":"M Hirschenberger","year":"2021","journal-title":"Cell"},{"issue":"1","key":"ppat.1010741.ref050","doi-asserted-by":"crossref","first-page":"9","DOI":"10.3892\/mmr.2020.11127","article-title":"Obesity-a risk factor for increased COVID-19 prevalence, severity and lethality","volume":"22","author":"D Petrakis","year":"2020","journal-title":"Mol Med Rep"},{"issue":"11","key":"ppat.1010741.ref051","doi-asserted-by":"crossref","first-page":"1839","DOI":"10.1093\/infdis\/jiab490","article-title":"Markers of Immune Activation and Inflammation in Individuals With Postacute Sequelae of Severe Acute Respiratory Syndrome Coronavirus 2 Infection","volume":"224","author":"MJ Peluso","year":"2021","journal-title":"J Infect Dis"},{"issue":"5","key":"ppat.1010741.ref052","doi-asserted-by":"crossref","DOI":"10.1136\/jitc-2021-002550","article-title":"Immune profiling of COVID-19: preliminary findings and implications for the pandemic","volume":"9","author":"HT Maecker","year":"2021","journal-title":"J Immunother Cancer"},{"issue":"1","key":"ppat.1010741.ref053","doi-asserted-by":"crossref","first-page":"16","DOI":"10.1038\/s41541-020-00279-z","article-title":"Protective efficacy of a SARS-CoV-2 DNA vaccine in wild-type and immunosuppressed Syrian hamsters","volume":"6","author":"RL Brocato","year":"2021","journal-title":"NPJ Vaccines"},{"issue":"11","key":"ppat.1010741.ref054","doi-asserted-by":"crossref","first-page":"1694","DOI":"10.1038\/s41591-020-1070-6","article-title":"Ad26 vaccine protects against SARS-CoV-2 severe clinical disease in hamsters","volume":"26","author":"LH Tostanoski","year":"2020","journal-title":"Nat Med"},{"issue":"9","key":"ppat.1010741.ref055","doi-asserted-by":"crossref","first-page":"e0257191","DOI":"10.1371\/journal.pone.0257191","article-title":"Vaccinia virus-based vaccines confer protective immunity against SARS-CoV-2 virus in Syrian hamsters","volume":"16","author":"R Kulkarni","year":"2021","journal-title":"PLoS ONE"},{"issue":"1","key":"ppat.1010741.ref056","doi-asserted-by":"crossref","first-page":"5868","DOI":"10.1038\/s41467-021-26178-y","article-title":"ChAdOx1 nCoV-19 (AZD1222) protects Syrian hamsters against SARS-CoV-2 B.1.351 and B.1.1.7","volume":"12","author":"RJ Fischer","year":"2021","journal-title":"Nat Commun"},{"issue":"1","key":"ppat.1010741.ref057","doi-asserted-by":"crossref","first-page":"1735","DOI":"10.1038\/s41467-021-21992-w","article-title":"Favipiravir antiviral efficacy against SARS-CoV-2 in a hamster model","volume":"12","author":"JS Driouich","year":"2021","journal-title":"Nat Commun"},{"issue":"1","key":"ppat.1010741.ref058","doi-asserted-by":"crossref","first-page":"2295","DOI":"10.1038\/s41467-021-22580-8","article-title":"Orally delivered MK-4482 inhibits SARS-CoV-2 replication in the Syrian hamster model","volume":"12","author":"K Rosenke","year":"2021","journal-title":"Nat Commun"},{"issue":"43","key":"ppat.1010741.ref059","doi-asserted-by":"crossref","first-page":"26955","DOI":"10.1073\/pnas.2014441117","article-title":"Favipiravir at high doses has potent antiviral activity in SARS-CoV-2-infected hamsters, whereas hydroxychloroquine lacks activity","volume":"117","author":"SJF Kaptein","year":"2020","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"8","key":"ppat.1010741.ref060","doi-asserted-by":"crossref","first-page":"e14122","DOI":"10.15252\/emmm.202114122","article-title":"Attenuation of clinical and immunological outcomes during SARS-CoV-2 infection by ivermectin","volume":"13","author":"GD de Melo","year":"2021","journal-title":"EMBO Mol Med"},{"issue":"1","key":"ppat.1010741.ref061","doi-asserted-by":"crossref","first-page":"5838","DOI":"10.1038\/s41467-020-19684-y","article-title":"STAT2 signaling restricts viral dissemination but drives severe pneumonia in SARS-CoV-2 infected hamsters","volume":"11","author":"R Boudewijns","year":"2020","journal-title":"Nat Commun"},{"issue":"9","key":"ppat.1010741.ref062","first-page":"2428","article-title":"Simulation of the Clinical and Pathological Manifestations of Coronavirus Disease 2019 (COVID-19) in a Golden Syrian Hamster Model: Implications for Disease Pathogenesis and Transmissibility","volume":"71","author":"JF Chan","year":"2020","journal-title":"Clin Infect Dis"},{"issue":"28","key":"ppat.1010741.ref063","doi-asserted-by":"crossref","first-page":"16587","DOI":"10.1073\/pnas.2009799117","article-title":"Syrian hamsters as a small animal model for SARS-CoV-2 infection and countermeasure development","volume":"117","author":"M Imai","year":"2020","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"1","key":"ppat.1010741.ref064","doi-asserted-by":"crossref","first-page":"503","DOI":"10.1128\/JVI.79.1.503-511.2005","article-title":"Severe acute respiratory syndrome coronavirus infection of golden Syrian hamsters","volume":"79","author":"A Roberts","year":"2005","journal-title":"J Virol"},{"issue":"1","key":"ppat.1010741.ref065","doi-asserted-by":"crossref","first-page":"2673","DOI":"10.1080\/22221751.2020.1858177","article-title":"Defining the Syrian hamster as a highly susceptible preclinical model for SARS-CoV-2 infection","volume":"9","author":"K Rosenke","year":"2020","journal-title":"Emerg Microbes Infect"},{"issue":"4","key":"ppat.1010741.ref066","doi-asserted-by":"crossref","DOI":"10.26508\/lsa.202000886","article-title":"SARS-CoV-2 infection induces protective immunity and limits transmission in Syrian hamsters","volume":"4","author":"P Selvaraj","year":"2021","journal-title":"Life Sci Alliance"},{"issue":"7818","key":"ppat.1010741.ref067","doi-asserted-by":"crossref","first-page":"834","DOI":"10.1038\/s41586-020-2342-5","article-title":"Pathogenesis and transmission of SARS-CoV-2 in golden hamsters","volume":"583","author":"SF Sia","year":"2020","journal-title":"Nature"},{"key":"ppat.1010741.ref068","doi-asserted-by":"crossref","first-page":"618891","DOI":"10.3389\/fmicb.2020.618891","article-title":"SARS-CoV-2 Causes a Systemically Multiple Organs Damages and Dissemination in Hamsters","volume":"11","author":"Z Song","year":"2020","journal-title":"Front Microbiol"},{"key":"ppat.1010741.ref069","doi-asserted-by":"crossref","first-page":"3781","DOI":"10.2147\/JIR.S323026","article-title":"Characterization of Virus Replication, Pathogenesis, and Cytokine Responses in Syrian Hamsters Inoculated with SARS-CoV-2","volume":"14","author":"SJ Yang","year":"2021","journal-title":"J Inflamm Res"},{"issue":"7","key":"ppat.1010741.ref070","doi-asserted-by":"crossref","first-page":"e1009705","DOI":"10.1371\/journal.ppat.1009705","article-title":"SARS-CoV-2 infection in the Syrian hamster model causes inflammation as well as type I interferon dysregulation in both respiratory and non-respiratory tissues including the heart and kidney","volume":"17","author":"ME Francis","year":"2021","journal-title":"PLoS Pathog"},{"issue":"2","key":"ppat.1010741.ref071","doi-asserted-by":"crossref","first-page":"312","DOI":"10.1016\/j.virol.2008.07.026","article-title":"An immunosuppressed Syrian golden hamster model for SARS-CoV infection","volume":"380","author":"SR Schaecher","year":"2008","journal-title":"Virology"},{"issue":"1","key":"ppat.1010741.ref072","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1016\/j.bbrc.2020.03.047","article-title":"Spike protein recognition of mammalian ACE2 predicts the host range and an optimized ACE2 for SARS-CoV-2 infection","volume":"526","author":"J Luan","year":"2020","journal-title":"Biochem Biophys Res Commun"},{"issue":"4","key":"ppat.1010741.ref073","doi-asserted-by":"crossref","first-page":"1058","DOI":"10.1016\/j.cell.2020.09.049","article-title":"A Therapeutic Non-self-reactive SARS-CoV-2 Antibody Protects from Lung Pathology in a COVID-19 Hamster Model","volume":"183","author":"J Kreye","year":"2020","journal-title":"Cell"},{"issue":"1","key":"ppat.1010741.ref074","doi-asserted-by":"crossref","first-page":"192","DOI":"10.1186\/s12879-021-05839-9","article-title":"Correlation between lung infection severity and clinical laboratory indicators in patients with COVID-19: a cross-sectional study based on machine learning","volume":"21","author":"X Wang","year":"2021","journal-title":"BMC Infect Dis"},{"issue":"11","key":"ppat.1010741.ref075","doi-asserted-by":"crossref","first-page":"1009","DOI":"10.1136\/thoraxjnl-2020-215314","article-title":"Respiratory follow-up of patients with COVID-19 pneumonia","volume":"75","author":"PM George","year":"2020","journal-title":"Thorax"},{"issue":"3","key":"ppat.1010741.ref076","doi-asserted-by":"crossref","first-page":"715","DOI":"10.1148\/radiol.2020200370","article-title":"Time Course of Lung Changes at Chest CT during Recovery from Coronavirus Disease 2019 (COVID-19)","volume":"295","author":"F Pan","year":"2020","journal-title":"Radiology"},{"key":"ppat.1010741.ref077","article-title":"SARS-CoV-2 infection in hamsters and humans results in lasting and unique systemic perturbations post recovery","author":"JJ Frere","year":"2022","journal-title":"Sci Transl Med"},{"issue":"1","key":"ppat.1010741.ref078","doi-asserted-by":"crossref","first-page":"6122","DOI":"10.1038\/s41467-020-19891-7","article-title":"Lethality of SARS-CoV-2 infection in K18 human angiotensin-converting enzyme 2 transgenic mice","volume":"11","author":"FS Oladunni","year":"2020","journal-title":"Nat Commun"},{"issue":"1","key":"ppat.1010741.ref079","doi-asserted-by":"crossref","DOI":"10.3390\/v13010132","article-title":"Neuroinvasion and Encephalitis Following Intranasal Inoculation of SARS-CoV-2 in K18-hACE2 Mice","volume":"13","author":"P Kumari","year":"2021","journal-title":"Viruses"},{"key":"ppat.1010741.ref080","first-page":"JVI0151121","article-title":"SARS-CoV-2 causes lung infection without severe disease in human ACE2 knock-in mice","author":"ES Winkler","year":"2021","journal-title":"J Virol"},{"issue":"1","key":"ppat.1010741.ref081","doi-asserted-by":"crossref","first-page":"2433","DOI":"10.1080\/22221751.2020.1838955","article-title":"Comparison of transgenic and adenovirus hACE2 mouse models for SARS-CoV-2 infection","volume":"9","author":"R Rathnasinghe","year":"2020","journal-title":"Emerg Microbes Infect"},{"issue":"11","key":"ppat.1010741.ref082","doi-asserted-by":"crossref","first-page":"1327","DOI":"10.1038\/s41590-020-0778-2","article-title":"SARS-CoV-2 infection of human ACE2-transgenic mice causes severe lung inflammation and impaired function","volume":"21","author":"ES Winkler","year":"2020","journal-title":"Nat Immunol"},{"issue":"7852","key":"ppat.1010741.ref083","doi-asserted-by":"crossref","first-page":"122","DOI":"10.1038\/s41586-021-03361-1","article-title":"SARS-CoV-2 spike D614G change enhances replication and transmission","volume":"592","author":"B Zhou","year":"2021","journal-title":"Nature"},{"issue":"3","key":"ppat.1010741.ref084","doi-asserted-by":"crossref","first-page":"744","DOI":"10.1016\/j.cell.2020.06.011","article-title":"A SARS-CoV-2 Infection Model in Mice Demonstrates Protection by Neutralizing Antibodies","volume":"182","author":"AO Hassan","year":"2020","journal-title":"Cell"},{"issue":"5","key":"ppat.1010741.ref085","doi-asserted-by":"crossref","first-page":"704","DOI":"10.1016\/j.chom.2020.03.023","article-title":"Infection and Rapid Transmission of SARS-CoV-2 in Ferrets","volume":"27","author":"YI Kim","year":"2020","journal-title":"Cell Host Microbe"},{"issue":"1","key":"ppat.1010741.ref086","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1038\/s41467-020-20439-y","article-title":"Dose-dependent response to infection with SARS-CoV-2 in the ferret model and evidence of protective immunity","volume":"12","author":"KA Ryan","year":"2021","journal-title":"Nat Commun"},{"issue":"5","key":"ppat.1010741.ref087","doi-asserted-by":"crossref","first-page":"e218","DOI":"10.1016\/S2666-5247(20)30089-6","article-title":"SARS-CoV-2 in fruit bats, ferrets, pigs, and chickens: an experimental transmission study","volume":"1","author":"K Schlottau","year":"2020","journal-title":"Lancet Microbe"},{"issue":"1","key":"ppat.1010741.ref088","doi-asserted-by":"crossref","first-page":"1260","DOI":"10.1038\/s41467-021-21389-9","article-title":"Comparison of rhesus and cynomolgus macaques as an infection model for COVID-19","volume":"12","author":"FJ Salguero","year":"2021","journal-title":"Nat Commun"},{"issue":"6494","key":"ppat.1010741.ref089","doi-asserted-by":"crossref","first-page":"1012","DOI":"10.1126\/science.abb7314","article-title":"Comparative pathogenesis of COVID-19, MERS, and SARS in a nonhuman primate model","volume":"368","author":"B Rockx","year":"2020","journal-title":"Science"},{"issue":"7824","key":"ppat.1010741.ref090","doi-asserted-by":"crossref","first-page":"268","DOI":"10.1038\/s41586-020-2324-7","article-title":"Respiratory disease in rhesus macaques inoculated with SARS-CoV-2","volume":"585","author":"VJ Munster","year":"2020","journal-title":"Nature"},{"issue":"11","key":"ppat.1010741.ref091","doi-asserted-by":"crossref","first-page":"e1008949","DOI":"10.1371\/journal.ppat.1008949","article-title":"Virulence and pathogenesis of SARS-CoV-2 infection in rhesus macaques: A non-human primate model of COVID-19 progression","volume":"16","author":"H Zheng","year":"2020","journal-title":"PLoS Pathog"},{"issue":"6505","key":"ppat.1010741.ref092","doi-asserted-by":"crossref","first-page":"812","DOI":"10.1126\/science.abc4776","article-title":"SARS-CoV-2 infection protects against rechallenge in rhesus macaques","volume":"369","author":"A Chandrashekar","year":"2020","journal-title":"Science"},{"issue":"1","key":"ppat.1010741.ref093","doi-asserted-by":"crossref","first-page":"86","DOI":"10.1038\/s41590-020-00835-8","article-title":"Establishment of an African green monkey model for COVID-19 and protection against re-infection","volume":"22","author":"C Woolsey","year":"2021","journal-title":"Nat Immunol"},{"issue":"4","key":"ppat.1010741.ref094","doi-asserted-by":"crossref","first-page":"e0097421","DOI":"10.1128\/mBio.00974-21","article-title":"Sex Differences in Lung Imaging and SARS-CoV-2 Antibody Responses in a COVID-19 Golden Syrian Hamster Model","volume":"12","author":"S Dhakal","year":"2021","journal-title":"MBio"},{"issue":"1","key":"ppat.1010741.ref095","doi-asserted-by":"crossref","first-page":"4869","DOI":"10.1038\/s41467-021-25030-7","article-title":"Temporal omics analysis in Syrian hamsters unravel cellular effector responses to moderate COVID-19","volume":"12","author":"G Nouailles","year":"2021","journal-title":"Nat Commun"},{"issue":"3","key":"ppat.1010741.ref096","doi-asserted-by":"crossref","first-page":"557","DOI":"10.1016\/j.immuni.2021.01.017","article-title":"Leveraging the antiviral type I interferon system as a first line of defense against SARS-CoV-2 pathogenicity","volume":"54","author":"DA Hoagland","year":"2021","journal-title":"Immunity"},{"key":"ppat.1010741.ref097","article-title":"SARS-CoV-2 Infected Cardiomyocytes Recruit Monocytes by Secreting CCL2","author":"S Chen","year":"2020","journal-title":"Res Sq"},{"key":"ppat.1010741.ref098","article-title":"Myocarditis in COVID-19 patients: current problems","author":"D Mele","year":"2021","journal-title":"Intern Emerg Med"},{"issue":"3","key":"ppat.1010741.ref099","doi-asserted-by":"crossref","first-page":"263","DOI":"10.1080\/15513815.2020.1761491","article-title":"Fatal Eosinophilic Myocarditis in a Healthy 17-Year-Old Male with Severe Acute Respiratory Syndrome Coronavirus 2 (SARS-CoV-2c)","volume":"39","author":"R Craver","year":"2020","journal-title":"Fetal Pediatr Pathol"},{"key":"ppat.1010741.ref100","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.73522","article-title":"Golden Syrian hamster as a model to study cardiovascular complications associated with SARS-CoV-2 infection","volume":"11","author":"ZA Rizvi","year":"2022","journal-title":"Elife"},{"issue":"596","key":"ppat.1010741.ref101","doi-asserted-by":"crossref","DOI":"10.1126\/scitranslmed.abf8396","article-title":"COVID-19-related anosmia is associated with viral persistence and inflammation in human olfactory epithelium and brain infection in hamsters","volume":"13","author":"GD de Melo","year":"2021","journal-title":"Sci Transl Med"},{"key":"ppat.1010741.ref102","doi-asserted-by":"crossref","first-page":"579","DOI":"10.1016\/j.bbi.2020.06.032","article-title":"Massive transient damage of the olfactory epithelium associated with infection of sustentacular cells by SARS-CoV-2 in golden Syrian hamsters","volume":"89","author":"B Bryche","year":"2020","journal-title":"Brain Behav Immun"},{"issue":"11","key":"ppat.1010741.ref103","doi-asserted-by":"crossref","first-page":"1555","DOI":"10.1021\/acschemneuro.0c00210","article-title":"Expression of the SARS-CoV-2 Entry Proteins, ACE2 and TMPRSS2, in Cells of the Olfactory Epithelium: Identification of Cell Types and Trends with Age","volume":"11","author":"K Bilinska","year":"2020","journal-title":"ACS Chem Nerosci"},{"issue":"1","key":"ppat.1010741.ref104","doi-asserted-by":"crossref","first-page":"21415","DOI":"10.1038\/s41598-020-78402-2","article-title":"SARS-CoV-2 receptor ACE2 is co-expressed with genes related to transmembrane serine proteases, viral entry, immunity and cellular stress","volume":"10","author":"W Wruck","year":"2020","journal-title":"Sci Rep"},{"issue":"2","key":"ppat.1010741.ref105","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1001\/archneur.64.2.185","article-title":"Neuronal damage in brain inflammation","volume":"64","author":"O Aktas","year":"2007","journal-title":"Arch Neurol"},{"issue":"1","key":"ppat.1010741.ref106","doi-asserted-by":"crossref","first-page":"628","DOI":"10.1038\/s41598-021-04622-9","article-title":"Recovery of anosmia in hamsters infected with SARS-CoV-2 is correlated with repair of the olfactory epithelium","volume":"12","author":"RA Reyna","year":"2022","journal-title":"Sci Rep"},{"issue":"33","key":"ppat.1010741.ref107","doi-asserted-by":"crossref","DOI":"10.1126\/sciimmunol.aau6085","article-title":"Structure and function of the immune system in the spleen","volume":"4","author":"SM Lewis","year":"2019","journal-title":"Sci Immunol"},{"key":"ppat.1010741.ref108","doi-asserted-by":"crossref","first-page":"579330","DOI":"10.3389\/fphar.2020.579330","article-title":"Tissue Distribution of ACE2 Protein in Syrian Golden Hamster (Mesocricetus auratus) and Its Possible Implications in SARS-CoV-2 Related Studies","volume":"11","author":"V Suresh","year":"2020","journal-title":"Front Pharmacol"},{"key":"ppat.1010741.ref109","article-title":"Differential susceptibility of SARS-CoV-2 in animals: Evidence of ACE2 host receptor distribution in companion animals, livestock and wildlife by immunohistochemical characterisation","author":"FZX Lean","year":"2021","journal-title":"Transbound Emerg Dis"},{"issue":"1","key":"ppat.1010741.ref110","doi-asserted-by":"crossref","first-page":"2539","DOI":"10.1038\/s41467-022-30195-w","article-title":"SARS-CoV-2 infection induces inflammatory bone loss in golden Syrian hamsters","volume":"13","author":"W Qiao","year":"2022","journal-title":"Nat Commun"},{"issue":"21","key":"ppat.1010741.ref111","doi-asserted-by":"crossref","first-page":"4851","DOI":"10.1111\/bph.15143","article-title":"Animal models of mechanisms of SARS-CoV-2 infection and COVID-19 pathology","volume":"177","author":"SJ Cleary","year":"2020","journal-title":"Br J Pharmacol"},{"issue":"7830","key":"ppat.1010741.ref112","doi-asserted-by":"crossref","first-page":"509","DOI":"10.1038\/s41586-020-2787-6","article-title":"Animal models for COVID-19","volume":"586","author":"C Munoz-Fontela","year":"2020","journal-title":"Nature"},{"issue":"6511","key":"ppat.1010741.ref113","doi-asserted-by":"crossref","first-page":"1603","DOI":"10.1126\/science.abc4730","article-title":"Adaptation of SARS-CoV-2 in BALB\/c mice for testing vaccine efficacy","volume":"369","author":"H Gu","year":"2020","journal-title":"Science"},{"issue":"7830","key":"ppat.1010741.ref114","doi-asserted-by":"crossref","first-page":"560","DOI":"10.1038\/s41586-020-2708-8","article-title":"A mouse-adapted model of SARS-CoV-2 to test COVID-19 countermeasures","volume":"586","author":"KH Dinnon","year":"2020","journal-title":"Nature"},{"issue":"7818","key":"ppat.1010741.ref115","doi-asserted-by":"crossref","first-page":"830","DOI":"10.1038\/s41586-020-2312-y","article-title":"The pathogenicity of SARS-CoV-2 in hACE2 transgenic mice","volume":"583","author":"L Bao","year":"2020","journal-title":"Nature"},{"issue":"1","key":"ppat.1010741.ref116","doi-asserted-by":"crossref","first-page":"e1009195","DOI":"10.1371\/journal.ppat.1009195","article-title":"K18-hACE2 mice develop respiratory disease resembling severe COVID-19","volume":"17","author":"CK Yinda","year":"2021","journal-title":"PLoS Pathog"},{"issue":"2","key":"ppat.1010741.ref117","doi-asserted-by":"crossref","first-page":"813","DOI":"10.1128\/JVI.02012-06","article-title":"Lethal infection of K18-hACE2 mice infected with severe acute respiratory syndrome coronavirus","volume":"81","author":"PB McCray","year":"2007","journal-title":"J Virol"},{"issue":"1","key":"ppat.1010741.ref118","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1016\/j.cell.2020.05.027","article-title":"Pathogenesis of SARS-CoV-2 in Transgenic Mice Expressing Human Angiotensin-Converting Enzyme 2","volume":"182","author":"RD Jiang","year":"2020","journal-title":"Cell"},{"issue":"7843","key":"ppat.1010741.ref119","doi-asserted-by":"crossref","first-page":"603","DOI":"10.1038\/s41586-020-2943-z","article-title":"COVID-19 treatments and pathogenesis including anosmia in K18-hACE2 mice","volume":"589","author":"J Zheng","year":"2021","journal-title":"Nature"},{"issue":"67","key":"ppat.1010741.ref120","doi-asserted-by":"crossref","first-page":"eabl9929","DOI":"10.1126\/sciimmunol.abl9929","article-title":"Administration of aerosolized SARS-CoV-2 to K18-hACE2 mice uncouples respiratory infection from fatal neuroinvasion","volume":"7","author":"V Fumagalli","year":"2022","journal-title":"Sci Immunol"},{"issue":"1","key":"ppat.1010741.ref121","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1093\/ilar.52.1.24","article-title":"The collaborative cross: a recombinant inbred mouse population for the systems genetic era","volume":"52","author":"DW Threadgill","year":"2011","journal-title":"ILAR J"},{"key":"ppat.1010741.ref122","first-page":"1","article-title":"Human-Immune-System (HIS) humanized mouse model (DRAGA: HLA-A2.HLA-DR4.Rag1KO.IL-2RgammacKO.NOD) for COVID-19","author":"TD Brumeanu","year":"2022","journal-title":"Hum Vaccin Immunother"},{"key":"ppat.1010741.ref123","article-title":"A humanized mouse model of chronic COVID-19","author":"E Sefik","year":"2021","journal-title":"Nat Biotechnol"},{"issue":"1","key":"ppat.1010741.ref124","doi-asserted-by":"crossref","first-page":"e1009287","DOI":"10.1371\/journal.ppat.1009287","article-title":"Baseline T cell immune phenotypes predict virologic and disease control upon SARS-CoV infection in Collaborative Cross mice","volume":"17","author":"JB Graham","year":"2021","journal-title":"PLoS Pathog"},{"key":"ppat.1010741.ref125","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1016\/j.virol.2015.03.017","article-title":"Ferret models of viral pathogenesis","volume":"479\u201380","author":"T Enkirch","year":"2015","journal-title":"Virology"},{"issue":"(3","key":"ppat.1010741.ref126","article-title":"The Power of First Impressions: Can Influenza Imprinting during Infancy Inform Vaccine Design?","volume":"8","author":"M Rioux","year":"2020","journal-title":"Vaccines (Basel)"},{"issue":"24","key":"ppat.1010741.ref127","doi-asserted-by":"crossref","first-page":"13187","DOI":"10.1128\/JVI.01456-12","article-title":"Differential pathological and immune responses in newly weaned ferrets are associated with a mild clinical outcome of pandemic 2009 H1N1 infection","volume":"86","author":"SS Huang","year":"2012","journal-title":"J Virol"},{"issue":"10","key":"ppat.1010741.ref128","doi-asserted-by":"crossref","first-page":"e1005173","DOI":"10.1371\/journal.ppat.1005173","article-title":"Influenza Transmission in the Mother-Infant Dyad Leads to Severe Disease, Mammary Gland Infection, and Pathogenesis by Regulating Host Responses","volume":"11","author":"SG Paquette","year":"2015","journal-title":"PLoS Pathog"},{"issue":"12","key":"ppat.1010741.ref129","doi-asserted-by":"crossref","first-page":"2389","DOI":"10.1016\/j.ajpath.2019.08.017","article-title":"Age-Related Pathology Associated with H1N1 A\/California\/07\/2009 Influenza Virus Infection","volume":"189","author":"SJ Bissel","year":"2019","journal-title":"Am J Pathol"},{"issue":"1","key":"ppat.1010741.ref130","first-page":"29","article-title":"The role of cellular susceptibility in the declining severity of respiratory Influenza of ferrets with age","volume":"65","author":"DM Coates","year":"1984","journal-title":"Br J Exp Pathol"},{"issue":"5","key":"ppat.1010741.ref131","first-page":"559","article-title":"The role of naturally-acquired bacterial infection in influenza-related death in neonatal ferrets","volume":"64","author":"RH Husseini","year":"1983","journal-title":"Br J Exp Pathol"},{"issue":"4","key":"ppat.1010741.ref132","doi-asserted-by":"crossref","first-page":"561","DOI":"10.1099\/00222615-13-4-561","article-title":"Studies of influenza virus infection in newborn ferrets","volume":"13","author":"MH Collie","year":"1980","journal-title":"J Med Microbiol"},{"issue":"2","key":"ppat.1010741.ref133","first-page":"113","article-title":"The pregnant ferret as a model for studying the congenital effects of influenza virus infection in utero: infection of foetal tissues in organ culture and in vivo","volume":"58","author":"C Sweet","year":"1977","journal-title":"Br J Exp Pathol"},{"issue":"Pt 6","key":"ppat.1010741.ref134","doi-asserted-by":"crossref","first-page":"1523","DOI":"10.1099\/0022-1317-70-6-1523","article-title":"Mechanism of immunity to Influenza: maternal and passive neonatal protection following immunization of adult ferrets with a live vaccinia-influenza virus haemagglutinin recombinant but not with recombinants containing other influenza virus proteins","volume":"70","author":"KJ Jakeman","year":"1989","journal-title":"J Gen Virol"},{"issue":"3","key":"ppat.1010741.ref135","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1002\/path.1711400302","article-title":"The effects of maternal influenzal viraemia in late gestation on the conceptus of the pregnant ferret","volume":"140","author":"DI Rushton","year":"1983","journal-title":"J Pathol"},{"issue":"4","key":"ppat.1010741.ref136","doi-asserted-by":"crossref","DOI":"10.1128\/mBio.01113-18","article-title":"Moving Forward. Recent Developments for the Ferret Biomedical Research Model","volume":"9","author":"RA Albrecht","year":"2018","journal-title":"mBio"},{"issue":"5","key":"ppat.1010741.ref137","doi-asserted-by":"crossref","first-page":"e0233794","DOI":"10.1371\/journal.pone.0233794","article-title":"Sequencing B cell receptors from ferrets (Mustela putorius furo)","volume":"15","author":"J Wong","year":"2020","journal-title":"PLoS ONE"},{"issue":"1","key":"ppat.1010741.ref138","doi-asserted-by":"crossref","first-page":"1864","DOI":"10.1038\/s41598-021-81389-z","article-title":"Screening and development of monoclonal antibodies for identification of ferret T follicular helper cells","volume":"11","author":"W Jiang","year":"2021","journal-title":"Sci Rep"},{"issue":"4","key":"ppat.1010741.ref139","doi-asserted-by":"crossref","first-page":"1957","DOI":"10.1128\/JVI.02476-12","article-title":"Sequencing, annotation, and characterization of the influenza ferret infectome","volume":"87","author":"AJ Leon","year":"2013","journal-title":"J Virol"},{"issue":"12","key":"ppat.1010741.ref140","doi-asserted-by":"crossref","first-page":"1250","DOI":"10.1038\/nbt.3079","article-title":"The draft genome sequence of the ferret (Mustela putorius furo) facilitates study of human respiratory disease","volume":"32","author":"X Peng","year":"2014","journal-title":"Nat Biotechnol"},{"issue":"43","key":"ppat.1010741.ref141","doi-asserted-by":"crossref","first-page":"26926","DOI":"10.1073\/pnas.2008203117","article-title":"Glycomic analysis of host response reveals high mannose as a key mediator of influenza severity","volume":"117","author":"DW Heindel","year":"2020","journal-title":"Proc Natl Acad Sci U S A"},{"key":"ppat.1010741.ref142","doi-asserted-by":"crossref","first-page":"519","DOI":"10.1007\/978-0-387-33012-9_93","article-title":"Identification of ferret ACE2 and its receptor function for SARS-coronavirus","volume":"581","author":"A Zamoto","year":"2006","journal-title":"Adv Exp Med Biol"},{"issue":"1","key":"ppat.1010741.ref143","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1016\/j.virol.2007.12.032","article-title":"The SARS-CoV ferret model in an infection-challenge study","volume":"374","author":"YK Chu","year":"2008","journal-title":"Virology"},{"issue":"9","key":"ppat.1010741.ref144","doi-asserted-by":"crossref","first-page":"e45842","DOI":"10.1371\/journal.pone.0045842","article-title":"Lack of innate interferon responses during SARS coronavirus infection in a vaccination and reinfection ferret model","volume":"7","author":"MJ Cameron","year":"2012","journal-title":"PLoS ONE"},{"issue":"Pt 9","key":"ppat.1010741.ref145","first-page":"2136","article-title":"Severe acute respiratory syndrome vaccine efficacy in ferrets: whole killed virus and adenovirus-vectored vaccines","volume":"89","author":"RH See","year":"2008","journal-title":"J Gen Virol"},{"issue":"1","key":"ppat.1010741.ref146","doi-asserted-by":"crossref","DOI":"10.1038\/s41467-020-17367-2","article-title":"SARS-CoV-2 is transmitted via contact and via the air between ferrets","volume":"11","author":"M Richard","year":"2020","journal-title":"Nat Commun"},{"issue":"1","key":"ppat.1010741.ref147","doi-asserted-by":"crossref","DOI":"10.1038\/s41467-021-21918-6","article-title":"SARS-CoV and SARS-CoV-2 are transmitted through the air between ferrets over more than one meter distance.","volume":"12","author":"JS Kutter","year":"2021","journal-title":"Nat Commun"},{"key":"ppat.1010741.ref148","article-title":"Male sex and age biases viral burden, viral shedding, and type 1 and 2 interferon responses during SARS-CoV-2 infection in ferrets","author":"M. E. RB Francis","year":"2021","journal-title":"Forthcoming at Sci Rep"},{"issue":"1","key":"ppat.1010741.ref149","doi-asserted-by":"crossref","first-page":"14536","DOI":"10.1038\/s41598-021-93855-9","article-title":"Sex and age bias viral burden and interferon responses during SARS-CoV-2 infection in ferrets","volume":"11","author":"ME Francis","year":"2021","journal-title":"Sci Rep"},{"issue":"1","key":"ppat.1010741.ref150","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1038\/s41467-021-27717-3","article-title":"Age-dependent pathogenic characteristics of SARS-CoV-2 infection in ferrets","volume":"13","author":"YI Kim","year":"2022","journal-title":"Nat Commun"},{"issue":"1","key":"ppat.1010741.ref151","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1002\/ame2.12108","article-title":"Age-related rhesus macaque models of COVID-19","volume":"3","author":"P Yu","year":"2020","journal-title":"Animal Model Exp Med"},{"key":"ppat.1010741.ref152","doi-asserted-by":"crossref","first-page":"754642","DOI":"10.3389\/fimmu.2021.754642","article-title":"Similarities and Differences in the Acute-Phase Response to SARS-CoV-2 in Rhesus Macaques and African Green Monkeys","volume":"12","author":"C Coleman","year":"2021","journal-title":"Front Immunol"},{"key":"ppat.1010741.ref153","article-title":"Distinct upper airway epithelium interferon-stimulated and profibrotic gene expression between adult and infant rhesus macaques infected with SARS-CoV-2","author":"SN Langel","year":"2022","journal-title":"BioRxiv"},{"issue":"8","key":"ppat.1010741.ref154","doi-asserted-by":"crossref","DOI":"10.3390\/v13081673","article-title":"The Post-Acute Phase of SARS-CoV-2 Infection in Two Macaque Species Is Associated with Signs of Ongoing Virus Replication and Pathology in Pulmonary and Extrapulmonary Tissues","volume":"13","author":"KP Boszormenyi","year":"2021","journal-title":"Viruses"},{"issue":"1","key":"ppat.1010741.ref155","doi-asserted-by":"crossref","first-page":"6078","DOI":"10.1038\/s41467-020-19967-4","article-title":"Cellular events of acute, resolving or progressive COVID-19 in SARS-CoV-2 infected non-human primates","volume":"11","author":"MD Fahlberg","year":"2020","journal-title":"Nat Commun"},{"issue":"1","key":"ppat.1010741.ref156","doi-asserted-by":"crossref","DOI":"10.3390\/v13010113","article-title":"Intranasal Infection of Ferrets with SARS-CoV-2 as a Model for Asymptomatic Human Infection","volume":"13","author":"HE Everett","year":"2021","journal-title":"Viruses"},{"issue":"1","key":"ppat.1010741.ref157","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1046\/j.1601-1848.2001.00005.x","article-title":"Behavioral profiles of inbred strains on novel olfactory, spatial and emotional tests for reference memory in mice","volume":"1","author":"A Holmes","year":"2002","journal-title":"Genes Brain Behav"},{"key":"ppat.1010741.ref158","unstructured":"Spencer N, Orvold IB, Kelvin AA, John G. How land Development of spontaneous tests of olfactory, object, and social discrimination behaviour in male golden hamsters: use in rodent models of SARS-CoV-2 infection?. Fourteenth Annual Meeting of the Canadian Association for Neuroscience Virtual Conference, August 2021."},{"issue":"2","key":"ppat.1010741.ref159","doi-asserted-by":"crossref","first-page":"e503","DOI":"10.1093\/cid\/ciaa995","article-title":"Severe Acute Respiratory Syndrome Coronavirus 2 Infects and Damages the Mature and Immature Olfactory Sensory Neurons of Hamsters","volume":"73","author":"AJ Zhang","year":"2021","journal-title":"Clin Infect Dis"},{"issue":"8","key":"ppat.1010741.ref160","doi-asserted-by":"crossref","first-page":"776","DOI":"10.1212\/WNL.0000000000001285","article-title":"Thalamus structure and function determine severity of cognitive impairment in multiple sclerosis","volume":"84","author":"MM Schoonheim","year":"2015","journal-title":"Neurology"},{"issue":"13","key":"ppat.1010741.ref161","doi-asserted-by":"crossref","first-page":"2318","DOI":"10.1089\/neu.2011.1763","article-title":"Thalamus and cognitive impairment in mild traumatic brain injury: a diffusional kurtosis imaging study","volume":"29","author":"EJ Grossman","year":"2012","journal-title":"J Neurotrauma"},{"issue":"2","key":"ppat.1010741.ref162","doi-asserted-by":"crossref","first-page":"277","DOI":"10.1016\/j.ejrad.2010.12.044","article-title":"Changes in thalamus connectivity in mild cognitive impairment: evidence from resting state fMRI","volume":"81","author":"Z Wang","year":"2012","journal-title":"Eur J Radiol"},{"issue":"1","key":"ppat.1010741.ref163","doi-asserted-by":"crossref","first-page":"11605","DOI":"10.1038\/s41598-017-10229-w","article-title":"Lipopolysaccharide-initiated persistent rhinitis causes gliosis and synaptic loss in the olfactory bulb","volume":"7","author":"S Hasegawa-Ishii","year":"2017","journal-title":"Sci Rep"},{"issue":"3","key":"ppat.1010741.ref164","doi-asserted-by":"crossref","first-page":"978","DOI":"10.1016\/j.jaci.2018.09.028","article-title":"Neuroplastic changes in the olfactory bulb associated with nasal inflammation in mice","volume":"143","author":"S Hasegawa-Ishii","year":"2019","journal-title":"J Allergy Clin Immunol"},{"issue":"24","key":"ppat.1010741.ref165","doi-asserted-by":"crossref","first-page":"2512","DOI":"10.1001\/jama.2020.8391","article-title":"Olfactory Dysfunction in COVID-19: Diagnosis and Management","volume":"323","author":"KL Whitcroft","year":"2020","journal-title":"JAMA"},{"issue":"7","key":"ppat.1010741.ref166","doi-asserted-by":"crossref","first-page":"674","DOI":"10.1001\/jamaoto.2020.0832","article-title":"Sudden and Complete Olfactory Loss of Function as a Possible Symptom of COVID-19","volume":"146","author":"M Eliezer","year":"2020","journal-title":"JAMA Otolaryngol Head Neck Surg"},{"key":"ppat.1010741.ref167","doi-asserted-by":"crossref","DOI":"10.1093\/chemse\/bjab044","article-title":"Relationship between depression and olfactory sensory function: a review","volume":"46","author":"A Athanassi","year":"2021","journal-title":"Chem Senses"},{"issue":"6","key":"ppat.1010741.ref168","doi-asserted-by":"crossref","first-page":"479","DOI":"10.1093\/chemse\/bjw061","article-title":"The Association Between Olfaction and Depression: A Systematic Review","volume":"41","author":"P Kohli","year":"2016","journal-title":"Chem Senses"},{"issue":"4","key":"ppat.1010741.ref169","doi-asserted-by":"crossref","first-page":"678","DOI":"10.3390\/v13040678","article-title":"The Intersection of Age and Influenza Severity: Utility of Ferrets for Dissecting the Age-Dependent Immune Responses and Relevance to Age-Specific Vaccine Development","volume":"13","author":"M Rioux","year":"2021","journal-title":"Viruses"},{"key":"ppat.1010741.ref170","article-title":"Historical H1N1 influenza virus imprinting increases vaccine protection by influencing the activity and sustained production of antibodies elicited at vaccination in ferrets","author":"MEMM Francis","year":"2019","journal-title":"Vaccine"},{"key":"ppat.1010741.ref171","doi-asserted-by":"crossref","first-page":"177","DOI":"10.1016\/j.virol.2014.07.013","article-title":"Impaired heterologous immunity in aged ferrets during sequential Influenza A H1N1 infection","volume":"464\u20135","author":"SG Paquette","year":"2014","journal-title":"Virology"},{"issue":"4","key":"ppat.1010741.ref172","article-title":"Historical H1N1 Influenza Virus Imprinting Increases Vaccine Protection by Influencing the Activity and Sustained Production of Antibodies Elicited at Vaccination in Ferrets","volume":"7","author":"ME Francis","year":"2019","journal-title":"Vaccines (Basel)."},{"key":"ppat.1010741.ref173","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1038\/s41541-019-0123-7","article-title":"Repeated vaccination against matched H3N2 influenza virus gives less protection than single vaccination in ferrets","volume":"4","author":"N Music","year":"2019","journal-title":"NPJ Vaccines"},{"issue":"2","key":"ppat.1010741.ref174","doi-asserted-by":"crossref","first-page":"274","DOI":"10.1016\/j.ajpath.2020.10.016","article-title":"Acute Respiratory Distress in Aged, SARS-CoV-2-Infected African Green Monkeys but Not Rhesus Macaques","volume":"191","author":"RV Blair","year":"2021","journal-title":"Am J Pathol"},{"key":"ppat.1010741.ref175","article-title":"Evidence mapping and review of long-COVID and its underlying pathophysiological mechanism","author":"A Umesh","year":"2022","journal-title":"Infection"},{"issue":"8","key":"ppat.1010741.ref176","doi-asserted-by":"crossref","first-page":"825","DOI":"10.1177\/0961203316678675","article-title":"Safety, pharmacokinetics, and pharmacodynamics of RSLV-132, an RNase-Fc fusion protein in systemic lupus erythematosus: a randomized, double-blind, placebo-controlled study","volume":"26","author":"DJ Burge","year":"2017","journal-title":"Lupus"},{"issue":"1","key":"ppat.1010741.ref177","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1002\/art.41489","article-title":"Improvement of Severe Fatigue Following Nuclease Therapy in Patients With Primary Sjogren\u2019s Syndrome: A Randomized Clinical Trial","volume":"73","author":"J Posada","year":"2021","journal-title":"Arthritis Rheumatol"},{"key":"ppat.1010741.ref178","doi-asserted-by":"crossref","first-page":"198088","DOI":"10.1016\/j.virusres.2020.198088","article-title":"Pyruvate affects inflammatory responses of macrophages during Influenza A virus infection","volume":"286","author":"H Abusalamah","year":"2020","journal-title":"Virus Res"},{"issue":"3","key":"ppat.1010741.ref179","article-title":"Inhalation of Sodium Pyruvate to Reduce the Symptoms and Severity of Respiratory Diseases Including COVID-19, Long COVID, and Pulmonary Fibrosis","volume":"3","author":"CRNR Lupfer","year":"2021","journal-title":"Eur J Respir Med"},{"issue":"6","key":"ppat.1010741.ref180","doi-asserted-by":"crossref","DOI":"10.3390\/nu12061616","article-title":"Nicotinamide Riboside-The Current State of Research and Therapeutic Uses","volume":"12","author":"M Mehmel","year":"2020","journal-title":"Nutrients"}],"container-title":["PLOS Pathogens"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.ppat.1010741","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,11,26]],"date-time":"2023-11-26T18:01:41Z","timestamp":1701021701000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.ppat.1010741"}},"subtitle":[],"editor":[{"given":"Seema","family":"Lakdawala","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2022,9,7]]},"references-count":180,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2022,9,7]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.ppat.1010741","relation":{},"ISSN":["1553-7374"],"issn-type":[{"value":"1553-7374","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,9,7]]}}}