{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,25]],"date-time":"2026-09-25T01:13:43Z","timestamp":1790298823401,"version":"4.1.0"},"reference-count":35,"publisher":"Springer Science and Business Media LLC","issue":"7940","license":[{"start":{"date-parts":[[2022,11,2]],"date-time":"2022-11-02T00:00:00Z","timestamp":1667347200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"},{"start":{"date-parts":[[2022,11,2]],"date-time":"2022-11-02T00:00:00Z","timestamp":1667347200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Nature"],"published-print":{"date-parts":[[2022,12,15]]},"DOI":"10.1038\/s41586-022-05482-7","type":"journal-article","created":{"date-parts":[[2022,11,2]],"date-time":"2022-11-02T07:19:32Z","timestamp":1667373572000},"page":"540-545","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":105,"title":["Characterization of SARS-CoV-2 Omicron BA.4 and BA.5 isolates in rodents"],"prefix":"10.1038","volume":"612","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-0890-1922","authenticated-orcid":false,"given":"Ryuta","family":"Uraki","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Peter J.","family":"Halfmann","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2258-9031","authenticated-orcid":false,"given":"Shun","family":"Iida","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7768-5157","authenticated-orcid":false,"given":"Seiya","family":"Yamayoshi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuri","family":"Furusawa","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maki","family":"Kiso","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mutsumi","family":"Ito","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8266-020X","authenticated-orcid":false,"given":"Kiyoko","family":"Iwatsuki-Horimoto","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sohtaro","family":"Mine","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Makoto","family":"Kuroda","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tadashi","family":"Maemura","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuko","family":"Sakai-Tagawa","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6557-0771","authenticated-orcid":false,"given":"Hiroshi","family":"Ueki","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rong","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yanan","family":"Liu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Deanna","family":"Larson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shuetsu","family":"Fukushi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shinji","family":"Watanabe","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3488-5439","authenticated-orcid":false,"given":"Ken","family":"Maeda","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3248-1761","authenticated-orcid":false,"given":"Andrew","family":"Pekosz","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3253-6961","authenticated-orcid":false,"given":"Ahmed","family":"Kandeil","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4397-7132","authenticated-orcid":false,"given":"Richard J.","family":"Webby","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2441-4729","authenticated-orcid":false,"given":"Zhongde","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6988-1975","authenticated-orcid":false,"given":"Masaki","family":"Imai","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tadaki","family":"Suzuki","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5061-8296","authenticated-orcid":false,"given":"Yoshihiro","family":"Kawaoka","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2022,11,2]]},"reference":[{"key":"5482_CR1","doi-asserted-by":"publisher","first-page":"687","DOI":"10.1038\/s41586-022-04441-6","volume":"603","author":"PJ Halfmann","year":"2022","unstructured":"Halfmann, P. J. et al. SARS-CoV-2 Omicron virus causes attenuated disease in mice and hamsters. Nature 603, 687\u2013692 (2022).","journal-title":"Nature"},{"key":"5482_CR2","doi-asserted-by":"publisher","first-page":"693","DOI":"10.1038\/s41586-022-04442-5","volume":"603","author":"H Shuai","year":"2022","unstructured":"Shuai, H. et al. Attenuated replication and pathogenicity of SARS-CoV-2 B.1.1.529 Omicron. Nature 603, 693\u2013699 (2022).","journal-title":"Nature"},{"key":"5482_CR3","doi-asserted-by":"publisher","first-page":"700","DOI":"10.1038\/s41586-022-04462-1","volume":"603","author":"R Suzuki","year":"2022","unstructured":"Suzuki, R. et al. Attenuated fusogenicity and pathogenicity of SARS-CoV-2 Omicron variant. Nature 603, 700\u2013705 (2022).","journal-title":"Nature"},{"key":"5482_CR4","doi-asserted-by":"publisher","first-page":"119","DOI":"10.1038\/s41586-022-04856-1","volume":"607","author":"R Uraki","year":"2022","unstructured":"Uraki, R. et al. Characterization and antiviral susceptibility of SARS-CoV-2 Omicron\/BA.2. Nature 607, 119\u2013127 (2022).","journal-title":"Nature"},{"key":"5482_CR5","doi-asserted-by":"publisher","unstructured":"Tegally, H. et al. Emergence of SARS-CoV-2 Omicron lineages BA.4 and BA.5 in South Africa. Nat. Med. https:\/\/doi.org\/10.1038\/s41591-022-01911-2 (2022).","DOI":"10.1038\/s41591-022-01911-2"},{"key":"5482_CR6","doi-asserted-by":"publisher","first-page":"1933","DOI":"10.1038\/s41591-022-01887-z","volume":"28","author":"JA Lewnard","year":"2022","unstructured":"Lewnard, J. A. et al. Clinical outcomes associated with SARS-CoV-2 Omicron (B.1.1.529) variant and BA.1\/BA.1.1 or BA.2 subvariant infection in southern California. Nat. Med. 28, 1933\u20131943 (2022).","journal-title":"Nat. Med."},{"key":"5482_CR7","doi-asserted-by":"publisher","first-page":"437","DOI":"10.1016\/S0140-6736(22)00017-4","volume":"399","author":"N Wolter","year":"2022","unstructured":"Wolter, N. et al. Early assessment of the clinical severity of the SARS-CoV-2 Omicron variant in South Africa: a data linkage study. Lancet 399, 437\u2013446 (2022).","journal-title":"Lancet"},{"key":"5482_CR8","doi-asserted-by":"publisher","first-page":"1286","DOI":"10.1001\/jama.2022.2274","volume":"327","author":"AC Ulloa","year":"2022","unstructured":"Ulloa, A. C., Buchan, S. A., Daneman, N. & Brown, K. A. Estimates of SARS-CoV-2 Omicron variant severity in Ontario, Canada. JAMA 327, 1286\u20131288 (2022).","journal-title":"JAMA"},{"key":"5482_CR9","doi-asserted-by":"publisher","first-page":"1124","DOI":"10.1016\/j.chom.2021.06.006","volume":"29","author":"C Motozono","year":"2021","unstructured":"Motozono, C. et al. SARS-CoV-2 spike L452R variant evades cellular immunity and increases infectivity. Cell Host Microbe 29, 1124\u20131136.e11 (2021).","journal-title":"Cell Host Microbe"},{"key":"5482_CR10","doi-asserted-by":"publisher","first-page":"76","DOI":"10.1038\/s41392-022-00941-z","volume":"7","author":"Y Zhang","year":"2022","unstructured":"Zhang, Y. et al. SARS-CoV-2 spike L452R mutation increases Omicron variant fusogenicity and infectivity as well as host glycolysis. Signal Transduct. Target Ther. 7, 76 (2022).","journal-title":"Signal Transduct. Target Ther."},{"key":"5482_CR11","doi-asserted-by":"publisher","first-page":"3426","DOI":"10.1016\/j.cell.2021.04.025","volume":"184","author":"X Deng","year":"2021","unstructured":"Deng, X. et al. Transmission, infectivity, and neutralization of a spike L452R SARS-CoV-2 variant. Cell 184, 3426\u20133437.e38 (2021).","journal-title":"Cell"},{"key":"5482_CR12","doi-asserted-by":"publisher","first-page":"16587","DOI":"10.1073\/pnas.2009799117","volume":"117","author":"M Imai","year":"2020","unstructured":"Imai, M. et al. Syrian hamsters as a small animal model for SARS-CoV-2 infection and countermeasure development. Proc. Natl Acad. Sci. USA 117, 16587\u201316595 (2020).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"5482_CR13","doi-asserted-by":"publisher","first-page":"834","DOI":"10.1038\/s41586-020-2342-5","volume":"583","author":"SF Sia","year":"2020","unstructured":"Sia, S. F. et al. Pathogenesis and transmission of SARS-CoV-2 in golden hamsters. Nature 583, 834\u2013838 (2020).","journal-title":"Nature"},{"key":"5482_CR14","doi-asserted-by":"publisher","first-page":"2428","DOI":"10.1093\/cid\/ciaa644","volume":"71","author":"JF Chan","year":"2020","unstructured":"Chan, J. F. et al. 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. Clin. Infect. Dis. 71, 2428\u20132446 (2020).","journal-title":"Clin. Infect. Dis."},{"key":"5482_CR15","doi-asserted-by":"publisher","unstructured":"Gilliland, T. et al. Protection of human ACE2 transgenic Syrian hamsters from SARS CoV-2 variants by human polyclonal IgG from hyper-immunized transchromosomic bovines. Preprint at bioRxiv https:\/\/doi.org\/10.1101\/2021.07.26.453840 (2021).","DOI":"10.1101\/2021.07.26.453840"},{"key":"5482_CR16","doi-asserted-by":"publisher","first-page":"813","DOI":"10.1128\/JVI.02012-06","volume":"81","author":"PB McCray Jr.","year":"2007","unstructured":"McCray, P. B. Jr. et al. Lethal infection of K18-hACE2 mice infected with severe acute respiratory syndrome coronavirus. J. Virol. 81, 813\u2013821 (2007).","journal-title":"J. Virol."},{"key":"5482_CR17","doi-asserted-by":"publisher","first-page":"1327","DOI":"10.1038\/s41590-020-0778-2","volume":"21","author":"ES Winkler","year":"2020","unstructured":"Winkler, E. S. et al. SARS-CoV-2 infection of human ACE2-transgenic mice causes severe lung inflammation and impaired function. Nat. Immunol. 21, 1327\u20131335 (2020).","journal-title":"Nat. Immunol."},{"key":"5482_CR18","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-31200-y","volume":"13","author":"F Armando","year":"2022","unstructured":"Armando, F. et al. SARS-CoV-2 Omicron variant causes mild pathology in the upper and lower respiratory tract of hamsters. Nat. Commun. 13, 3519 (2022).","journal-title":"Nat. Commun."},{"key":"5482_CR19","doi-asserted-by":"publisher","first-page":"706","DOI":"10.1038\/s41586-022-04474-x","volume":"603","author":"B Meng","year":"2022","unstructured":"Meng, B. et al. Altered TMPRSS2 usage by SARS-CoV-2 Omicron impacts infectivity and fusogenicity. Nature 603, 706\u2013714 (2022).","journal-title":"Nature"},{"key":"5482_CR20","doi-asserted-by":"publisher","first-page":"271","DOI":"10.1016\/j.cell.2020.02.052","volume":"181","author":"M Hoffmann","year":"2020","unstructured":"Hoffmann, M. et al. SARS-CoV-2 cell entry depends on ACE2 and TMPRSS2 and is blocked by a clinically proven protease inhibitor. Cell 181, 271\u2013280.e8 (2020).","journal-title":"Cell"},{"key":"5482_CR21","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1038\/s41580-021-00418-x","volume":"23","author":"CB Jackson","year":"2022","unstructured":"Jackson, C. B., Farzan, M., Chen, B. & Choe, H. Mechanisms of SARS-CoV-2 entry into cells. Nat. Rev. Mol. Cell Biol. 23, 3\u201320 (2022).","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"5482_CR22","doi-asserted-by":"publisher","first-page":"100847","DOI":"10.1016\/j.jbc.2021.100847","volume":"297","author":"JP Evans","year":"2021","unstructured":"Evans, J. P. & Liu, S. L. Role of host factors in SARS-CoV-2 entry. J. Biol. Chem. 297, 100847 (2021).","journal-title":"J. Biol. Chem."},{"key":"5482_CR23","doi-asserted-by":"publisher","first-page":"134","DOI":"10.1038\/s41392-021-00558-8","volume":"6","author":"MM Zhao","year":"2021","unstructured":"Zhao, M. M. et al. Cathepsin L plays a key role in SARS-CoV-2 infection in humans and humanized mice and is a promising target for new drug development. Signal Transduct. Target Ther. 6, 134 (2021).","journal-title":"Signal Transduct. Target Ther."},{"key":"5482_CR24","doi-asserted-by":"publisher","first-page":"715","DOI":"10.1038\/s41586-022-04479-6","volume":"603","author":"KPY Hui","year":"2022","unstructured":"Hui, K. P. Y. et al. SARS-CoV-2 Omicron variant replication in human bronchus and lung ex vivo. Nature 603, 715\u2013720 (2022).","journal-title":"Nature"},{"key":"5482_CR25","doi-asserted-by":"crossref","unstructured":"Kimura, I. et al. Virological characteristics of the SARS-CoV-2 Omicron BA.2 subvariants, including BA.4 and BA.5 Cell 185, 3992\u20134007.e16 (2022).","DOI":"10.1016\/j.cell.2022.09.018"},{"key":"5482_CR26","doi-asserted-by":"publisher","unstructured":"Davies, M.-A. et al. Outcomes of laboratory-confirmed SARS-CoV-2 infection during resurgence driven by Omicron lineages BA.4 and BA.5 compared with previous waves in the Western Cape Province, South Africa. Preprint at medRxiv https:\/\/doi.org\/10.1101\/2022.06.28.22276983 (2022).","DOI":"10.1101\/2022.06.28.22276983"},{"key":"5482_CR27","doi-asserted-by":"publisher","unstructured":"Lewnard, J. A., Hong, V. & Tartof, S. Y. Association of SARS-CoV-2 BA.4\/BA.5 Omicron lineages with immune escape and clinical outcome. Preprint at medRxiv https:\/\/doi.org\/10.1101\/2022.07.31.22278258 (2022).","DOI":"10.1101\/2022.07.31.22278258"},{"key":"5482_CR28","doi-asserted-by":"publisher","DOI":"10.21203\/rs.3.rs-1792132\/v1","author":"N Wolter","year":"2022","unstructured":"Wolter, N. et al. Clinical severity of SARS-CoV-2 Omicron BA.4 and BA.5 lineages in South Africa. Res. Sq. https:\/\/doi.org\/10.21203\/rs.3.rs-1792132\/v1 (2022).","journal-title":"Res. Sq."},{"key":"5482_CR29","doi-asserted-by":"publisher","first-page":"7001","DOI":"10.1073\/pnas.2002589117","volume":"117","author":"S Matsuyama","year":"2020","unstructured":"Matsuyama, S. et al. Enhanced isolation of SARS-CoV-2 by TMPRSS2-expressing cells. Proc. Natl Acad. Sci. USA 117, 7001\u20137003 (2020).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"5482_CR30","doi-asserted-by":"publisher","first-page":"1475","DOI":"10.1056\/NEJMc2201933","volume":"386","author":"E Takashita","year":"2022","unstructured":"Takashita, E. et al. Efficacy of antiviral agents against the SARS-CoV-2 Omicron subvariant BA.2. N. Engl. J. Med. 386, 1475\u20131477 (2022).","journal-title":"N. Engl. J. Med."},{"key":"5482_CR31","doi-asserted-by":"publisher","first-page":"113","DOI":"10.1016\/j.cell.2021.12.002","volume":"185","author":"M Gagne","year":"2022","unstructured":"Gagne, M. et al. Protection from SARS-CoV-2 delta one year after mRNA-1273 vaccination in rhesus macaques coincides with anamnestic antibody response in the lung. Cell 185, 113\u2013130.e15 (2022).","journal-title":"Cell"},{"key":"5482_CR32","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1095\/biolreprod.113.116905","volume":"90","author":"Z Li","year":"2014","unstructured":"Li, Z. et al. Generation of transgenic pigs by cytoplasmic injection of piggyBac transposase-based pmGENIE-3 plasmids. Biol. Reprod. 90, 93 (2014).","journal-title":"Biol. Reprod."},{"key":"5482_CR33","doi-asserted-by":"crossref","unstructured":"Quick, J. nCoV-2019 sequencing protocol. Protocol.io https:\/\/www.protocols.io\/view\/ncov-2019-sequencing-protocol-v3-locost-bh42j8ye?version_warning=no (2020).","DOI":"10.17504\/protocols.io.bbmuik6w"},{"key":"5482_CR34","doi-asserted-by":"publisher","first-page":"e0239403","DOI":"10.1371\/journal.pone.0239403","volume":"15","author":"K Itokawa","year":"2020","unstructured":"Itokawa, K., Sekizuka, T., Hashino, M., Tanaka, R. & Kuroda, M. Disentangling primer interactions improves SARS-CoV-2 genome sequencing by multiplex tiling PCR. PLoS ONE 15, e0239403 (2020).","journal-title":"PLoS ONE"},{"key":"5482_CR35","doi-asserted-by":"crossref","unstructured":"Itokawa, K. et al. nCoV-2019 sequencing protocol for illumina V.5. Protocol.io https:\/\/www.protocols.io\/view\/ncov-2019-sequencing-protocol-for-illumina-b2msqc6e?version_warning=no (2021).","DOI":"10.17504\/protocols.io.betejeje"}],"container-title":["Nature"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41586-022-05482-7.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41586-022-05482-7","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41586-022-05482-7.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,12,14]],"date-time":"2022-12-14T11:04:27Z","timestamp":1671015867000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41586-022-05482-7"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,11,2]]},"references-count":35,"journal-issue":{"issue":"7940","published-print":{"date-parts":[[2022,12,15]]}},"alternative-id":["5482"],"URL":"https:\/\/doi.org\/10.1038\/s41586-022-05482-7","relation":{"has-preprint":[{"id-type":"doi","id":"10.21203\/rs.3.rs-1820048\/v1","asserted-by":"object"}]},"ISSN":["0028-0836","1476-4687"],"issn-type":[{"value":"0028-0836","type":"print"},{"value":"1476-4687","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,11,2]]},"assertion":[{"value":"3 July 2022","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"25 October 2022","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"2 November 2022","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Y.K. has received unrelated funding support from Daiichi Sankyo Pharmaceutical, Toyama Chemical, Tauns Laboratories, Inc., Shionogi & Co. LTD, Otsuka Pharmaceutical, KM Biologics, Kyoritsu Seiyaku, Shinya Corporation and Fuji Rebio. The remaining authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}},{"value":"This content has been made available to all.","name":"free","label":"Free to read"}]}}