Samarasekera, U. (2021). India grapples with second wave of COVID-19. *Lancet Microbe* 2:e238. doi: 10.1016/S2666-5247(21)00123-3

Shang, J., Ye, G., Shi, K., Wan, Y., Luo, C., Aihara, H., et al. (2020). Structural basis of receptor recognition by SARS-CoV-2. *Nature* 581, 221–224. doi: 10.1038/s41586-020-2179-y

Shen, X., Tang, H., Pajon, R., Smith, G., Glenn, G. M., Shi, W., et al. (2021). Neutralization of SARS-CoV-2 variants B1.429 and B1.351. *New Engl. J. Med.* 384:24. doi: 10.1056/NEJMc2103740

Sheward, D. J., Kim, C., Ehling, R. A., Pankow, A., Castro Dopico, X., Dyrdak, R., et al. (2022). Neutralisation sensitivity of the SARS-CoV-2 omicrom (.B1.1.529) variant: a cross-sectional study. *Lancet Infect. Dis.* 22, 813–820. doi: 10.1016/S1473-3099(22)00129-3

Shi, J., Wen, Z., Zhong, G., Yang, H., Wang, C., Huang, B., et al. (2020). Susceptibility of ferrets, cats, dogs, and other domesticated animals to SARS-coronavirus-2. *Science* 368, 1016–1020. doi: 10.1126/science.abb7015

Shukla, N., Roelle, S. M., Suzart, V. G., Bruchez, A. M., and Matreyek, K. A. (2021). Mutants of human ACE2 differentially promote SARS-CoV and SARS-CoV-2 spike mediated infection. *PLoS Pathog.* 17:e1009715. doi: 10.1371/journal.ppat.1009715

Simas, P. V. M., de Souza Barnabé, A. C., Durães-Carvalho, R., de Lima, F., Neto, D., Caserta, L., et al. (2015). Bat coronavirus in Brazil related to appalachian ridge and porcine epidemic diarrhea viruses. *Emerg. Infect. Dis.* 21, 729–731. doi: 10.3201/eid2104.141783

Snouwaert, J. N., Jania, L. A., Nguyen, T., Martinez, D. R., Schäfer, A., Catanzaro, N. J., et al. (2023). Human ACE2 expression, a major tropism determinant for SARS-CoV-2, is regulated by upstream and intragenic elements. *PLoS Pathog.* 19:e1011168. doi: 10.1371/journal.ppat.1011168

Song, H. D., Tu, C. C., Zhang, G. W., Wang, S. Y., Zheng, K., Lei, L. C., et al. (2005). (2005) cross-host evolution of severe acute respiratory syndrome coronavirus in palm civet and human. *Proc. Natl. Acad. Sci. U. S. A.* 102, 2430–2435. doi: 10.1073/pnas.0409608102

Subbarao, K., McAuliffe, J., Vogel, L., Fahle, G., Fischer, S., Tatti, K., et al. (2004). Prior infection and passive transfer of neutralizing antibody prevent replication of severe acute respiratory syndrome coronavirus in the respiratory tract of mice. *J. Virol.* 78, 3572–3537. doi: 10.1128/JVI.78.7.3572-3577.2004

Sun, F., Wang, X., Tan, S., Dan, Y., Lu, Y., Zhang, J., et al. SARS-CoV-2 quasispecies provides an advantage mutation pool for the epidemic variants. *Microbiol Spectr.* (2021) 9:e00261-e00221. doi: 10.1128/Spectrum.00261-21

Suryamohan, K., Diwanji, D., Stawiski, E. W., Gupta, R., Miersch, S., Liu, J., et al. (2021). Human ACE2 receptor polymorphisms and altered susceptibility to SARS-CoV-2. *Comm. Biol.* 4:475. doi: 10.1038/s42003-021-02030-3

Takada, K., Takahashi Ueda, M., Shichinohe, S., Kida, Y., Ono, C., Matsuura, Y., et al. (2023). Genomic diversity of SARS-CoV-2 can be accelerated by a mutation in the nsp14 gene. *iScience* 26:106210. doi: 10.1016/j.isci.2023.106210

Tang, J. W., Tambyah, P. A., and Hui, D. S. (2020). Emergence of a new SARS-CoV-2 variant in the UK. *J. Infect.* 82, e27–e28. doi: 10.1016/j.jinf.2020.12.024

Tegally, H., Wilkinson, E., Giovanetti, M., Iranzadeh, A., Fonseca, V., Giandhari, J., et al. (2021). Detection of a SARS-CoV-2 variant of concern in South Africa. *Nature* 592, 438–443. doi: 10.1038/s41586-021-03402-9

Thye, A. Y. K., Law, J. W. F., Pusparajah, P., Letchumanan, V., Chan, K. G., and Lee, L. H. (2021). Emerging SARS-CoV-2 variants of concern (VOCs): An impending global crisis. *Biomedicine* 9:1303. doi: 10.3390/biomedicines9101303

Towler, P., Staker, B., Prasad, S. G., Menon, S., Tang, J., Parsons, T., et al. (2004). ACE2 X-ray structures reveal a large hinge-bending motion important for inhibitor binding and catalysis. *J. Biol. Chem.* 279, 17996–18007. doi: 10.1074/jbc.M311191200

Turner, A. J., and Hooper, N. M. (2002). The angiotensin-converting enzyme gene family: genomics and pharmacology. *Trends Pharmacol. Sci.* 23, 177–183. doi: 10.1016/
s0165-6147(00)01994-5

Wacharalpluesadee, S., Tan, C. W., Maneeorn, P., Duengkae, P., Zhu, F., Joyjinda, Y., et al. (2021). Evidence for SARS-CoV-2 related coronaviruses circulating in bats and pangolins in Southeast Asia. *Nature Com* 12:972. doi: 10.1038/s41467-021-21240-1

Wang, Q., Iketani, S., Li, Z., Liu, L., Guo, Y., Huang, Y., et al. (2023). Alarming antibody evasion properties of rising SARS-CoV-2 BQ and XBB subvariants. *Cells* 186, 1–8. doi: 10.1016/j.cell.2022.12.018

Wei, C., Shan, K. J., Wang, W., Zhang, S., Huan, Q., and Qian, W. (2021). Evidence for a mouse origin of the SARS-CoV-2 omicron variant. *J. Genet. Genom.* 48, 1111–1121. doi: 10.1016/j.jgg.2021.12.003

Weissman, D., Alameh, M. G., de Silva, T., Collini, P., Hornsby, H., Brown, R., et al. (2021). D614G spike mutation increases SARS-CoV-2 susceptibility to neutralization. *Cell Host Microbe* 29, 23–31.e4. doi: 10.1016/j.chom.2020.11.012

Wentworth, D. E., Gillim-Ross, L., Espina, N., and Bernard, K. A. (2004). Mice susceptible to SARS coronavirus. *Emerg. Infect. Dis.* 10, 1293–1296. doi: 10.3201/eid1007.031119

Wertheim, J. O., Chu, D. K., Peiris, J. S., Kosakovsky Pond, S. L., and Poon, L. L. (2013). A case for the ancient origin of coronaviruses. *J. Virol* 87, 7039–7045. doi: 10.1128/JVI.03273-12

Woo, P. C., Lau, S. K., Lam, C. S., Lau, C. C., Tsang, A. K., Lau, J. H., et al. (2012). Discovery of seven novel mammalian and avian coronaviruses in the genus deltacoronavirus supports bat coronaviruses as the gene source of alphacoronavirus and betacoronavirus and avian coronaviruses as the gene source of gamma-coronavirus and delta-coronavirus. *J. Virol.* 86, 3995–4008. doi: 10.1128/JVI.06540-11

Xu, D., Zhang, Z., and Wang, F. S. (2004). SARS-associated coronavirus quasispecies in individual patients. *N. Engl. J. Med.* 350, 1366–1367. doi: 10.1056/NEJMc032421

Yan, R., Zhang, Y., Li, Y., Xia, L., Guo, Y., Zhou, Q., et al. (2020). Structural basis for the recognition of the SARS-CoV-2 by full-length human ACE2. *Science* 367, 1444–1448. doi: 10.1126/science.abb2762

Yang, M., Zhao, J., Xing, L., and Shi, L. (2015). The association between angiotensinconverting enzyme 2 polymorphisms and essential hypertension risk: a meta-analysis involving 14,122 patients. *J. Renin-Angiotensin-Aldosterone Syst.* 16, 1240–1244. doi: 10.1177/1470320314549221

Yin, X., Popa, H., Stapon, A., Bouda, E., and Garcia-Diaz, M. (2023). Fidelity of ribocleotide incorporation by the SARS-CoV-2 replication complex. *J. Mol. Biol.* 435:167973. doi: 10.1016/j.jmb.2023.167973

Yue, C., Li, Z., Lin, Y., Yang, Y., Yuan, M., Pan, Z., et al. (2021). Sensitivity of SARS-CoV-2 variants to neutralization by convalescent sera and a VH3-30 monoclonal antibody. *Front. Immunol.* 12:751584. doi: 10.3389/fimmu.2021.751584

Yue, C., Song, W., Wang, L., Jian, F., Chen, X., Gao, F., et al. (2023). ACE2 binding and antibody evasion in enhanced transmissibility of XBB.1.5. *Lancet Inf. Dis.* 23, 278–280. doi: 10.1016/S1473-3099(23)00010-5

Zhou, H., Chen, X., Hu, T., Li, J., Song, H., Liu, Y., et al. (2020). A novel bat coronavirus closely related to SARS-CoV-2 contains natural insertions at the S1/S2 cleavage site of the spike protein. *Curr. Biol.* 30, 2196–2203. doi: 10.1016/j.cub.2020.05.023

Zhou, P., Fan, H., Lan, T., Yang, X. L., Shi, W. F., Zhang, W., et al. (2018). Fatal swine acute diarrhoea syndrome caused by an HKU2-related coronavirus of bat origin. *Nature* 556, 255–258. doi: 10.1038/s41586-018-0010-9

Zhou, P., Yang, X. L., Wang, X. G., Hu, B., Zhang, L., Zhang, W., et al. (2020). A pneumonia outbreak associated with a new coronavirus of probable bat origin. *Nature* 579, 270–273. doi: 10.1038/s41586-020-2012-7

Zhu, A., Wei, P., Man, M., Liu, X., Ji, T., Chen, J., et al. (2023). Antigenic characterization of SARS-CoV-2 omicron subvariants XBB.1.5, BQ.1, BQ.1.1, BF.7 and BA.2.75.2. Signal Transduct. Target. *Therapy* 8:125. doi: 10.1038/s41392-023-01391-x

Zhu, N., Zhang, D., Wang, W., Li, X., Yang, B., Song, J., et al. (2020). A novel coronavirus from patients with pneumonia in China, 2019. *New Engl. J. Med.* 382, 727–733. doi: 10.1056/NEJMoa2001017
Samarasekera, U. (2021). India grapples with second wave of COVID-19. *Lancet Microbe* 2:e238. doi: 10.1016/S2666-5247(21)00123-3

Shang, J., Ye, G., Shi, K., Wan, Y., Luo, C., Aihara, H., et al. (2020). Structural basis of receptor recognition by SARS-CoV-2. *Nature* 581, 221–224. doi: 10.1038/s41586-020-2179-y

Shen, X., Tang, H., Pajon, R., Smith, G., Glenn, G. M., Shi, W., et al. (2021). Neutralization of SARS-CoV-2 variants B1.429 and B1.351. *New Engl. J. Med.* 384:24. doi: 10.1056/NEJMc2103740

Sheward, D. J., Kim, C., Ehling, R. A., Pankow, A., Castro Dopico, X., Dyrdak, R., et al. (2022). Neutralisation sensitivity of the SARS-CoV-2 omicrom (.B1.1.529) variant: a cross-sectional study. *Lancet Infect. Dis.* 22, 813–820. doi: 10.1016/S1473-3099(22)00129-3

Shi, J., Wen, Z., Zhong, G., Yang, H., Wang, C., Huang, B., et al. (2020). Susceptibility of ferrets, cats, dogs, and other domesticated animals to SARS-coronavirus-2. *Science* 368, 1016–1020. doi: 10.1126/science.abb7015

Shukla, N., Roelle, S. M., Suzart, V. G., Bruchez, A. M., and Matreyek, K. A. (2021). Mutants of human ACE2 differentially promote SARS-CoV and SARS-CoV-2 spike mediated infection. *PLoS Pathog.* 17:e1009715. doi: 10.1371/journal.ppat.1009715

Simas, P. V. M., de Souza Barnabé, A. C., Durães-Carvalho, R., de Lima, F., Neto, D., Caserta, L., et al. (2015). Bat coronavirus in Brazil related to appalachian ridge and porcine epidemic diarrhea viruses. *Emerg. Infect. Dis.* 21, 729–731. doi: 10.3201/eid2104.141783

Snouwaert, J. N., Jania, L. A., Nguyen, T., Martinez, D. R., Schäfer, A., Catanzaro, N. J., et al. (2023). Human ACE2 expression, a major tropism determinant for SARS-CoV-2, is regulated by upstream and intragenic elements. *PLoS Pathog.* 19:e1011168. doi: 10.1371/journal.ppat.1011168

Song, H. D., Tu, C. C., Zhang, G. W., Wang, S. Y., Zheng, K., Lei, L. C., et al. (2005). (2005) cross-host evolution of severe acute respiratory syndrome coronavirus in palm civet and human. *Proc. Natl. Acad. Sci. U. S. A.* 102, 2430–2435. doi: 10.1073/pnas.0409608102

Subbarao, K., McAuliffe, J., Vogel, L., Fahle, G., Fischer, S., Tatti, K., et al. (2004). Prior infection and passive transfer of neutralizing antibody prevent replication of severe acute respiratory syndrome coronavirus in the respiratory tract of mice. *J. Virol.* 78, 3572–3537. doi: 10.1128/JVI.78.7.3572-3577.2004

Sun, F., Wang, X., Tan, S., Dan, Y., Lu, Y., Zhang, J., et al. SARS-CoV-2 quasispecies provides an advantage mutation pool for the epidemic variants. *Microbiol Spectr.* (2021) 9:e00261-e00221. doi: 10.1128/Spectrum.00261-21

Suryamohan, K., Diwanji, D., Stawiski, E. W., Gupta, R., Miersch, S., Liu, J., et al. (2021). Human ACE2 receptor polymorphisms and altered susceptibility to SARS-CoV-2. *Comm. Biol.* 4:475. doi: 10.1038/s42003-021-02030-3

Takada, K., Takahashi Ueda, M., Shichinohe, S., Kida, Y., Ono, C., Matsuura, Y., et al. (2023). Genomic diversity of SARS-CoV-2 can be accelerated by a mutation in the nsp14 gene. *iScience* 26:106210. doi: 10.1016/j.isci.2023.106210

Tang, J. W., Tambyah, P. A., and Hui, D. S. (2020). Emergence of a new SARS-CoV-2 variant in the UK. *J. Infect.* 82, e27–e28. doi: 10.1016/j.jinf.2020.12.024

Tegally, H., Wilkinson, E., Giovanetti, M., Iranzadeh, A., Fonseca, V., Giandhari, J., et al. (2021). Detection of a SARS-CoV-2 variant of concern in South Africa. *Nature* 592, 438–443. doi: 10.1038/s41586-021-03402-9

Thye, A. Y. K., Law, J. W. F., Pusparajah, P., Letchumanan, V., Chan, K. G., and Lee, L. H. (2021). Emerging SARS-CoV-2 variants of concern (VOCs): An impending global crisis. *Biomedicine* 9:1303. doi: 10.3390/biomedicines9101303

Towler, P., Staker, B., Prasad, S. G., Menon, S., Tang, J., Parsons, T., et al. (2004). ACE2 X-ray structures reveal a large hinge-bending motion important for inhibitor binding and catalysis. *J. Biol. Chem.* 279, 17996–18007. doi: 10.1074/jbc.M311191200

Turner, A. J., and Hooper, N. M. (2002). The angiotensin-converting enzyme gene family: genomics and pharmacology. *Trends Pharmacol. Sci.* 23, 177–183. doi: 10.1016/
s0165-6147(00)01994-5

Wacharalpluesadee, S., Tan, C. W., Maneeorn, P., Duengkae, P., Zhu, F., Joyjinda, Y., et al. (2021). Evidence for SARS-CoV-2 related coronaviruses circulating in bats and pangolins in Southeast Asia. *Nature Com* 12:972. doi: 10.1038/s41467-021-21240-1

Wang, Q., Iketani, S., Li, Z., Liu, L., Guo, Y., Huang, Y., et al. (2023). Alarming antibody evasion properties of rising SARS-CoV-2 BQ and XBB subvariants. *Cells* 186, 1–8. doi: 10.1016/j.cell.2022.12.018

Wei, C., Shan, K. J., Wang, W., Zhang, S., Huan, Q., and Qian, W. (2021). Evidence for a mouse origin of the SARS-CoV-2 omicron variant. *J. Genet. Genom.* 48, 1111–1121. doi: 10.1016/j.jgg.2021.12.003

Weissman, D., Alameh, M. G., de Silva, T., Collini, P., Hornsby, H., Brown, R., et al. (2021). D614G spike mutation increases SARS-CoV-2 susceptibility to neutralization. *Cell Host Microbe* 29, 23–31.e4. doi: 10.1016/j.chom.2020.11.012

Wentworth, D. E., Gillim-Ross, L., Espina, N., and Bernard, K. A. (2004). Mice susceptible to SARS coronavirus. *Emerg. Infect. Dis.* 10, 1293–1296. doi: 10.3201/eid1007.031119

Wertheim, J. O., Chu, D. K., Peiris, J. S., Kosakovsky Pond, S. L., and Poon, L. L. (2013). A case for the ancient origin of coronaviruses. *J. Virol* 87, 7039–7045. doi: 10.1128/JVI.03273-12

Woo, P. C., Lau, S. K., Lam, C. S., Lau, C. C., Tsang, A. K., Lau, J. H., et al. (2012). Discovery of seven novel mammalian and avian coronaviruses in the genus deltacoronavirus supports bat coronaviruses as the gene source of alphacoronavirus and betacoronavirus and avian coronaviruses as the gene source of gamma-coronavirus and delta-coronavirus. *J. Virol* 86, 3995–4008. doi: 10.1128/JVI.06540-11

Xu, D., Zhang, Z., and Wang, F. S. (2004). SARS-associated coronavirus quasispecies in individual patients. *N. Engl. J. Med.* 350, 1366–1367. doi: 10.1056/NEJMc032421

Yan, R., Zhang, Y., Li, Y., Xia, L., Guo, Y., Zhou, Q., et al. (2020). Structural basis for the recognition of the SARS-CoV-2 by full-length human ACE2. *Science* 367, 1444–1448. doi: 10.1126/science.abb2762

Yang, M., Zhao, J., Xing, L., and Shi, L. (2015). The association between angiotensinconverting enzyme 2 polymorphisms and essential hypertension risk: a meta-analysis involving 14,122 patients. *J. Renin-Angiotensin-Aldosterone Syst.* 16, 1240–1244. doi: 10.1177/1470320314549221

Yin, X., Popa, H., Stapon, A., Bouda, E., and Garcia-Diaz, M. (2023). Fidelity of ribocleotide incorporation by the SARS-CoV-2 replication complex. *J. Mol. Biol.* 435:167973. doi: 10.1016/j.jmb.2023.167973

Yue, C., Li, Z., Lin, Y., Yang, Y., Yuan, M., Pan, Z., et al. (2021). Sensitivity of SARS-CoV-2 variants to neutralization by convalescent sera and a VH3-30 monoclonal antibody. *Front. Immunol.* 12:751584. doi: 10.3389/fimmu.2021.751584

Yue, C., Song, W., Wang, L., Jian, F., Chen, X., Gao, F., et al. (2023). ACE2 binding and antibody evasion in enhanced transmissibility of XBB.1.5. *Lancet Inf. Dis.* 23, 278–280. doi: 10.1016/S1473-3099(23)00010-5

Zhou, H., Chen, X., Hu, T., Li, J., Song, H., Liu, Y., et al. (2020). A novel bat coronavirus closely related to SARS-CoV-2 contains natural insertions at the S1/S2 cleavage site of the spike protein. *Curr. Biol.* 30, 2196–2203. doi: 10.1016/j.cub.2020.05.023

Zhou, P., Fan, H., Lan, T., Yang, X. L., Shi, W. F., Zhang, W., et al. (2018). Fatal swine acute diarrhoea syndrome caused by an HKU2-related coronavirus of bat origin. *Nature* 556, 255–258. doi: 10.1038/s41586-018-0010-9

Zhou, P., Yang, X. L., Wang, X. G., Hu, B., Zhang, L., Zhang, W., et al. (2020). A pneumonia outbreak associated with a new coronavirus of probable bat origin. *Nature* 579, 270–273. doi: 10.1038/s41586-020-2012-7

Zhu, A., Wei, P., Man, M., Liu, X., Ji, T., Chen, J., et al. (2023). Antigenic characterization of SARS-CoV-2 omicron subvariants XBB.1.5, BQ.1, BQ.1.1, BF.7 and BA.2.75.2. Signal Transduct. Target. *Therapy* 8:125. doi: 10.1038/s41392-023-01391-x

Zhu, N., Zhang, D., Wang, W., Li, X., Yang, B., Song, J., et al. (2020). A novel coronavirus from patients with pneumonia in China, 2019. *New Engl. J. Med.* 382, 727–733. doi: 10.1056/NEJMoa2001017