Generation of G9 genotype porcine rotavirus using reverse genetics system and its application for antiviral screen and vaccine development

Nan Han; Ran Tao; Xi Cheng; Hui Deng; Jianxin Wang; Xianyu Bian; Jinzhu Zhou; Xuejiao Zhu; Min Sun; Xuehan Zhang; Mi Hu; Rongli Guo; Wei Wang; Siyuan Ding; Bin Li

doi:10.1016/j.virs.2026.04.005

April 2026

Citation: Nan Han, Ran Tao, Xi Cheng, Hui Deng, Jianxin Wang, Xianyu Bian, Jinzhu Zhou, Xuejiao Zhu, Min Sun, Xuehan Zhang, Mi Hu, Rongli Guo, Wei Wang, Siyuan Ding, Bin Li. Generation of G9 genotype porcine rotavirus using reverse genetics system and its application for antiviral screen and vaccine development .VIROLOGICA SINICA, 2026, 41(2) : 439-454. http://dx.doi.org/10.1016/j.virs.2026.04.005

Generation of G9 genotype porcine rotavirus using reverse genetics system and its application for antiviral screen and vaccine development

Nan Han ^a,b,c,d ,
Ran Tao ^a,b,c ,
Xi Cheng ^a,b,c,e ,
Hui Deng ^a,b,c ,
Jianxin Wang ^a,b,c ,
Xianyu Bian ^a,b,c ,
Jinzhu Zhou ^a,b,c ,
Xuejiao Zhu ^a,b,c ,
Min Sun ^a,b,c ,
Xuehan Zhang ^a,b,c ,
Mi Hu ^a,b,c ,
Rongli Guo ^a,b,c ,
Wei Wang ^a,b,c ,
Siyuan Ding ^{f
,,} ,
Bin Li ^{a,b,c,d,e
,,}

a. Institute of Veterinary Medicine, Jiangsu Academy of Agricultural Sciences, Key Laboratory of Veterinary Biological Engineering and Technology, Ministry of Agriculture, Jiangsu Key Laboratory for Food Quality and Safety-State Key Laboratory Cultivation Base of Ministry of Science and Technology, Nanjing 210014, China;
b. Jiangsu Co-Innovation Center for the Prevention and Control of Important Animal Infectious Diseases and Zoonoses, Jiangsu Key Laboratory of Zoonoses, Yangzhou University, Yangzhou 225009, China;
c. GuoTai (Taizhou) Center of Technology Innovation for Veterinary Biologicals, Taizhou 225300, China;
d. College of Veterinary Medicine, Yangzhou University, Yangzhou 225009, China;
e. College of Veterinary Medicine, Nanjing Agricultural University, Nanjing 210095, China;
f. Department of Molecular Microbiology, Washington University School of Medicine in St. Louis, St. Louis 63110, Missouri, USA

Corresponding author: Siyuan Ding, siyuan.ding@wustl.edu
Bin Li, libinana@126.com
Received Date: 19 October 2025
Accepted Date: 10 April 2026
Available online: 15 April 2026

Abstract

Group A rotaviruses (RVs) continue to be one of the most important pathogens causing severe acute gastroenteritis in infants and young animals worldwide. Recently, the prevalence of porcine RV (PoRV) from pig farms has strikingly increased, adversely affecting the swine industry, particularly with the G9 genotype of PoRV VP7 emerging as the predominant genotype spreading in China. Current vaccines against PoRV fail to provide sufficient protective immunity, necessitating urgent development of effective vaccines and antiviral drugs against PoRV. Here, we successfully established and improved the entirely plasmid-based reverse genetics (RG) system to rescue a G9 genotype of recombinant PoRV AHFY2022 strain (G9P[23]). Using the improved RG system, we rescued recombinant AHFY2022 harboring the fluorescent UnaG protein or nano-luciferase (NLuc) reporter within gene segment 7 that encodes non-structural protein 3 (NSP3). Furthermore, we adopted the UnaG reporter virus to screen anti-PoRV drugs and identified two promising antiviral drugs, C8 and C9. Moreover, we generated the recombinant PoRV (rAHFY2022-G5-VP7) containing G5 genotype of VP7 from PoRV-positive samples in the backbone of AHFY2022 strain. The reassortant strain exhibited efficient replication and genetic stability. Mouse models were utilized to evaluate the immune responses elicited by rAHFY2022-G5-VP7 strain in vivo, revealing similar neutralizing antibodies and cellular immune response compared to the parental AHFY2022 strain in mice. Together, this study provides an important tool for screening potential anti-PoRV drugs and developing novel vaccines against prevalent PoRV strains.
- Reverse genetics
- , Reporter virus
- , Porcine rotavirus (PoRV)
- , Antivirals
- , Vaccine development

Electronic Supplementary Material

10.1016j.virs.2026.04.005-ESM.docx
References
1. Blutt, S.E., Crawford, S.E., Warfield, K.L., Lewis, D.E., Estes, M.K., Conner, M.E., 2004. The VP7 outer capsid protein of rotavirus induces polyclonal B-cell activation. J Virol 78, 6974-6981.
2. de Sautu, M., Herrmann, T., Scanavachi, G., Jenni, S., Harrison, S.C., 2024. The rotavirus VP5∗/VP8∗ conformational transition permeabilizes membranes to Ca2. PLoS Pathog 20, e1011750.
3. Desselberger, U., 2020. Potential of plasmid only based reverse genetics of rotavirus for the development of next-generation vaccines. Curr Opin Virol 44, 1-6.
4. Diebold, O., Gonzalez, V., Venditti, L., Sharp, C., Blake, R.A., Tan, W.S., Stevens, J., Caddy, S., Digard, P., Borodavka, A., Gaunt, E., 2022. Using Species a Rotavirus Reverse Genetics to Engineer Chimeric Viruses Expressing SARS-CoV-2 Spike Epitopes. J Virol 96, e0048822.
5. Fukuda, S., Kugita, M., Kumamoto, K., Akari, Y., Higashimoto, Y., Nagao, S., Murata, T., Yoshikawa, T., Taniguchi, K., Komoto, S., 2024. Generation of Recombinant Authentic Live Attenuated Human Rotavirus Vaccine Strain RIX4414 (Rotarix((R))) from Cloned cDNAs Using Reverse Genetics. Viruses 16(8): 1198.
6. Ghonaim, A.H., Rouby, S.R., Nageeb, W.M., Elgendy, A.A., Xu, R., Jiang, C., Ghonaim, N.H., He, Q., Li, W., 2025. Insights into recent advancements in human and animal rotavirus vaccines: Exploring new frontiers. Virol Sin 40, 1-14.
7. Graham, K.L., Halasz, P., Tan, Y., Hewish, M.J., Takada, Y., Mackow, E.R., Robinson, M.K., Coulson, B.S., 2003. Integrin-using rotaviruses bind alpha2beta1 integrin alpha2 I domain via VP4 DGE sequence and recognize alphaXbeta2 and alphaVbeta3 by using VP7 during cell entry. J Virol 77, 9969-9978.
8. Hamajima, R., Lusiany, T., Minami, S., Nouda, R., Nurdin, J.A., Yamasaki, M., Kobayashi, N., Kanai, Y., Kobayashi, T., 2022. A reverse genetics system for human rotavirus G2P[4]. J Gen Virol 103, 12.
9. Han, X., Si, L.L., Shi, Y.Y., Fan, Z.B., Wang, S.X., Tian, Z.Y., Li, M., Sun, J.Q., Jiao, P.X., Ran, F.X., Zhang, Y.M., Zhou, D.M., Xiao, S.L., 2017. Synthesis and In Vitro Anti-Influenza Virus Evaluation of Novel Sialic Acid (C-5 and C-9)-Pentacyclic Triterpene Derivatives. Molecules 22.
10. Hassan Mir, R., Godavari, G., Siddiqui, N.A., Ahmad, B., Mothana, R.A., Ullah, R., Almarfadi, O.M., Jachak, S.M., Masoodi, M.H., 2021. Design, Synthesis, Molecular Modelling, and Biological Evaluation of Oleanolic Acid-Arylidene Derivatives as Potential Anti-Inflammatory Agents. Drug Des Devel Ther 15, 385-397.
11. Hatazawa, R., Fukuda, S., Kumamoto, K., Matsushita, F., Nagao, S., Murata, T., Taniguchi, K., Matsui, T., Komoto, S., 2021. Strategy for generation of replication-competent recombinant rotaviruses expressing multiple foreign genes. J Gen Virol 102(4).
12. He, S., Shi, J., Hu, H., Zhang, X., Ning, M., Wu, W., Wen, Y., Wang, Y., Wu, R., Zhao, Q., Du, S., Cao, S., Huang, X., Zhao, S., Lang, Y., Zeng, N., Yan, Q., 2025. Molecular Epidemiological Investigation and Viral Isolation of Porcine Rotavirus in Southwest China During 2024-2025. Vet Sci 12(12):1137.
13. Hou, G., Zeng, Q., Matthijnssens, J., Greenberg, H.B., Ding, S., 2021. Rotavirus NSP1 Contributes to Intestinal Viral Replication, Pathogenesis, and Transmission. mBio 12, e0320821.
14. Hou, W., Fan, M., Zhu, Z., Li, X., 2023. Establishment and Application of a Triplex Real-Time RT-PCR Assay for Differentiation of PEDV, PoRV, and PDCoV. Viruses 15(6), 1238.
15. Jannus, F., Sainz, J., Reyes-Zurita, F.J., 2024. Principal Bioactive Properties of Oleanolic Acid, Its Derivatives, and Analogues. Molecules 29(14), 3291.
16. Jung, J., Bae, J., Park, J.S., Lee, S.W., Jeong, J.H., Park, S.J., 2024. In Vitro Anti-Rotaviral Activity of Bavachin Isolated from Psoralea corylifolia L. (Fabaceae). Vet Sci 11(5):188.
17. Kanai, Y., Kawagishi, T., Nouda, R., Onishi, M., Pannacha, P., Nurdin, J.A., Nomura, K., Matsuura, Y., Kobayashi, T., 2019. Development of Stable Rotavirus Reporter Expression Systems. J Virol 93(4), e01774-18.
18. Kanai, Y., Kobayashi, T., 2021. Rotavirus reverse genetics systems: Development and application. Virus Res 295, 198296.
19. Kanai, Y., Komoto, S., Kawagishi, T., Nouda, R., Nagasawa, N., Onishi, M., Matsuura, Y., Taniguchi, K., Kobayashi, T., 2017. Entirely plasmid-based reverse genetics system for rotaviruses. Proc Natl Acad Sci U S A 114, 2349-2354.
20. Kanai, Y., Onishi, M., Kawagishi, T., Pannacha, P., Nurdin, J.A., Nouda, R., Yamasaki, M., Lusiany, T., Khamrin, P., Okitsu, S., Hayakawa, S., Ebina, H., Ushijima, H., Kobayashi, T., 2020. Reverse Genetics Approach for Developing Rotavirus Vaccine Candidates Carrying VP4 and VP7 Genes Cloned from Clinical Isolates of Human Rotavirus. J Virol 95(2):e01374-20.
21. Kanda, M., Fukuda, S., Hamada, N., Nishiyama, S., Masatani, T., Fujii, Y., Izumi, F., Okajima, M., Taniguchi, K., Sugiyama, M., Komoto, S., Ito, N., 2022. Establishment of a reverse genetics system for avian rotavirus A strain PO-13. J Gen Virol 103(6).
22. Kawagishi, T., Nurdin, J.A., Onishi, M., Nouda, R., Kanai, Y., Tajima, T., Ushijima, H., Kobayashi, T., 2020. Reverse Genetics System for a Human Group A Rotavirus. J Virol 94(2):e00963-19.
23. Kawagishi, T., Sanchez-Tacuba, L., Feng, N., Costantini, V.P., Tan, M., Jiang, X., Green, K.Y., Vinje, J., Ding, S., Greenberg, H.B., 2023. Mucosal and systemic neutralizing antibodies to norovirus induced in infant mice orally inoculated with recombinant rotaviruses. Proc Natl Acad Sci U S A 120, e2214421120.
24. Khwaza, V., Oyedeji, O.O., Aderibigbe, B.A., 2018. Antiviral Activities of Oleanolic Acid and Its Analogues. Molecules 23(9), 2300.
25. Kinra, M., Joseph, A., Nampoothiri, M., Arora, D., Mudgal, J., 2021. Inhibition of NLRP3-inflammasome mediated IL-1beta release by phenylpropanoic acid derivatives: in-silico and in-vitro approach. Eur J Pharm Sci 157, 105637.
26. Komoto, S., Fukuda, S., Ide, T., Ito, N., Sugiyama, M., Yoshikawa, T., Murata, T., Taniguchi, K., 2018. Generation of Recombinant Rotaviruses Expressing Fluorescent Proteins by Using an Optimized Reverse Genetics System. J Virol 92(13), e00588-18.
27. Komoto, S., Fukuda, S., Kugita, M., Hatazawa, R., Koyama, C., Katayama, K., Murata, T., Taniguchi, K., 2019. Generation of Infectious Recombinant Human Rotaviruses from Just 11 Cloned cDNAs Encoding the Rotavirus Genome. J Virol 93(8):e02207-e02218.
28. Li, C., Lu, H., Geng, C., Yang, K., Liu, W., Liu, Z., Yuan, F., Gao, T., Wang, S., Wen, P., Song, H., Tian, Y., Zhou, D., 2022. Epidemic and Evolutionary Characteristics of Swine Enteric Viruses in South-Central China from 2018 to 2021. Viruses 14(7):1420.
29. Li, J., Xiao, L., Chen, Z., Fan, L., Wang, W., Guo, R., He, Z., Hu, H., Jiang, J., Zhao, L., Zhong, T., Fan, B., Zhu, X., Li, B., 2024. A spike-based mRNA vaccine that induces durable and broad protection against porcine deltacoronavirus in piglets. J Virol 98, e0053524.
30. Liu, H., Tian, H., Hao, P., Du, H., Wang, K., Qiu, Y., Yin, X., Wu, N., Du, Q., Tong, D., Huang, Y., 2024a. PoRVA G9P[23] and G5P[7] infections differentially promote PEDV replication by reprogramming glutamine metabolism. PLoS Pathog 20, e1012305.
31. Liu, X., Li, S., Yu, J., Chai, P., Xie, Z., Pang, L., Li, J., Zhu, W., Ren, W., Duan, Z., 2024b. Establishment of a Reverse Genetics System for Rotavirus Vaccine Strain LLR and Developing Vaccine Candidates Carrying VP7 Gene Cloned From Human Strains Circulating in China. J Med Virol 96, e70065.
32. Liu, Z., Gu, S., Zhu, X., Liu, M., Cao, Z., Qiu, P., Li, S., Liu, S., Song, G., 2022. Discovery and optimization of new 6, 7-dihydroxy-1, 2, 3, 4-tetrahydroisoquinoline derivatives as potent influenza virus PA(N) inhibitors. Eur J Med Chem 227, 113929.
33. Lopez, S., Arias, C.F., 2004. Multistep entry of rotavirus into cells: a Versaillesque dance. Trends Microbiol 12, 271-278.
34. Lv, Y., Tong, Z., Liu, J., Zhang, Z., Wang, C., Zeng, Y., Liu, P., Zong, X., Chen, G., Chen, H., Tan, C., 2024. Molecular Characterization and Pathogenicity Analysis of Porcine Rotavirus A. Viruses 16(12), 1842.
35. Makabe, T., Komaniwa, H., Kishi, Y., Yataya, K., Imagawa, H., Sato, K., Inaba, Y., 1985. Isolation of ovine rotavirus in cell cultures. Brief report. Arch Virol 83, 123-127.
36. Matthijnssens, J., Ciarlet, M., Rahman, M., Attoui, H., Banyai, K., Estes, M.K., Gentsch, J.R., Iturriza-Gomara, M., Kirkwood, C.D., Martella, V., Mertens, P.P., Nakagomi, O., Patton, J.T., Ruggeri, F.M., Saif, L.J., Santos, N., Steyer, A., Taniguchi, K., Desselberger, U., Van Ranst, M., 2008. Recommendations for the classification of group A rotaviruses using all 11 genomic RNA segments. Arch Virol 153, 1621-1629.
37. Matthijnssens, J., Otto, P.H., Ciarlet, M., Desselberger, U., Van Ranst, M., Johne, R., 2012. VP6-sequence-based cutoff values as a criterion for rotavirus species demarcation. Arch Virol 157, 1177-1182.
38. Meng, L., Su, Y., Yang, F., Xiao, S., Yin, Z., Liu, J., Zhong, J., Zhou, D., Yu, F., 2019. Design, synthesis and biological evaluation of amino acids-oleanolic acid conjugates as influenza virus inhibitors. Bioorg Med Chem 27, 115147.
39. Pan, Y., Li, Z., Miao, Q., Shi, H., Guo, L., Feng, L., Tian, J., 2025. Phylogenitc analysis and immunogenicity comparison of porcine genotype G9 rotavirus in China from 2020-2023. Virol Sin 40, 176-185.
40. Park, J.G., Alfajaro, M.M., Cho, E.H., Kim, J.Y., Soliman, M., Baek, Y.B., Park, C.H., Lee, J.H., Son, K.Y., Cho, K.O., Kang, M.I., 2019. Development of a live attenuated trivalent porcine rotavirus A vaccine against disease caused by recent strains most prevalent in South Korea. Vet Res 50, 2.
41. Patton, J.T., Silvestri, L.S., Tortorici, M.A., Vasquez-Del Carpio, R., Taraporewala, Z.F., 2006. Rotavirus genome replication and morphogenesis: role of the viroplasm. Curr Top Microbiol Immunol 309, 169-187.
42. Philip, A.A., Hu, S., Dai, J., Patton, J.T., 2023. Recombinant rotavirus expressing the glycosylated S1 protein of SARS-CoV-2. J Gen Virol 104.
43. Philip, A.A., Patton, J.T., 2020. Expression of Separate Heterologous Proteins from the Rotavirus NSP3 Genome Segment Using a Translational 2A Stop-Restart Element. J Virol 94(18):e00959-20.
44. Philip, A.A., Perry, J.L., Eaton, H.E., Shmulevitz, M., Hyser, J.M., Patton, J.T., 2019. Generation of Recombinant Rotavirus Expressing NSP3-UnaG Fusion Protein by a Simplified Reverse Genetics System. J Virol 93(24):e01616-e01619.
45. Qiao, M., Li, M., Li, Y., Wang, Z., Hu, Z., Qing, J., Huang, J., Jiang, J., Jiang, Y., Zhang, J., Gao, C., Yang, C., Li, X., Zhou, B., 2024. Recent Molecular Characterization of Porcine Rotaviruses Detected in China and Their Phylogenetic Relationships with Human Rotaviruses. Viruses 16(3):453.
46. Qin, S.K., Li, K.H., Liu, B.J., Cao, C., Yu, D.B., Jiang, Z.G., Wang, J., Han, Y.X., Wang, F., Qi, Y.L., Sun, C., Yu, L., Chang, J.T., Yin, X., 2024. Efficient and robust reverse genetics system for bovine rotavirus generation and its application for antiviral screening. Virol Sin 39, 917-928.
47. Sanchez-Tacuba, L., Feng, N., Meade, N.J., Mellits, K.H., Jais, P.H., Yasukawa, L.L., Resch, T.K., Jiang, B., Lopez, S., Ding, S., Greenberg, H.B., 2020. An Optimized Reverse Genetics System Suitable for Efficient Recovery of Simian, Human, and Murine-Like Rotaviruses. J Virol 94(18):e01294-20.
48. Sanekata, T., Kuwamoto, Y., Akamatsu, S., Sakon, N., Oseto, M., Taniguchi, K., Nakata, S., Estes, M.K., 1996. Isolation of group B porcine rotavirus in cell culture. J Clin Microbiol 34, 759-761.
49. Sato, K., Tokuhisa, S., Inaba, Y., 1995. Effect of enzymes on the growth of human and animal rotaviruses. J Vet Med Sci 57, 569-570.
50. Snyder, A.J., Agbemabiese, C.A., Patton, J.T., 2024. Production of OSU G5P[7] Porcine Rotavirus Expressing a Fluorescent Reporter via Reverse Genetics. Viruses 16(3):411.
51. Son, J., Huang, S., Zeng, Q., Bricker, T.L., Case, J.B., Zhou, J., Zang, R., Liu, Z., Chang, X., Darling, T.L., Xu, J., Harastani, H.H., Chen, L., Gomez Castro, M.F., Zhao, Y., Kohio, H.P., Hou, G., Fan, B., Niu, B., Guo, R., Rothlauf, P.W., Bailey, A.L., Wang, X., Shi, P.Y., Martinez, E.D., Brody, S.L., Whelan, S.P.J., Diamond, M.S., Boon, A.C.M., Li, B., Ding, S., 2022. JIB-04 Has Broad-Spectrum Antiviral Activity and Inhibits SARS-CoV-2 Replication and Coronavirus Pathogenesis. mBio 13, e0337721.
52. Song, W., Feng, Y., Zhang, J., Kong, D., Fan, J., Zhao, M., Hua, L., Xiang, J., Tang, X., Xiao, S., Peng, Z., Wu, B., 2024. Development of a multiplex reverse transcription-quantitative PCR (qPCR) method for detecting common causative agents of swine viral diarrhea in China. Porcine Health Manag 10, 12.
53. Tang, X., Li, S., Zhou, J., Bian, X., Wang, J., Han, N., Zhu, X., Tao, R., Wang, W., Sun, M., Li, P., Zhang, X., Li, B., 2024. Recombinant bivalent subunit vaccine combining truncated VP4 from P[7] and P[23] induces protective immunity against prevalent porcine rotaviruses. J Virol 98, e0021224.
54. Tao, R., Chang, X., Zhou, J., Zhu, X., Yang, S., Li, K., Gu, L., Zhang, X., Li, B., 2023. Molecular epidemiological investigation of group A porcine rotavirus in East China. Front Vet Sci 10, 1138419.
55. Tao, R., Cheng, X., Gu, L., Zhou, J., Zhu, X., Zhang, X., Guo, R., Wang, W., Li, B., 2024. Lipidomics reveals the significance and mechanism of the cellular ceramide metabolism for rotavirus replication. J Virol 98, e0006424.
56. Troeger, C., Khalil, I.A., Rao, P.C., Cao, S., Blacker, B.F., Ahmed, T., Armah, G., Bines, J.E., Brewer, T.G., Colombara, D.V., Kang, G., Kirkpatrick, B.D., Kirkwood, C.D., Mwenda, J.M., Parashar, U.D., Petri, W.A., Jr., Riddle, M.S., Steele, A.D., Thompson, R.L., Walson, J.L., Sanders, J.W., Mokdad, A.H., Murray, C.J.L., Hay, S.I., Reiner, R.C., Jr., 2018. Rotavirus Vaccination and the Global Burden of Rotavirus Diarrhea Among Children Younger Than 5 Years. JAMA Pediatr 172, 958-965.
57. Uprety, T., Wang, D., Li, F., 2021. Recent advances in rotavirus reverse genetics and its utilization in basic research and vaccine development. Arch Virol 166, 2369-2386.
58. Valusenko-Mehrkens, R., Gadicherla, A.K., Johne, R., Falkenhagen, A., 2023. Strain-Specific Interactions between the Viral Capsid Proteins VP4, VP7 and VP6 Influence Rescue of Rotavirus Reassortants by Reverse Genetics. Int J Mol Sci 24(6):5670.
59. Vlasova, A.N., Amimo, J.O., Saif, L.J., 2017. Porcine Rotaviruses: Epidemiology, Immune Responses and Control Strategies. Viruses 9(3), 48.
60. Wang, J., Zhou, J., Zhu, X., Bian, X., Han, N., Fan, B., Gu, L., Cheng, X., Li, S., Tao, R., Li, J., Zhang, X., Li, B., 2024. Isolation and characterization of a G9P[23] porcine rotavirus strain AHFY2022 in China. Microb Pathog 190, 106612.
61. Woode, G.N., Bridger, J., Hall, G.A., Jones, J.M., Jackson, G., 1976. The isolation of reovirus-like agents (rota-viruses) from acute gastroenteritis of piglets. J Med Microbiol 9, 203-209.
62. Xue, R., Tian, Y., Zhang, Y., Zhang, M., Li, Z., Chen, S., Liu, Q., 2018. Diversity of group A rotavirus of porcine rotavirus in Shandong province China. Acta Virol 62, 229-234.
63. Zhang, F., Luo, S., Gu, J., Li, Z., Li, K., Yuan, W., Ye, Y., Li, H., Ding, Z., Song, D., Tang, Y., 2019. Prevalence and phylogenetic analysis of porcine diarrhea associated viruses in southern China from 2012 to 2018. BMC Vet Res 15, 470.
64. Zhang, F., Luo, Y., Lin, C., Tan, M., Wan, P., Xie, B., Xiong, L., Ji, H., 2024. Epidemiological monitoring and genetic variation analysis of pathogens associated with porcine viral diarrhea in southern China from 2021 to 2023. Front Microbiol 15, 1303915.
65. Zhao, Y., Fan, B., Song, X., Gao, J., Guo, R., Yi, C., He, Z., Hu, H., Jiang, J., Zhao, L., Zhong, T., Li, B., 2024. PEDV-spike-protein-expressing mRNA vaccine protects piglets against PEDV challenge. mBio 15, e0295823.
66. Zhou, J., Huang, S., Fan, B., Niu, B., Guo, R., Gu, J., Gao, S., Li, B., 2021. iTRAQ-based proteome analysis of porcine group A rotavirus-infected porcine IPEC-J2 intestinal epithelial cells. J Proteomics 248, 104354.
67. Zhou, X., Li, Y., Li, T., Cao, J., Guan, Z., Xu, T., Jia, G., Ma, G., Zhao, R., 2023. Portulaca oleracea L. Polysaccharide Inhibits Porcine Rotavirus In Vitro. Animals (Basel) 13(14):2306.
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