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Therefore, developing safe, effective and affordable vaccines remains essential to meet the global demand, particularly in developing countries. Transgenic plants have emerged as a promising platform to express recombinant proteins for pharmaceutical and vaccine applications. Two binary vectors, pCAMBIA1300Gt1\u2010S1 and pCAMBIA1300Actin\u2010S1, containing distinct promoters, were constructed and transformed into rice via\n                    <jats:italic>Agrobacterium<\/jats:italic>\n                    . Overall, 56 independent transgenic rice lines were regenerated. Expression analysis revealed that the rice\u2010derived S1 (rS1) protein could be expressed in pGt1::S1 transgenic rice seeds. rS1 protein expression levels reached up to 282\u2009\u03bcg\/g dry weight, with S1 gene insertion having no effect on grain size and weight. The rS1 protein exhibited a high affinity for human angiotensin\u2010converting enzyme 2 (ACE2)\n                    <jats:italic>in vitro<\/jats:italic>\n                    . Moreover, the immunogenicity of purified rS1 protein co\u2010administered with various adjuvants demonstrated that mice vaccinated with Alum\u2010adjuvant rS1 generated enhanced humoral immune responses with high serum IgG, IgG1 and neutralizing antibody levels.\n                    <jats:italic>Salmonella<\/jats:italic>\n                    Typhimurium flagellin (FliC)\u2010adjuvanted rS1 elicited stronger S1\u2010specific IgG2a levels, promoted splenocyte proliferation and induced mixed Th1\/Th2\/Th17 cytokine responses. This was evidenced by increased proportions of antigen\u2010specific interferon (IFN)\u2010\u03b3, interleukin\u20104 (IL\u20104) and IL\u201017A\u2010positive CD4\n                    <jats:sup>+<\/jats:sup>\n                    T lymphocytes, suggesting its potential to induce both humoral and cellular immune responses. These findings suggest that rS1 protein offers a promising approach for affordable COVID\u201019 subunit vaccine production, and this strategy can be universally applied to other viral vaccines.\n                  <\/jats:p>","DOI":"10.1111\/pbi.70077","type":"journal-article","created":{"date-parts":[[2025,4,5]],"date-time":"2025-04-05T09:25:10Z","timestamp":1743845110000},"page":"2570-2582","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":4,"title":["Rice\u2010derived\n                    <scp>SARS<\/scp>\n                    \u2010\n                    <scp>CoV<\/scp>\n                    \u20102 glycoprotein\n                    <scp>S1<\/scp>\n                    subunit vaccine elicits humoral and cellular immune responses"],"prefix":"10.1111","volume":"23","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-0108-5715","authenticated-orcid":false,"given":"Li","family":"Song","sequence":"first","affiliation":[{"name":"Jiangsu Key Laboratory of Zoonosis, Jiangsu Co\u2010innovation Center for Prevention and Control of Important Animal Infectious Diseases and Zoonoses, Key Laboratory of Prevention and Control of Biological Hazard Factors (Animal Origin) for Agrifood Safety and Quality of the Ministry of Agriculture and Rural Affairs of China, Joint International Research Laboratory of Agriculture and Agri\u2010product Safety of the Ministry of Education, Yangzhou University  Yangzhou China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yaya","family":"Wen","sequence":"additional","affiliation":[{"name":"Jiangsu Key Laboratory of Zoonosis, Jiangsu Co\u2010innovation Center for Prevention and Control of Important Animal Infectious Diseases and Zoonoses, Key Laboratory of Prevention and Control of Biological Hazard Factors (Animal Origin) for Agrifood Safety and Quality of the Ministry of Agriculture and Rural Affairs of China, Joint International Research Laboratory of Agriculture and Agri\u2010product Safety of the Ministry of Education, Yangzhou University  Yangzhou China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yu","family":"Zhou","sequence":"additional","affiliation":[{"name":"Jiangsu Key Laboratory of Crop Genomics and Molecular Breeding, Key Laboratory of Plant Functional Genomics of the Ministry of Education, College of Agriculture and Co\u2010Innovation Center for Modern Production Technology of Grain Crops of Jiangsu Province Yangzhou University  Yangzhou China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hui","family":"Zhang","sequence":"additional","affiliation":[{"name":"Jiangsu Key Laboratory of Zoonosis, Jiangsu Co\u2010innovation Center for Prevention and Control of Important Animal Infectious Diseases 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