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mRNA vaccines against H10N8 and H7N9 influenza viruses of pandemic potential are immunogenic and well tolerated in healthy adults in phase 1 randomized clinical trials
Open accessAbstract
BACKGROUND: We evaluated safety and immunogenicity of the first mRNA vaccines against potentially pandemic avian H10N8 and H7N9 influenza viruses. METHODS: Two randomized, placebo-controlled, double-blind, phase 1 clinical trials enrolled participants between December 2015 and August 2017 at single centers in Germany (H10N8) and USA (H7N9). Healthy adults (ages 18-64 years for H10N8 study; 18-49 years for H7N9 study) participated. Participants received vaccine or placebo in a 2-dose vaccination series 3 weeks apart. H10N8 intramuscular (IM) dose levels of 25, 50, 75, 100, and 400 µg and intradermal dose levels of 25 and 50 µg were evaluated. H7N9 IM 10-, 25-, and 50-µg dose levels were evaluated; 2-dose series 6 months apart was also evaluated. Primary endpoints were safety (adverse events) and tolerability. Secondary immunogenicity outcomes included humoral (hemagglutination inhibition [HAI], microneutralization [MN] assays) and cell-mediated responses (ELISPOT assay). RESULTS: H10N8 and H7N9 mRNA IM vaccines demonstrated favorable safety and reactogenicity profiles. No vaccine-related serious adverse event was reported. For H10N8 (N = 201), 100-µg IM dose induced HAI titers ≥ 1:40 in 100% and MN titers ≥ 1:20 in 87.0% of participants. The 25-µg intradermal dose induced HAI titers > 1:40 in 64.7% of participants compared to 34.5% of participants receiving the IM dose. For H7N9 (N = 156), IM doses of 10, 25, and 50 µg achieved HAI titers ≥ 1:40 in 36.0%, 96.3%, and 89.7% of participants, respectively. MN titers ≥ 1:20 were achieved by 100% in the 10- and 25-µg groups and 96.6% in the 50-µg group. Seroconversion rates were 78.3% (HAI) and 87.0% (MN) for H10N8 (100 µg IM) and 96.3% (HAI) and 100% (MN) in H7N9 (50 µg). Significant cell-mediated responses were not detected in either study. CONCLUSIONS: The first mRNA vaccines against H10N8 and H7N9 influenza viruses were well tolerated and elicited robust humoral immune responses. ClinicalTrials.gov NCT03076385 and NCT03345043.
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“Those studies showed that solicited systemic adverse events tended to be more frequent and more severe with higher doses and after the second vaccination.”
From An mRNA Vaccine against SARS-CoV-2 — Preliminary Report (2020) · cites this paper as [11]
Experimental — accuracy not yet measuredThe cited source says what the claim attributes to it.
In the citing paper
“This descriptive safety profile is similar to that described in a report of two trials of avian influenza mRNA vaccines (influenza A/H10N8 and influenza A/H7N9) that were manufactured by Moderna with the use of an earlier lipid nanoparticle capsule formulation11 and is consistent with an interim report of a phase 1–2 evaluation of a Covid-19 mRNA vaccine encoding the S receptor-binding domain.6”
From An mRNA Vaccine against SARS-CoV-2 — Preliminary Report (2020) · cites this paper as [11]
In the citing paper
“Two RNA vaccine studies encoding other antigens did not see durable antibody titers in humans or non-human primates”
From Adaptive immunity to SARS-CoV-2 and COVID-19 (2021) · cites this paper as Feldman et al., 2019
In the citing paper
“Current clinical trials showed that the mRNA-1440 vaccine against H10N8 and the mRNA-1851 vaccine against H7N9 influenza viruses were well tolerated and elicited robust humoral immune responses when tested separately (Table 4).360”
From Advances in COVID-19 mRNA vaccine development (2022) · cites this paper as [360]
In the citing paper
“Adverse effects were limited to pain at the injection site, redness, muscle pain, joint pain, headache, fatigue and chills/common-cold-like symptoms, indicating that the vaccine is safe and well tolerated150.”
From mRNA vaccines for infectious diseases: principles, delivery and clinical translation (2021) · cites this paper as [150]
In the citing paper
“in phase I trials of the Moderna influenza H10N8 vaccine, severe adverse events from the 400 μg dose stopped further evaluation of that dose150.”
From mRNA vaccines for infectious diseases: principles, delivery and clinical translation (2021) · cites this paper as [150]
In the citing paper
“Moderna already reported on the good tolerability of an intramuscular/intradermal nucleoside-modified mRNA vaccine against two Influenza viruses (H10N8 and H7N9). Typical adverse events related to immune activation such as myalgia, fatigue and headache were reported which were mostly mild to moderate in severity and short-lived, not requiring medical interventions [16]”
From The dawn of mRNA vaccines: the COVID-19 case (2021) · cites this paper as [16]
In the citing paper
“Indeed, both i.m. and i.d. administration of lipid nanoparticle–mRNA vaccines produce robust immune responses at a well-tolerated dose in human trials156,157.”
From Lipid nanoparticles for mRNA delivery (2021) · cites this paper as [157]
In the citing paper
“Haemagglutinin is an essential surface antigen of influenza viruses156,157.”
From Lipid nanoparticles for mRNA delivery (2021) · cites this paper as [157]
In the citing paper
“Lipid nanoparticle–mRNA vaccines are also being investigated as influenza vaccines29,41,156,157,159,184,189–191, with some formulations in clinical trials (Table 1).”
From Lipid nanoparticles for mRNA delivery (2021) · cites this paper as [157]
In the citing paper
“A 100-μg dose level induced 78.3% (HAI) and 87.0% (MN) seroconversion for H10N8 (refs156,157) and a 50-μg dose level resulted in 96.3% (HAI) and 100% (MN) seroconversion for H7N9 (refs156,157).”
From Lipid nanoparticles for mRNA delivery (2021) · cites this paper as [157]
In the citing paper
“A 100-μg dose level induced 78.3% (HAI) and 87.0% (MN) seroconversion for H10N8 (refs156,157) and a 50-μg dose level resulted in 96.3% (HAI) and 100% (MN) seroconversion for H7N9 (refs156,157).”
From Lipid nanoparticles for mRNA delivery (2021) · cites this paper as [157]
In the citing paper
“The mRNA-1440 and mRNA-1851 vaccines, which are composed of lipid nanoparticles and mRNA encoding haemagglutinin from H10N8 and H7N9 influenza viruses, respectively, have completed phase I clinical studies (NCT03076385 and NCT03345043)156,157.”
From Lipid nanoparticles for mRNA delivery (2021) · cites this paper as [157]
In the citing paper
“Initial results with lipid-nanoparticle-formulated mRNA vaccines against H10N8 and H7N9 influenza viruses and SARS-CoV-2 indicate that these vaccines are well tolerated and that they produce a robust humoral response 76,77 .”
From Nanotechnology approaches for global infectious diseases (2021) · cites this paper as [77]
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Induction of broadly reactive influenza antibodies increases susceptibility to autoimmunity
Labombarde, J. G., Pillai, M. R., Wehenkel, M., Lin, C.-Y., Keating, R., Brown, S. A., Crawford, J. C., Brice, D. C., Castellaw, A. H., Mandarano, A. H., et al. (2022) · Cell Reports · doi:10.1016/j.celrep.2022.110482 · Cited by 20
“In support of this, there is evidence of a significant decline in the levels of broadly reactive influenza antibodies following different vaccine strategies to novel influenza strains (Andrews et al., 2017; Feldman et al., 2019; Fukuyama et al., 2020; Wheatley et al., 2015).”
BackgroundEdwards, D. K., & Carfı́, A. (2022) · Current Opinion in Immunology · doi:10.1016/j.coi.2022.102214 · Cited by 18
“Both vaccines were well-tolerated and raised protective immune responses [75], demonstrating that mRNA–LNP vaccines can be well-tolerated and immunogenic in humans.”
BackgroundHu, X., Liu, C., Rcheulishvili, N., Wang, Y., Xiong, T., Xie, F., Wang, X., Chen, R., Wang, P. G., & He, Y. (2025) · Microbiology Spectrum · doi:10.1128/spectrum.02909-24 · Cited by 14
Kordyukova, L. V., & Shanko, A. V. (2021) · Biochemistry (Moscow) · doi:10.1134/s0006297921070026 · Cited by 13
Recent Advances in Messenger Ribonucleic Acid (mRNA) Vaccines and Their Delivery Systems: A Review
Belay, W. Y., Nibret, G., & Aschale, Y. (2023) · Clinical Pharmacology Advances and Applications · doi:10.2147/cpaa.s418314 · Cited by 16
“Infusing immunogenic material into the bloodstream may cause a cytokine storm, or the overwhelming synthesis of cytokines, which can result in shock and death.305 The most popular method of administering vaccines to patients is through intramuscular injection.306,307 After IM injection, the LNPs are efficiently taken…”
BackgroundA Rift Valley fever mRNA vaccine elicits strong immune responses in mice and rhesus macaques
Bian, T., Hao, M., Zhao, X., Zhao, C., Luo, G., Zhang, Z., Fu, G., Lu, Y., Chen, Y., Wang, Y., et al. (2023) · npj Vaccines · doi:10.1038/s41541-023-00763-2 · Cited by 14
Kayesh, M. E. H., Kohara, M., & Tsukiyama–Kohara, K. (2024) · International Journal of Molecular Sciences · doi:10.3390/ijms25115909 · Cited by 14
“In a Phase 1 clinical trial, mRNA vaccines against H10N8 and H7N9 were found to be well-tolerated and to elicit robust humoral immune responses [132].”
BackgroundAdvances in Development of mRNA-Based Therapeutics
Huang, L., Zhang, L., Li, W., Li, S., Wen, J., Li, H., & Liu, Z. (2020) · Current topics in microbiology and immunology · doi:10.1007/82_2020_222 · Cited by 16
Message in a bottle: mRNA vaccination for influenza
Shartouny, J. R., & Lowen, A. C. (2022) · Journal of General Virology · doi:10.1099/jgv.0.001765 · Cited by 15
“A phase I trial of lipid nanoparticle mRNA vaccines encoding the full- length HA of H10N8 or H7N9 IAV, avian viruses with pandemic potential, was performed to examine safety of both intramuscular and intradermal administration [86].”
BackgroundSofroniou, A., & Ridley, A. (2025) · British Journal of Biomedical Science · doi:10.3389/bjbs.2025.14557 · Cited by 15
“The earliest in fl uenza mRNA-based vaccines to enter clinical trials, mRNA-H10N8 (NCT03076385) [26] and mRNA-H7N9 (NCT03345043) [27], were well tolerated with strong humoral immune responses in humans when administered intramuscularly as a two-dose regimen [108].”
Han, A. R., Lee, D., Kim, S. K., Choo, C., Park, J. C., Lee, J. R., Choi, W. J., Jun, J. H., Rhee, J. H., Kim, S. H., et al. (2022) · Daehan saengsik uihak hoeji/Clinical and experimental reproductive medicine · doi:10.5653/cerm.2022.05225 · Cited by 12
“It has been reported that the existing mRNA vaccines against influenza virus, rabies virus, and Zika virus are safe during pregnancy and have good immunogenicity profiles [33-35].”
BackgroundAgent-based model of the impact of higher influenza vaccine efficacy on seasonal influenza burden
Krauland, M. G., Zimmerman, R. K., Williams, K. V., Raviotta, J. M., Harrison, L. H., Williams, J. V., & Roberts, M. S. (2022) · Vaccine X · doi:10.1016/j.jvacx.2022.100249 · Cited by 12
Considerations on the Design of Lipid-based mRNA Vaccines Against Cancer
Meulewaeter, S., Zhang, Y., Wadhwa, A., Fox, K., Lentacker, I., Harder, K. W., Cullis, P. R., Smedt, S. C. D., Cheng, M. H. Y., & Verbeke, R. (2023) · Journal of Molecular Biology · doi:10.1016/j.jmb.2023.168385 · Cited by 13
Reneer, Z. B., Bergeron, H. C., Reynolds, S., Thornhill-Wadolowski, E., Feng, L., Bugno, M., Truax, A. D., & Tripp, R. A. (2024) · PLoS ONE · doi:10.1371/journal.pone.0297833 · Cited by 16
Parveen, A., & Elkordy, A. (2024) · Pathogens · doi:10.3390/pathogens13060500 · Cited by 13
“The initial mRNA vaccinations against the influenza viruses H7N9 and H10N8 produced strong humoral immune responses and were well tolerated [53].”
BackgroundMuraduzzaman, A. K. M., Illing, P. T., Mifsud, N. A., & Purcell, A. W. (2022) · Viruses · doi:10.3390/v14112578 · Cited by 13
Influenza vaccines: where we are, where we are going.
Khalil, N., & Bernstein, D. I. (2022) · PubMed · doi:10.1097/mop.0000000000001103 · Cited by 9
Kok, A., Fouchier, R. A. M., & Richard, M. (2021) · Vaccines · doi:10.3390/vaccines9121465 · Cited by 11
“6% against H7N9 and 100% and 87% against H10N8, as measured by HI assay (titers 1 >= 40) or MN (titers >= 20) assay, respectively [79].”
MethodCAR‐T cell therapeutic avenue for fighting cardiac fibrosis: Roadblocks and perspectives
Abdalla, A. M. E., Yu, M., Ahmed, A. I. M., Meng, N., & Ouyang, C. (2024) · Cell Biochemistry and Function · doi:10.1002/cbf.3955 · Cited by 16
“147 However, LNP ‐ mRNA formulations may require different features as vaccines 148,149 4E, which may affect oxygen and nutrient exchange in the heart.”
BackgroundmRNA Vaccines Against COVID-19 as Trailblazers for Other Human Infectious Diseases
Brandi, R., Paganelli, A., D’Amelio, R., Giuliani, P., Lista, F. R., Salemi, S., & Paganelli, R. (2024) · Vaccines · doi:10.3390/vaccines12121418 · Cited by 13
Clinical Development of mRNA Vaccines: Challenges and Opportunities
August, A., Brito, L., Paris, R., & Zaks, T. Z. (2022) · Current topics in microbiology and immunology · doi:10.1007/82_2022_259 · Cited by 12
Self-citationA Thermal-Stable Protein Nanoparticle That Stimulates Long Lasting Humoral Immune Response
Wong, T., Liou, G., & Kan, M.-C. (2023) · Vaccines · doi:10.3390/vaccines11020426 · Cited by 12
A Review of the Progress of COVID-19 Vaccine Development
Ullah, S., Al-Sehemi, A. G., Klemeš, J. J., SAQIB, S., GONDAL, S. M. A., SAQIB, S., ARSHAD, A., Saqib, H., Mukhtar, A., IBRAHIM, M., et al. (2021) · Düzce Tıp Fakültesi Dergisi · doi:10.18678/dtfd.890089 · Cited by 12
“The specificity of antigen plans is important for having both relaxed and imperative reactions to the production of hazards of pandemics and scourges (81).”
BackgroundMethodMeyer, M., Wang, Y., Edwards, D. K., Smith, G. R., Rubenstein, A. B., Ramanathan, P., Mire, C. E., Pietzsch, C. A., Chen, X., Ge, Y., et al. (2021) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2021.01.25.428136 · Cited by 12
“Nucleoside modified mRNA, a relatively new addition to the arsenal of vaccine platforms, has shown promise against numerous viral infectious diseases in preclinical trials (1, 2), and phase 1 and 2 trials that are completed (3, 4) or currently underway (5-7).”
BackgroundResearch progress of mRNA vaccines for infectious diseases
Zhang, Z., Du, J., Zhang, D., Han, R., Wu, X., & Liang, Y. (2025) · European journal of medical research · doi:10.1186/s40001-025-03060-x · Cited by 19
“They are not only well tolerated but also elicit a robust humoral immune response, offering a novel method for the prevention of future pandemic influenza [145].”
Sandelius, Å., Naseer, H., Lindqvist, J., Wilson, A., & Henderson, N. (2024) · Bioanalysis · doi:10.1080/17576180.2024.2360361 · Cited by 12
Advantages of Broad-Spectrum Influenza mRNA Vaccines and Their Impact on Pulmonary Influenza
Cheng, Z., Ma, J., & Zhao, C. (2024) · Vaccines · doi:10.3390/vaccines12121382 · Cited by 11
Zhang, M., Van, L. M., & Amiji, M. M. (2025) · Molecular Therapy — Nucleic Acids · doi:10.1016/j.omtn.2025.102686 · Cited by 11
Non-capsular based immunization approaches to prevent Streptococcus pneumoniae infection
Silva, P. H., Vázquez, Y., Campusano, C., Retamal-Díaz, A. R., Lay, M. K., Muñoz, C. A., González, P. A., Kalergis, A. M., & Bueno, S. M. (2022) · Frontiers in Cellular and Infection Microbiology · doi:10.3389/fcimb.2022.949469 · Cited by 12
“Several groups have worked in RNA vaccines before in treatments for cancer and other viruses like influenza (Kranz et al., 2016; Alberer et al., 2017; Feldman et al., 2019).”
BackgroundLau, Y. M. A., Pang, J., Tilstra, G., Couture‐Senécal, J., & Khan, O. F. (2022) · Expert Opinion on Drug Delivery · doi:10.1080/17425247.2022.2144827 · Cited by 11
Elalouf, A., Kedarya, T., Elalouf, H., & Rosenfeld, A. (2023) · Journal of Genetic Engineering and Biotechnology · doi:10.1186/s43141-023-00574-x · Cited by 11
“Using mRNA can also improve pharmaceutical manufacturing since mRNA production is less complex than recombinant technology [39].”
BackgroundRNA Vaccines against Infectious Diseases: Vital Progress with Room for Improvement
Abdelzaher, H. M., Gabr, A. S., Saleh, B. M., Gawad, R. M. A., Nour, A. A., & Abdelnaser, A. (2021) · Vaccines · doi:10.3390/vaccines9111211 · Cited by 12
“Safety and reactogenicity profiles for doses up to 100 μg of H10N8 and H7N9 mRNA vaccines appeared to be comparable to that of more traditional adjuvanted and unadjuvanted influenza vaccines [71].”
BackgroundMethodmRNA vaccines: hope beneath the hype
Extance, A. (2021) · BMJ · doi:10.1136/bmj.n2744 · Cited by 11
Applications and challenges of biomaterial mediated mRNA delivery
Fang, H., & Chen, Q. (2022) · Exploration of Targeted Anti-tumor Therapy · doi:10.37349/etat.2022.00093 · Cited by 12
“administrations of mRNA vaccines were able to induce robust immune response at a well-tolerated dose [92, 93].”
BackgroundYin, J., Chen, Y., Li, Y., Zhang, X., & Wang, C. (2022) · Human Vaccines & Immunotherapeutics · doi:10.1080/21645515.2022.2119763 · Cited by 11
Vaccines on demand, part II: future reality
Geall, A. J., Kis, Z., & Ulmer, J. B. (2023) · Expert Opinion on Drug Discovery · doi:10.1080/17460441.2022.2147501 · Cited by 10
Qoura, L. A., Kostyushev, D. S., Parodi, A., Boyarintsev, D. I., Chulanov, V., & Pokrovsky, V. S. (2026) · Cellular and Molecular Bioengineering · doi:10.1007/s12195-026-00888-z · Cited by 12
du Preez, H. N., Lin, J., Maguire, G. E. M., Aldous, C. M., & Kruger, H. G. (2024) · European Journal of Clinical Investigation · doi:10.1111/eci.14296 · Cited by 12
“45,46 The Pfizer and Moderna COVID-19 mRNA vaccines are synthetic nucleoside-modified mRNA vaccines formulated in LNPs, which encode either the tri-merized receptor-binding domain (RBD) of the Sp in S1 (BNT162b1) or the prefusion stabilized full-length Sp of SARS- CoV-2 (Moderna and BNT162b2).”
A Closer Look at the Avian Influenza Virus H7N9: A Calm before the Storm?
Nakhaie, M., Rukerd, M. R. Z., Shahpar, A., Pardeshenas, M., Khoshnazar, S. M., Khazaeli, M., Bashash, D., Nezhad, N. Z., & Charostad, J. (2024) · Journal of Medical Virology · doi:10.1002/jmv.70090 · Cited by 11
“…cell culture ‐ based antigen vaccines, live attenuated influenza vaccines, figure vaccines, mRNA vaccines, phylogenetically related seed strain vaccines, recombinant baculovirus expression vaccines, and adjuvant ‐ containing H7 ‐ based vaccines, could enhance vaccination capabilities [17, 19, 20].”
Levine, M. Z., Holiday, C., Bai, Y., Zhong, W., Liu, F., Jefferson, S. N., Gross, F. L., Tzeng, W., Fries, L. F., Smith, G., et al. (2022) · Vaccines · doi:10.3390/vaccines10111856 · Cited by 10
“The VLP vaccine with ISCOMATRIX, has also been shown to elicit strong N9 NAI responses [12].”
BackgroundFlu RNA Vaccine: A Game Changer?
Meurens, F. (2020) · Vaccines · doi:10.3390/vaccines8040760 · Cited by 9
“Both approaches are in the pipelines of the three current major players in the field of RNA vaccines: Moderna Therapeutics (Cambridge, MA, USA) [30], EpiVax (Providence, RI, USA) and CureVac AG (Tübingen, Germany).”
BackgroundAlishetty, S., Carrasco, M. J., Alameh, M., Paige, M. A., Said, H., Wright, L., Narayanan, A., Alem, F., Hernandez, K. R., Gillevet, P. M., et al. (2021) · Research Square · doi:10.21203/rs.3.rs-798453/v1 · Cited by 10
Caution before fractionating COVID-19 vaccines
Wilder‐Smith, A., Desai, S. N., Cravioto, A., Nohynek, H., & Hombach, J. (2021) · Nature Medicine · doi:10.1038/s41591-021-01534-z · Cited by 10
La nueva generación de vacunas de ARN mensajero (ARNm) frente a la gripe
Reina, J. (2021) · Enfermedades Infecciosas y Microbiología Clínica · doi:10.1016/j.eimc.2021.07.009 · Cited by 11
Delivery of gene editing therapeutics
Kevadiya, B. D., Islam, F. S., Deol, P., Zaman, L. A., Mosselhy, D. A., Ashaduzzaman, M., Bajwa, N., Routhu, N. K., Singh, P. A., Dawre, S., et al. (2023) · Nanomedicine Nanotechnology Biology and Medicine · doi:10.1016/j.nano.2023.102711 · Cited by 9
mRNA vaccine-a desirable therapeutic strategy for surmounting COVID-19 pandemic
Chen, P., Shi, X., He, W., Zhong, G., Tang, Y., Wang, H., & Zhang, P. (2022) · Human Vaccines & Immunotherapeutics · doi:10.1080/21645515.2022.2040330 · Cited by 10
Metadata
- Authors
- Feldman, R. A., Fuhr, R., Smolenov, I., (Mick) Ribeiro, A., Panther, L., Watson, M., Senn, J. J., Smith, M., Almarsson, Ӧ., Pujar, H. S., Laska, M. E., Thompson, J., Zaks, T., & Ciaramella, G.
- Affiliations
- Moderna Therapeutics (United States) (US)
- Venue
- Vaccine
- Publisher
- Elsevier BV
- Published
- 2019-05
- Volume
- 37
- Issue
- 25
- Pages
- 3326-3334
- ISSN
- 0264-410X (ISSN-L)
- OpenAlex
- W2944588091
- Cited by
- 454
- References
- 36
- Citations by year
- 2026: 27 · 2025: 41 · 2024: 66 · 2023: 60 · 2022: 87 · 2021: 114 · 2020: 53 · 2019: 6
- FWCI
- 49.46
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Licence
- Locations
- Vaccine (publishedVersion, cc-by-nc-nd)
- PubMed (publishedVersion)
- Funders
- Moderna
- Topics
- Health Sciences › Medicine › Epidemiology › Influenza Virus Research Studies
- Health Sciences › Medicine › Infectious Diseases › SARS-CoV-2 and COVID-19 Research
- Life Sciences › Immunology and Microbiology › Microbiology › Bacterial Infections and Vaccines
- Keywords
- mRNA vaccines, H7N9 influenza virus, pandemic influenza preparedness, vaccine safety, vaccine immunogenicity, intradermal vaccination, intramuscular vaccination, dose escalation, hemagglutination inhibition, microneutralization, cell-mediated immunity, healthy adults, reactogenicity, healthy volunteers, vaccine tolerability
- MeSH terms
- Immunogenicity, Vaccine, Adolescent, Adult, Antibodies, Viral, Dose-Response Relationship, Drug, Double-Blind Method, Female, Humans, Influenza, Human, Influenza Vaccines, Male, Middle Aged, RNA, Viral, Young Adult, Healthy Volunteers, Influenza A Virus, H7N9 Subtype, Influenza A Virus, H10N8 Subtype
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