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Efficacy of Liposomal M2e Vaccines in Combination With EFFICACY OF LIPOSOMAL M2E VACCINES IN COMBINATION WITH ADJUVANTS OR LIPOSOMAL ADJUVANT VACCINES WITHOUT M2E IN MICE INFECTED WITH INFLUENZA H1N1 OR H3N2 A Thesis Presented to the Faculty of California State Polytechnic University, Pomona In Partial Fulfillment Of the Requirements for the Degree Master of Science In Biological Sciences By Paulina G. Villanueva 2017 SIGNATURE PAGE THESIS: EFFICACY OF LIPOSOMAL M2E VACCINES IN COMBINATION WITH ADJUVANTS OR LIPOSOMAL ADJUVANT VACCINES WITHOUT M2E IN MICE INFECTED WITH INFLUENZA H1N1 OR H3N2 AUTHOR: Paulina G. Villanueva DATE SUBMITTED: Spring 2017 Biological Sciences Department Dr. Jill Adler-Moore Thesis Committee Chair Biological Sciences Dr. Nancy Buckley Biological Sciences Dr. Jamie Snyder Biological Sciences ii ACKNOWLEDGEMENTS I want to thank all the members of Dr. Adler’s laboratory for always being supportive. Thank you Jenny, Hernan, Jon, Eden, Fady, Cole, and Sherleen for always helping me during my spleen collections. Jenny, thank you for becoming one of my great friends. I am so happy to know that we will pursue our dreams in Los Angeles. Don’t forget to come visit me. Thank you Fady for being the best influenza team partner I could have asked for. You are a kind, intelligent, and a thoughtful person and I am so happy to know that your dream of becoming a physician is coming true. You deserve all the good that is coming your way. Thank you Eden for coming into the lab during the most challenging time of my career at Cal Poly Pomona. I know that you are going to do amazing things for the influenza team. Most importantly, thank you for being my friend. Jonathan, thank you for being my research mentor and friend. I will never forget that you always made time to help me during your most stressful times at Cal Poly Pomona. Dr. Adler, thank you for accepting me as a graduate student. Without your mentorship and support, my dream of becoming a physician would not be a reality. Thank you for always pushing me to do more and for always making time to get to know me as a person. To my committee members, Dr. Adler, Dr. Buckley, and Dr. Snyder, thank you from the bottom of my heart for working so hard to help me complete my thesis. I feel blessed to have been mentored by three incredibly accomplished women in science. I came to Cal Poly Pomona to find the kind of mentorship that you have given me. I hope that one day, I can shape someone’s future the way you help me shape mine. To my boyfriend, Nick Shadwell, thank you for always supporting me and for believing in my dreams. It is because of your motivation and support that I will soon be a medical student at UCLA. iii Thank you for always being by my side. I am so happy for everything that is coming our way. Lastly, Mom and Dad, thank you for helping me become the person that I am today. You brought me to the United States with big dreams for me. Because of your sacrifices, my dreams are coming true. You are my inspiration. iv ABSTRACT Introduction: Influenza A viruses are a major cause of acute respiratory diseases in humans. Different influenza A strains have been responsible for seasonal infections (epidemics) and worldwide pandemics. All of these pandemics and epidemics arise from the mutations or genetic reassortment of viral RNA coding for hemagglutinin (HA) and neuraminidase (NA) surface proteins of influenza A viruses, which inevitably leads to antigenic shift (pandemics) and antigenic drift (epidemics). Because of this, we have worked in collaboration with Molecular Express Inc. on the VesiVax® delivery system which incorporates the viral protein M2e epitope, a 15 amino acid (aa) residue at the amino end of the highly conserved region of the M2 protein, into a unilamellar liposome less than 100 nm in size. The M2e protein can be conjugated to the surface of the liposomes via a maleimide linker (CALV) or fused to a hydrophobic protein (HD), which allows the M2e to be incorporated into the liposome bilayer. This technology was used to investigate various combinations of vaccines composed of M2e and different immunological adjuvants or adjuvants without the M2e protein for their efficacy in Swiss Webster female mice against challenge with Influenza A/Puerto Rico/8/1934 (PR8) H1N1 or X-31 H3N2. Methods: Study 1, the vaccine formulations were as follows: L-CMI-100 μg mycoviral dsRNA-M2e, L-CMI-15 μg MPL-M2e, L-CMI-15μg MPL, L-CMI-100 μg mycoviral dsRNA and PBS. The mycoviral dsRNA was obtained from the mycovirus present in Saccharomyces cerevisiae. Mice were vaccinated subcutaneously on d0 and intranasally (IN) boosted on d28 and d56. Sera (n=5/gp) were collected 3 days after the last boost and tested for anti-whole virus IgG and anti-M2e IgG isotype concentrations. The remaining v mice were IN challenged 2 weeks post-boost with 10X LD50 Influenza A/Puerto Rico/8/1934 (PR8) H1N1 or 10X LD50 (X-31) H3N2. Lungs were collected 5 days after infection (n=5/gp) and tested for viral burden via foci assay. The remaining mice (n=10/gp) were monitored for morbidity to day 28 post-challenge. In Study 2, the vaccine formulations were as follows: L-CMI-50 μg mycoviral dsRNA (UC)-M2e, L-CMI-25 μg mycoviral dsRNA (UC)-M2e, L-CMI-25 μg mycoviral dsRNA(C)-M2e, L-CMI-15 μg mycoviral dsRNA (C)-M2e, L-CMI-5 μg mycoviral dsRNA (C)-M2e, Non-CMI-50 μg mycoviral dsRNA (UC) and PBS. The experiment was performed as in Study 1, but only the H1N1 challenge was used and lungs were not collected for this study. In Study 3, the vaccine formulations were as follows: L-MPL-M2e (1-12), L-MPL-M2e (1-12)cD, L- mycoviral dsRNA-M2e (1-12), L-MPL-M2e (1-15), L-6 μg LT1-M2e (1-15), L-3 μg LT1/7.5 μg MPL-M2e (1-15), L-3 μg LT1-M2e (1-15), L-Pam3CAG-M2e (1-15), L-MPL (No CMI), and PBS. As in Study 3, mice were only challenged with H1N1. In Study 4, the vaccine formulations were as follows: L-MPL-M2e-HD, L-CMI-Pam3CAG, L-CMI- MPL, L-CMI-mycoviral dsRNA, L-CMI-1V270, L-CMI-LT1 and L-MPL (No CMI). Mice were primed and boosted as in Study 1, but viral challenge was done 1 week after the last boost with 10X LD50 H1N1. Mice (n=10/gp) were monitored for morbidity to day 28 post-challenge. Results: L-CMI-100 μg mycoviral dsRNA-M2e provided significantly better protection compared to MPL and PBS control groups against an H1N1 and H3N2 challenge as measured by survival and decreased viral burden (p<0.014). L-CMI-100 μg mycoviral dsRNA also provided significantly better protection compared to the MPL control group (p<0.024) against H1N1 challenge and MPL and PBS control groups (p<0.01) and vi against H3N2 challenge as measured by survival (p<0.024). L-CMI-50 μg mycoviral dsRNA (UC)-M2e provided significantly better protection compared to L-CMI-25 μg mycoviral dsRNA (C)-M2e as measured by survival against an H1N1 challenge (p<0.024). Vaccine formulations L-Pam3CAG-M2e (1-15), L-LT1/MPL-M2e (1-15) and L-mycoviral dsRNA-M2e (1-12) provided the most protection as measured by survival. Lastly, L-MPL-M2e-HD, L-CMI-Pam3CAG, L-CMI-mycoviral dsRNA and L-CMI-MPL provided significantly better protection L-MPL (No CMI) as measured by survival, against an H1N1 challenge (p<0.048). Conclusions: These studies demonstrated for the first time that a CMI liposomal M2e vaccine formulation containing 100 μg of mycoviral dsRNA could provide cross protection against two strains of influenza virus (H1N1 and H3N2) in Swiss Webster female mice. We also showed that the antigen M2e (1-15) was more effective than the shorter antigen M2e (1-12) when incorporated into a CMI liposomal vaccine. Additionally, we observed that CMI liposomal vaccine formulations containing only an adjuvant without a protein antigen could provide protection against a lethal challenge of influenza virus. vii TABLE OF CONTENTS Signature Page…………………………………………………………………………….ii Acknowledgments……………………………………………………………………..iii-iv Abstract………………………………………………………………………………..v-vii List of Tables ………………………………………………………………………xvi-xxi List of Figures ……………………………………………………………………xxii-xxvi Chapter 1: Introduction……………………………………………………………...…1-68 I. Influenza Infection: Transmission and Disease Symptoms…………………..1-5 II. Orthomyxoviridae Viruses …………………………………………………..5-9 A. Types of Influenza Viruses……………………………………………5-9 1. Influenza A…………………………………………………...6-7 2. Influenza B …………………………………………………..7-8 3. Influenza C …………………………………………………..8-9 III. Influenza A Outbreaks……………………………………………………..9-14 A. Overview…………………………………………………………….9-10 B. Influenza Pandemic of 1918………………………………………..10-11 C. Influenza Pandemic of 1957………………………………………..11-12 D. Influenza Pandemic of 1968………………………………………..12-13 E. Influenza Pandemic of 2009 …………………………………………..13 F. Continuous Threats: H5N1 and H7N9 Influenza Virus ……………13-14 IV. Antigen Drift and Antigen Shift in Influenza…………………………….14-16 V. Influenza Viral Proteins ………………………………………………….16-21 VI. Viral Nomenclature………………………………………………………21-22 viii VII. Influenza Replication Cycle……………………………………………..22-25 A. Viral Entry………………………………………………………….22-23 B. Intracellular Replication……………………………………………23-24 C. Viral Budding………………………………………………………24-25 VIII. Antivirals…………………………………………………………….…25-29 A. Amantadine and Rimantadine……………………………………...26-27 B. Oseltamivir and Zanamivir…………………………………………27-29 IX. Immunity to Influenza……………………………………………………29-46 A. Innate Immune System……………………………………………..30-37 1. Cellular Innate Immune Response…………………………30-34 2. Inflammasome……………………………………………..34-35 3. Complement………………………………………………..36-37 B. Adaptive Immune Response………………………………………..38-46 1. T Cell Maturation………………………………………….39-40 2. Th1 Response/Cell Mediated………………………………40-42 3. Th2 Response/Antibody Mediated………………………...42-44 4. Th17/Inflammatory Response……………………………..44-45 5.
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