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Enhancing Myoblast Fusion for Therapy of Muscular Dystrophies

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Enhancing Myoblast Fusion for Therapy of Muscular Dystrophies Enhancing Myoblast Fusion for Therapy of Muscular Dystrophies The Harvard community has made this article openly available. Please share how this access benefits you. Your story matters Citation Wu, Melissa P. 2013. Enhancing Myoblast Fusion for Therapy of Muscular Dystrophies. Doctoral dissertation, Harvard University. Citable link http://nrs.harvard.edu/urn-3:HUL.InstRepos:11151535 Terms of Use This article was downloaded from Harvard University’s DASH repository, and is made available under the terms and conditions applicable to Other Posted Material, as set forth at http:// nrs.harvard.edu/urn-3:HUL.InstRepos:dash.current.terms-of- use#LAA Copyright © 2013 by Melissa P. Wu All Rights Reserved Dissertation Advisor: Professor Emanuela Gussoni Melissa P. Wu Enhancing Myoblast Fusion for Therapy of Muscular Dystrophies Abstract Skeletal muscle is a major organ comprising 30-40% of the human body mass. The coordination of processes resulting in mature muscle requires many genes, and their loss can result in debilitating muscle disorders. Of the strategies being developed to cure muscle diseases, enhancement of the natural process of muscle cell fusion in existing or introduced myogenic cells has great therapeutic potential. In this work, we determined whether a drug that stimulates proliferation and fusion of myoblasts could alleviate murine Duchenne muscular dystrophy. We also studied the necessity of a gene that is upregulated in early fusing human myoblast cultures and its role in muscle disease development. Carbamylated erythropoietin (C-EPO) is an analog of erythropoietin, which promotes myoblast proliferation and inhibits fibrosis. We treated mdx mice with intraperitoneal injections of C-EPO for 4 and 12 weeks, and monitored weight, serum creatine kinase levels, and changes in muscle histology. We observed moderate histological improvement at 4 weeks that did not translate into a clinically significant improvement at 12 weeks. At the dosages tested, C-EPO was not an effective long-term therapeutic for the treatment of a Duchenne muscular dystrophy. We also studied the role of GPR56 in muscle. Loss of GPR56 causes bilateral frontoparietal polymicrogyria, which shares the brain phenotype of dystroglycanopathies but not the severe muscle defects. We found that GPR56 is transiently upregulated in iii differentiating myocytes and nascent myotubes. In muscle cultures, its loss results in decreased myoblast fusion and smaller myotubes. During in vivo muscle regeneration, the expression of myogenic regulators is delayed in GPR56 knockout mice; however, there are no gross morphological changes. These mild defects likely result from decreased signaling from GPR56 to SRE and NFAT-directed pathways for myoblast differentiation. Therefore, GPR56 promotes myoblast differentiation and fusion through activation of SRE and NFAT signaling pathways, but its loss in muscle in vivo is compensated for by other complementary factors. In summary, these studies add to the existing literature on the role of a specific gene in muscle cell fusion and disease development, and of a specific drug in augmenting myoblast fusion in the context of muscle disease. iv Table of Contents List of Figures and Tables ........................................................................................................... viii Acknowledgments ............................................................................................................................ x Chapter 1: Introduction ................................................................................................................. 1 Skeletal Muscle ................................................................................................................................................2 Skeletal Muscle Development ...................................................................................................2 Skeletal Muscle Structure ...........................................................................................................4 Genetic Disorders of Skeletal Muscle ...................................................................................................7 Therapies for Duchenne Muscular Dystrophy .............................................................................. 12 Hypothesis and Overview of Dissertation ...................................................................................... 16 References ...................................................................................................................................................... 18 Chapter 2: Carbamylated erythropoietin does not alleviate signs of dystrophy in mdx mice ................................................................................................................................................... 31 Attribution of Collaborator Contributions...................................................................................... 32 Abstract ........................................................................................................................................................... 33 Introduction .................................................................................................................................................. 33 Materials and Methods ............................................................................................................................. 36 Results .............................................................................................................................................................. 38 Discussion ....................................................................................................................................................... 42 Acknowledgements.................................................................................................................................... 47 References ...................................................................................................................................................... 48 Chapter 3: Establishing a role for GPR56 in the muscle:..................................................... 52 Attribution of Collaborator Contributions...................................................................................... 53 v Abstract ........................................................................................................................................................... 54 Introduction .................................................................................................................................................. 55 Transcription Factor Regulation of Myogenic Differentiation ............................... 55 Cell-surface Receptor Regulation of Myogenic Differentiation ............................. 57 Identification of Novel Regulators of Myogenesis ....................................................... 58 The G-protein Coupled Receptor 56 May Play a Role in Myogenesis ................. 59 Methods ........................................................................................................................................................... 63 Results .............................................................................................................................................................. 78 Discussion ....................................................................................................................................................... 97 References ................................................................................................................................................... 109 Chapter 4: Summary of Accomplishments and Significance of Findings ..................... 120 Advances in Our Understanding of Myoblast Differentiation and Fusion.................... 121 Implications of Our Studies on Developing Therapies For Clinical Use ........................ 122 Lessons From Testing the Administration of C-EPO in Mdx Mice..................... 122 Lessons From Studying the Effects of Loss of GPR56 on Muscle Development ............................................................................................................................................................ 125 Future Directions ..................................................................................................................................... 127 Developing Therapeutics for Muscular Dystrophies ............................................... 127 Understanding the Process of Muscle Cell Differentiation and Fusion........... 127 References ................................................................................................................................................... 129 Appendix I: Investigation of the Appearance of Tubular Aggregates in GPR56 knockout mouse tissue.............................................................................................................. 132 Attribution of Collaborator Contributions................................................................................... 133 vi Introduction ............................................................................................................................................... 134 Methods ........................................................................................................................................................ 139 Results ..........................................................................................................................................................
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