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Murine Metapodophalangeal Sesamoid Bone Mineralization: a Light and Electron Microscopy Study

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Murine Metapodophalangeal Sesamoid Bone Mineralization: a Light and Electron Microscopy Study MURINE METAPODOPHALANGEAL SESAMOID BONE MINERALIZATION: A LIGHT AND ELECTRON MICROSCOPY STUDY A thesis submitted to Kent State University in partial fulfillment of the requirements for the degree of Master of Arts by Alison R. H. Doherty December, 2007 Thesis written by Alison R. H. Doherty B.A., University of Wyoming, 2003 M.A., Kent State University, 2007 Approved by ________________________________________ William J. Landis, Ph.D, Advisor ________________________________________ C. Owen Lovejoy, Ph.D, Co-Advisor ________________________________________ Richard S. Meindl, Ph.D, Chair, Department of Anthropology ________________________________________ John R. Stalvey, Ph.D, Dean, College of Arts and Sciences ii Table of Contents List of Figures ................................................................................................................ iv List of Tables................................................................................................................... v Acknowledgments .......................................................................................................... vi Introduction ..................................................................................................................... 1 Literature Review ........................................................................................................ 2 Vesicle-Mediated Mineralization ............................................................................. 3 Collagen-Mediated Mineralization ........................................................................... 5 Sesamoid Bones....................................................................................................... 7 Fibrocartilage........................................................................................................... 9 Materials and Methods .................................................................................................. 12 Sample Acquisition ................................................................................................... 12 Radiography .............................................................................................................. 13 Histology ................................................................................................................... 13 Whole Mount Staining ............................................................................................... 20 Transmission Electron Microscopy Sample Determination ........................................ 23 Transmission Electron Microscopy ............................................................................ 23 Results ........................................................................................................................... 27 Radiography .............................................................................................................. 27 Histology ................................................................................................................... 29 Whole Mount Staining ............................................................................................... 38 Transmission Electron Microscopy Sample Determination ........................................ 45 Transmission Electron Microscopy ............................................................................ 46 Discussion ..................................................................................................................... 54 Conclusion .................................................................................................................... 60 Future Research ......................................................................................................... 61 Literature Cited ............................................................................................................. 64 Appendix ....................................................................................................................... 69 iii List of Figures Figure 1: Murine Front and Hind Paw Radiographs .................................................... 28 Figure 2: Murine Metatarsophalangeal Joint Anatomy ................................................ 30 Figure 3: Two-Week-Old Metatarsophalangeal Sesamoid ........................................... 31 Figure 4: Two-Week-Old Metatarsophalangeal Sesamoid Under Polarized Light ....... 32 Figure 5: Metatarsophalangeal Sesamoid Fibrocartilaginous Insertion Site ................. 33 Figure 6: Proteoglycan Content of the One-Week Metatarsophalangeal Sesamoid ...... 34 Figure 7: One-Week-Old Metacarpophalangeal Sesamoid .......................................... 35 Figure 8: One-Week-Old Metatarsophalangeal Sesamoid ........................................... 35 Figure 9: Two-Week-Old Metacarpophalangeal Sesamoid .......................................... 36 Figure 10: Three-Week-Old Metacarpophalangeal Sesamoid ........................................ 37 Figure 11: Three-Week-Old Metatarsophalangeal Sesamoid ......................................... 37 Figure 12: Whole Mount Double-Stained One-Week-Old Mouse ................................. 39 Figure 13: One-Week-Old Alizarin Red S Stained Front and Hind Paws....................... 40 Figure 14: Double-Stained One-Week-Old Hind Paw ................................................... 41 Figure 15: The First Mineralized Sesamoid ................................................................... 42 Figure 16: Secondary Ossification Centers in the One-Week-Old Hind Paw ................. 43 Figure 17: Three-Week-Old Alizarin Red S Stained Front and Hind Paws .................... 44 Figure 18: Thick Section of the Two-Week-Old Metacarpophalangeal Sesamoid.......... 47 Figure 19: Matrix Vesicles in the Extracellular Space between Cells ............................ 48 Figure 20: Matrix Vesicles near Chondrocytes .............................................................. 49 Figure 21: Chondrocytes within Calcifying Cartilage .................................................... 50 Figure 22: Perpendicular Collagen Fiber Orientation .................................................... 51 Figure 23: Areas of Parallel Collagen Fiber Orientation ................................................ 51 Figure 24: Thick Section of the Three-Week-Old Metacarpophalangeal Sesamoid ........ 52 iv List of Tables Table 1: Calcium Oxalate Endpoint Test ....................................................................... 14 14 Table 2: Tissue Processing ............................................................................................ 15 15 Table 3: Toluidine Blue Stain ....................................................................................... 17 17 Table 4: Picrosirius Red Stain ....................................................................................... 18 18 Table 5: Safranin-O Red Stain ...................................................................................... 19 19 Table 6: Whole Mount Staining .................................................................................... 22 22 Table 7: TEM Processing and Embedding .................................................................... 24 24 Table 8: Formvar Coated Grids ..................................................................................... 25 25 Table 9: Positive-Contrast Staining Grids ..................................................................... 26 26 v Acknowledgments I express my deepest gratitude to my loving and supporting husband, Adam Doherty, for standing behind me all the way in this endeavor. I could not have made it without his encouragement and willingness to edit. I am also very appreciative to Elizabeth Lowder and the training she gave me in all aspects of histology, radiography, and general laboratory procedures. When I ran into roadblocks, Beth was always my brainstorming partner and backup plan expert. Special thanks go to my advisor, Dr. William Landis, for his guidance and suggestions in this research project, as well as to Dr. Owen Lovejoy and Dr. Richard Meindl for their support. In addition, this project would not have been feasible without the training and expertise of Jeanette Killius, the NEOUCOM Electron Microscopy Laboratory Coordinator. Finally, I thank Dr. Hans Thewissen, Dr. Chris Vinyard, Robin Jacquet, Janet Hamilton, my fellow graduate colleagues, and my parents for their role in supporting this thesis project. Without funding from the NEOUCOM Skeletal Biology Focus Area and the Graduate Student Senate this project would not have been possible. vi Introduction Extracellular matrix vesicles have been documented as initiating mineralization of many vertebrate calcified structures, including calcifying cartilage, tendon, fibrocartilage, and bone (Ali et al., 1970; Hsu and Anderson, 1978; Landis, 1986; Lowenstam and Weiner, 1989; Yamada, 1976). In calcifying cartilage and bone, mineral crystal growth first occurs in matrix vesicles and then quickly engulfs the surrounding organic matrix. However, vesicles in normally mineralized turkey tendons do not appear to be the only site of mineral formation. In these tendons, collagen initiates mineral growth in areas isolated from matrix vesicles, but these nucleation events always follow vesicle presence in the tissue. Therefore, the role of matrix vesicles in initiating mineralization is still under investigation in vertebrate tissues (Boskey, 1998; Landis et al., 1996; Lowenstam and Weiner, 1989).
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