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Morphological Pattern and Molecular Signaling During Intervertebral and Epiphyseal Fusion in Cetaceans and Terrestrial Mammals

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Morphological Pattern and Molecular Signaling During Intervertebral and Epiphyseal Fusion in Cetaceans and Terrestrial Mammals MORPHOLOGICAL PATTERN AND MOLECULAR SIGNALING DURING INTERVERTEBRAL AND EPIPHYSEAL FUSION IN CETACEANS AND TERRESTRIAL MAMMALS A dissertation submitted To Kent State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy by Meghan Marie Moran August 2012 Dissertation written by Meghan Marie Moran B.A., University of Illinois, Urbana-Champaign, 2002 M.A., Western Michigan University, 2004 Ph.D., Kent State University, 2012 Approved by ____________________________, Chair, Doctoral Dissertation Committee J.G.M. Thewissen ____________________________, Members, Doctoral Dissertation Committee Christopher J. Vinyard ____________________________, Walter E. Horton, Jr. ____________________________, Samuel D. Crish Accepted by ____________________________, Director, School of Biomedical Sciences Robert V. Dorman ____________________________, Dean, College of Arts and Sciences John R. D. Stalvey ii TABLE OF CONTENTS LIST OF FIGURES ............................................................................................... iv LIST OF TABLES ................................................................................................ vii ACKNOWLEDGEMENTS ................................................................................... viii Chapter Page I INTRODUCTION ....................................................................................... 1 II SACRAL ONTOGENETIC ENDOCHONDRAL OSSIFICATION OF THE INTERVERTEBRAL DISC IN MICE (MUS MUSCULUS) .......... 13 III PATTERNS OF INTERVERTEBRAL AND EPIPHYSEAL FUSION IN CETACEANS COMPARED TO TERRESTRIAL MAMMALS .............. 56 IV INTERVERTEBRAL CERVICAL FUSION IN CETACEANS COMPARED TO INTERVERTEBRAL SACRAL FUSION IN TERRESTRIAL MAMMALS ..................................................................... 97 V IDENTIFICATION OF THE SACRUM IN MODERN CETACEANS ....... 122 VI A REVIEW OF INTERVERTEBRAL AND EPIPHYSEAL FUSION IN CETACEANS AND TERRESTRIAL MAMMAL ................... 128 REFERENCES ................................................................................................. 138 iii LIST OF FIGURES Figures Page 1 Intervertebral disc diagram ........................................................................ 6 2 Regions of the intervertebral disc ............................................................ 14 3 Protein signaling ...................................................................................... 20 4 Histology of the lumbosacral joint (L6S1) in mouse ................................. 32 5 Histology of the first sacral joint (S1S2) in mouse .................................... 34 6 Growth plate and blood vessels of intervertebral disc .............................. 36 7 Immunohistochemistry of the lumbosacral joint (L6S1) in mouse ............ 38 8 Immunohistochemistry of the lumbosacral joint (L6S1) in mouse ............ 40 9 Immunohistochemistry of the first sacral joint (S1S2) in mouse ............... 42 10 Immunohistochemistry of the first sacral joint (S1S2) in mouse ............... 44 11 Protein expression similar in non-fusing and fusing intervertebral joints .................................................................................. 46 12 protein expression different between non-fusing and fusing intervertebral joints ........................................................................ 48 13 VonKossa histology for mineralization in mouse ...................................... 51 14 Land mammal sacra ................................................................................ 68 15 Archaic cetacean sacra............................................................................ 70 16 Bowhead whale cervical specimens and cervical fusion maps ................ 72 17 Mouse postnatal ontogenetic fusion maps ............................................... 74 iv LIST OF FIGURES (Continued) Figure Page 18 Pig postnatal ontogenetic fusion maps .................................................... 76 19 Archaic cetacean fusion maps ................................................................. 77 20 Bowhead and beluga whale fusion maps................................................. 79 21 Four articulation points altered by evolution mapped on an alpaca pelvis ............................................................................................ 81 22 Cetacean phylogeny of sacral fusion ....................................................... 82 23 Cleared and stained Stenella attenuata ................................................. 87 24 Phylogeny of cervical fusion related to neck length ................................. 90 25 Semicircular canal size overlaid on the cervical fusion and neck length data ...................................................................................... 93 26 Intervertebral disc diagram ...................................................................... 98 27 MMP13 protein expression in bowhead cervical intervertebral discs ................................................................................. 105 28 TIE2 protein expression in bowhead cervical intervertebral discs ................................................................................. 107 29 GDF5 protein expression in bowhead cervical intervertebral discs ................................................................................. 108 30 BMP2/4 protein expression in bowhead cervical intervertebral discs ................................................................................. 110 31 Skeletonized bowhead whale ontogenetic cervical series ..................... 111 32 CT scans of bowhead neck .................................................................... 113 v LIST OF FIGURES CONTINUED Figure Page 33 Bowhead body length compared to neck length .................................... 114 34 Cleared and stained bowhead fetus ...................................................... 114 35 Intervertebral fusion patterns of bowhead whale and mouse ................. 115 36 Mouse and bowhead immunohistochemistry results ............................. 124 37 Pudendal nerve of modern cetaceans ................................................... 125 38 Cleared and stained cetacean fetuses ................................................... 132 vi LIST OF TABLES Table Page 1 Bowhead Whale (Balaena mysticetus) Samples ................................... 102 2 Mouse and Bowhead Immunohistochemistry Results............................ 117 vii ACKNOWLEDGEMENTS First I would like to thank my advisor, Dr. Hans Thewissen for his unwavering support and encouragement during my progress as a graduate student and during the completion of this dissertation. Hans provides unique research opportunities, excellent discussion, and a genuine attitude towards his students and science. I am truly thankful to him for the wonderful opportunities he has provided for me. I thank Dr. Walt Horton for accepting me into his lab in my first year of graduate school and providing me with an excellent environment to learn lab techniques and engage in scientific discovery. Dr. Chris Vinyard is thanked for his excellent advice, instruction, and discussions on science. Dr. Sam Crish is thanked for serving on my committee and his support during this dissertation process. Thank you to Dr. Mary Ann Raghanti for acting as Graduate Student Representative and Moderator during my defense. Others I would like to extend a thank you to are, Dr. Dean Dluzen of NEOMED who helped me start my first mouse colony; Dr. Bill Lynch and his graduate students Jaclyn Stenger and Sandra Cardona also of NEOMED for donating mouse cadavers; Dr. Makarand Risbud and his technician Renata Skubutyte of Thomas Jefferson University for their completion of ANK immunohistochemistry and collaboration; Dr. Terrance Demos of Loyola viii University Medical Center for his willingness to donate his and his departments services in CT scanning fossil and modern whale vertebrae; Dr. John Mort and Dr. Eunice Lee of Shriners Hospital Laboratories, McGill University, Montreal, Canada for their generous donation of Collagen X antibody; Jason Herreman of the North Slope Borough Department of Wildlife Management for his hard work and heavy lifting during Balaena mysticetus sample collection; Denise McBurney of NEOMED for all her scientific support, friendship, and laughs; Sharon Usip of NEOMED for her patience, guidance, and teaching in the lab; Beth Lowder for troubleshooting Von Kossa staining; and Lee Brucato, my last summer student, who worked on mouse immunohistochemistry. I received excellent administration and support from Diane Kehner, Debbie Heeter, Diana Dillon, Debbie Severt, Margaret Weakland, and Judy Wearden. I would like to thank the library staff especially Lisa Barker, Laura Colwell, and Denise Cardon for all their hard work in collecting sources during this project and their willingness to search for those hard to find pieces of literature. I also thank the Comparative Medicine Unit staff, Lora Nicholson, Debbie Dutton, Linda McCort, and Dr. Walter Horne for their support and guidance with my mouse colony and IACUC protocols. I would like to thank Dr. J. Craig George and Dr. Robert Suydam of the Department of Wildlife Management in Barrow, AK; the Alaska Eskimo Whaling ix Commission, and local subsistence hunters; Dr.Terrence Demos and Dr. John F. Moran of Loyola University Medical Center for their roles in completing the CT scans of the whales; Dr. Sirpa Nummela of the University of Helsinki for her role in this project; Dr. Larry
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