SKELETAL BIOMECHANICS of the FLORIDA MANATEE (Trichechus Manatus Latirostris) by KARI BETH CLIFTON a DISSERTATION PRESENTED TO
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SKELETAL BIOMECHANICS OF THE FLORIDA MANATEE (Trichechus manatus latirostris) By KARI BETH CLIFTON A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2005 Copyright 2005 by Kari Beth Clifton to all those—people and animal alike—who have loved, supported, encouraged, instructed, amazed, inspired, and empowered me ACKNOWLEDGMENTS This project was an enormous undertaking–one that I could not have completed alone. First I would like to acknowledge my supervisory committee chair (Roger Reep) and co-chair (Jack Mecholsky). I thank Roger for his unqualified generosity, encouragement, and guidance; and for feeding the spirit with food and wine. I benefited greatly from being allowed to wander off the path in pursuit of new ideas. I thank Jack for his patience in teaching a biologist to think like an engineer, and for the tailgates. My supervisory committee members (MaryBeth Horodyski, Donna Wheeler, and Tom Wronski) provided guidance, instruction, and support for which I am grateful. Many individuals contributed to the success of this project. While I was working at the Florida Department of Environmental Protection, Dr. Sentiel Rommel suggested I take a look at manatee bone. He and Dr. Tom Koob at the Shriners Hospital for Children introduced me to the field of bone mechanics. The Save the Manatee Club funded a pilot study that led to the inception of this project. I am indebted to Maggie Stoll for her invaluable technical expertise and administrative support. To my fellow graduate students in the fracture mechanics group, I am grateful for the weekly discussions of ceramics and other things. I learned a great deal about fractography and bone mechanics from Jiahau Yan. From the College of Engineering, Chuck Broward provided access to the air cannon, as well as his expert engineering skills. Billy Schultz performed the strain gauge testing. Bryan Conrad from the Department of Orthopaedics taught me to operate the iv biomechanical testing equipment, and provided invaluable technical support. Joan Yonchek lent her amazing expertise in hard and soft tissue histology. I thank Allyson Barrett and Ben Lee from the Department of Dental Biomaterials for their assistance with technical issues and data collection. This project made great strides forward with the help of veterinary students Julia Golden and Beth Carson; and undergraduate volunteers Johnattan, Rebecca, Jessica, Stephanie, Jaime, Travis, and Chris. Tissue samples were provided by the FWC’s Marine Mammal Pathobiology Lab in St. Petersburg, and by Sea World of Florida. I extend my thanks and appreciation to Dr. Sentiel Rommel, Mr. Tom Pitchford, Mr. Ken Arrison and the staff of the MMPL, and to Dr. Beth Chittick of Sea World. I wish to thank the college of Veterinary Medicine, the Marine Animal Health program, and in particular Dr. Charles Courtney for his interest and enthusiasm for my project. This project was funded by a College of Veterinary Medicine Consolidated Faculty Research award and a Marine Conservation Trust Fund award. Additional funding was provided by grants from the Sigma Xi Scientific Research Society, a College of Veterinary Medicine Graduate Student Seed Grant, and a Merck-Merial Veterinary Scholars Research Grant. The University’s Graduate Student Council and Graduate School provided travel awards so that I could attend conferences. Last, but not at all least, I would like to express my gratitude and appreciation for the love and support of my family throughout my many years of schooling. I am indebted to my nephews David, Adam, and Brain, for never, ever letting me get a lick of work done while at their house. There are, after all, more important things in life. v TABLE OF CONTENTS page ACKNOWLEDGMENTS ................................................................................................. iv LIST OF TABLES........................................................................................................... viii LIST OF FIGURES ........................................................................................................... ix ABSTRACT....................................................................................................................... xi CHAPTER 1 INTRODUCTION ........................................................................................................1 Skeletogenesis...............................................................................................................5 Bone Shape and Macrostructure...................................................................................7 Microscopic Structure of Bone.....................................................................................8 Mechanical Properties of Bone Related to Microscopic Structure.............................11 Variations in the Microscopic Structure of Vertebrate Bone .....................................14 Manatees in Perspective .............................................................................................24 2 MATERIAL PROPERTIES OF MANATEE RIB BONE.........................................29 Introduction.................................................................................................................29 Materials and Methods ...............................................................................................32 Bone Sample Collection......................................................................................32 Specimen Preparation..........................................................................................32 Material Properties Testing .................................................................................33 Bone Density and Mineral Content Determination.............................................35 Statistical Analyses..............................................................................................36 Results.........................................................................................................................36 Within-Animal Comparisons...............................................................................37 Material Properties Tests.....................................................................................37 Bone Density and Mineral Content .....................................................................38 Discussion...................................................................................................................39 The Material Properties of Manatee Bone...........................................................39 The Effect of Mineral Content on the Material Properties of Bone ....................45 Other Factors Influencing Bone Material Properties...........................................46 Summary..............................................................................................................50 vi 3 STRUCTURAL PROPERTIES OF MANATEE RIBS TESTED IN IMPACT........59 Introduction.................................................................................................................59 Materials and Methods ...............................................................................................60 Bone Sample Collection......................................................................................60 Whole Rib Impact Testing...................................................................................61 Strain Gauge Testing ...........................................................................................62 Fractal Dimensional Increment ...........................................................................63 Kinetic Energy Calculations................................................................................64 Statistical Analyses..............................................................................................65 Results.........................................................................................................................65 Whole Rib Impact Tests ......................................................................................65 Strain Gauge Tests...............................................................................................66 Kinetic Energy Calculations................................................................................67 Discussion...................................................................................................................68 The Fractal Dimensional Increment ....................................................................68 Fractal Dimensional Increment and Fracture Toughness....................................70 R-curve Behavior and Viscoelasticity in Bone ...................................................71 Impact Energy Absorption of Soft Tissues .........................................................73 Summary..............................................................................................................74 4 CONCLUSIONS AND FUTURE RESEARCH ........................................................85 Summary and Conclusions .........................................................................................85 Further Research.........................................................................................................87 Additional Avenues of Research ................................................................................90 APPENDIX A DATA TABLES .........................................................................................................92