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Trabecular Bone Variation in Mid and Upper Craniofacial TRABECULAR BONE VARIATION IN MID AND UPPER CRANIOFACIAL REGIONS OF FOUR ANTHROPOIDS LESLIE C. PRYOR SMITH Presented to the Faculty of the Graduate School of The University of Texas at Arlington in Partial Fulfillment of the Requirements for the Degree of MASTER OF ARTS IN ANTHROPOLOGY THE UNIVERSITY OF TEXAS AT ARLINGTON May 2011 Copyright © by Leslie C. Pryor Smith 2011 All Rights Reserved ACKNOWLEDGEMENTS I would like to thank my committee chair and advisor Dr. Shelley Smith for her commitment to the betterment of my thesis project and manuscript. I am grateful for all the support and encouragement she has given me during my time at UT Arlington. I would also like to thank my committee member, Dr. Naomi Cleghorn, at UT Arlington for her comments and suggestions to help improve this work. This thesis could not have been possible without the help of Dr. Paul Dechow at Baylor College of Dentistry. I have benefited tremendously from his impressive wealth of knowledge and experience, not just in primate anatomy and biomechanics, but in the broader range of biomedical sciences. I want to thank Dr. Lynne Opperman, professor at Baylor College of Dentistry, for her time, advice and encouragement throughout this project. I also thank Allen Mortimer and Mitra Bolouri for their work in the initial stages of this project. It would be ungracious not to mention the love and support I have received from my friends and family since I began this project. In particular, I would like to thank my father, Gerald, my mother, Claire, and my brother, Jerry for nurturing my curiosity and truth seeking nature. Last but not least, I thank my best friend and husband, Colby. He has shown me unrelenting patience, encouragement, and support throughout my time in graduate school. I am grateful to him for keeping me laughing even during the most stressful of times. I dedicate this thesis to my mother, Claire, for teaching me the importance of diligence. April 8, 2011 iii ABSTRACT TRABECULAR BONE VARIATION IN MID AND UPPER CRANIOFACIAL REGIONS OF FOUR ANTHROPOIDS Leslie Claire Pryor Smith, M.A. The University of Texas at Arlington, 2011 Supervising Professor: Shelley L. Smith Some posit that robust supraorbital tori seen in fossil hominin species are an adaptation to hard object feeding, while others believe that supraorbital morphology is overdesigned for mastication in robust species. Strain gage analysis of primate craniofacial regions finds that strain is much lower in the supraorbital region relative to the zygomatic region, however, the mechanisms behind bone adaptation to loads is not fully understood. While variations in cortical bone material properties in a few primate species have been determined, no one has explored the variation in cranial trabecular bone. This study used microcomputed tomography to assess the variation in trabecular bone structure in the supraorbital and zygomatic regions of 10 humans, 4 baboons, 1 chimpanzee, and 1 gorilla. Significant differences in the degree of anisotropy, an indication of trabecular bone strength relative to orthogonal axes, were region specific, but significant differences in bone volume fraction, positively correlated with density, were species specific. Furthermore, differences in the degree of anisotropy by region were consistent with differences in cortical iv bone surface strain determined by other researchers using strain gage analysis. Trabecular bone structural variables also differed in distribution by species and region. The results of this study suggest that trabeculae adapt their primary orientation in response to relative strain endured during mastication, but that other variables related to density and strength may have a genetic component. Moreover, the results of this study support the hypothesis that robust craniofacial features may be genetic or epigenetic adaptations to fallback feeding strategies. There is a tremendous amount of variation in trabecular bone morphometric variables among species and between regions, suggesting that trabecular bone orientation and structure plays an essential role in craniofacial bone adaptation. v TABLE OF CONTENTS ACKNOWLEDGEMENTS ................................................................................................................iii ABSTRACT ..................................................................................................................................... iv LIST OF ILLUSTRATIONS.............................................................................................................. ix LIST OF TABLES ............................................................................................................................ xi Chapter Page 1. INTRODUCTION……………………………………..………..….. ..................................... 1 1.1 The Adaptive Significance of the Supraorbital Region in Primates ................. 1 1.2 Hypotheses ...................................................................................................... 5 1.3 Conclusion ........................................................................................................ 5 2. BACKGROUND .............................................................................................................. 7 2.1 Introduction....................................................................................................... 7 2.2 The Mechanical Properties of Trabecular Bone .............................................. 8 2.2.1 Introduction ...................................................................................... 8 2.2.2 Bone: a Dynamic Multi-level Structure ............................................. 8 2.2.3 Introduction to Bone Biomechanics ............................................... 11 2.3 Historical Outline and Technique Development ............................................. 14 2.3.1 Pre 20 th Century: Discovering the Relationship between Bone Form and Function .................................................................................. 14 2.3.2 Trabecular Bone Research in the Last 50 Years ........................... 15 2.3.3 Conclusion ..................................................................................... 26 2.4 The Mechanical Significance of Supraorbital and Zygomatic Regions in Anthropoids .......................................................................................................... 27 2.4.1 Introduction .................................................................................... 27 vi 2.4.2 Endo and the Beam Model............................................................. 28 2.4.3 Conclusion ..................................................................................... 30 2.5 Finite Element Analysis and Primate Evolution ............................................. 31 2.5.1 Introduction .................................................................................... 31 2.5.2 Experimentation ............................................................................. 32 2.6 The Mechanical Properties of Cortical Bone in Humans and Primates ........ 34 2.7 The Importance of Cellular Accommodation to Bone Remodeling ................ 37 2.8 Conclusion ..................................................................................................... 40 3. MATERIALS AND METHODS ..................................................................................... 42 3.1 Sample ........................................................................................................... 42 3.2 Methods .......................................................................................................... 43 3.2.1 MicroCT Scanning and Analysis ................................................... 43 3.3 Anatomical Regions of Interest ...................................................................... 50 3.3.1 Supraorbital Region (SO) ............................................................... 50 3.3.2 The Zygoma ................................................................................... 51 3.3.3 The Zygomatic Arch ....................................................................... 51 3.4 Statistical Analysis ......................................................................................... 53 4. RESULTS ..................................................................................................................... 55 4.1 Univariate Analysis and Significance Testing ................................................ 55 4.1.1 Bone Volume/Total Volume Density Analysis ................................ 55 4.1.2 Trabecular Bone Morphometric Analysis ....................................... 64 4.2 Principle Components Analysis: Patterning of Trabecular Bone Properties by Species ........................................................................................................... 74 4.2.1 Supraorbital Region ....................................................................... 76 4.2.2 Zygoma .......................................................................................... 77 4.2.3 Zygomatic Arch .............................................................................. 78 vii 5. DISCUSSION AND CONCLUSION……………………………………..………..….. ....... 80 5.1 Discussion ...................................................................................................... 80 5.1.1 Strain Gradients in Primate Supraorbital and Zygomatic Regions ................................................................................ 80 5.1.2 Intra and Interspecific
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