Putting Stress to the Test: a Critical Evaluation of the Biological Response and Physical

Putting Stress to the Test: A Critical Evaluation of the Biological Response and Physical Manifestation of Stress in the Human Skeleton by Amy B. Scott A Thesis submitted to the Faculty of Graduate Studies of The University of Manitoba in partial fulfilment of the requirements of the degree of DOCTORATE OF PHILOSOPHY Department of Anthropology University of Manitoba Winnipeg, MB, Canada Copyright © 2015 Amy B. Scott ABSTRACT Using traditional osteological methods of stress analysis, specifically skeletal lesions and growth and development patterns, this research focuses on the timing and manifestation of stress in the hard tissues of the body and the interaction between these various indicators of poor health. The Danish Black Friars cemetery population spanning the medieval and post-medieval periods (13th-17th centuries) is used for this research to explore changing health trends during a period of socioeconomic disruption and expanding urban dwelling. The results of this study show a distinct trend between the late and early post-medieval periods where stress was elevated after the Reformation (AD 1536). Additionally, females generally show higher levels of stress than males. The poorer health experienced in the post-medieval period was likely influenced by the changing living conditions of the late 16th century where urban dwelling increased in Denmark introducing new pathogens, poorer living conditions and new environmental and working stressors, all contributing to poorer health. While the overall level of stress appears to increase into the post-medieval period, the average age-at-death is higher after the Reformation suggesting that while these individuals may have been exposed to more prolonged periods of stress, they were able to adapt and survive in spite of these hardships. This research also examines the ability to analyse stress through the extraction of ancient proteins from bone. Using MircoBCA and enzyme-linked immunoassay (ELISA) analysis, it is shown that protein preservation is good in the Black Friars population and the level of protein reflects the expected fluctuations associated with the demography of the population (i.e. age, sex) and not necessarily the influence of stressors. Overall this study provides a comprehensive examination of stress from its initial biological signal through to skeletal disruption in a climate of expanding urban development and changing socioeconomics in Denmark. ii ACKNOWLEDGEMENTS I remember the day that Rob Hoppa asked me to come to the University of Manitoba to do my PhD with him; it was before I had even begun my MA and had no idea where my life would take me. All these years later, and I truly could not have made a better choice. Words can never express how incredibly thankful I am to have had a mentor like Rob helping me navigate the academic waters; he was my life vest on many occasions and I am so thankful he was there. I do not know how he put up with me over these past five years, but am so grateful that he did and will always look back fondly on my time here at the University of Manitoba with him. I wouldn’t have made it this far without you Rob and cannot say how much your faith in me has meant. Thank you. I would also like to thank my PhD committee, Dr. Jesper Boldsen, Dr. Tom Klonisch and Dr. Linda Larcombe, for all of their guidance, patience and support for this project. It truly takes a great committee to build a successful dissertation and I know, as a team, we have done just that. Thank you to the Odense City Museums for access to the Black Friars collection and the ADBOU team, Bodil Theilade, Dorthe Dangvard Pedersen, Peter Tarp and Ulla Høg Freund in Denmark for your help during and after the data collection process. Dr. Lillian Skytte and Dr. Kaare Rasmussen, thank you for your guidance and help with my biochemical sample collection. Thank you also to the radiography team at the Odense University Hospital for your enthusiasm and participation in this research, Susanne Vestbjerg, Annika Hansen and Hanne Kjærby. A special thank you to my biochemical collaborators in the Department of Immunology at the University of Manitoba, Grace Choi, Dr. Neeloffer Mookherjee and Peyman Ezzati. Without your expertise and contribution, the biochemical aspect of this research would have been impossible, it truly was a pleasure working with all of you. Additionally, I would like to thank iii Dr. Matthew Collins at the University of York for his expertise and advice. A special thank you also to Dr. Mai-Britt Mosbech for her translation work on the Danish archaeological site reports. I would also like to thank the various funding resources that contributed to this research: SSHRC Vanier Graduate Scholarship (770-2012-0159); SSHRC Michael Smith Foreign Study Supplement (771-2012-0113); SSHRC Standard Research Grant (410-2011-1408); Dr. Emöke J.E. Szathmáry Graduate Fellowship in Biological Anthropology; Manitoba Graduate Scholarship; and at the University of Manitoba the Department of Anthropology, the Faculty of Arts, the Faculty of Graduate Studies. I am humbled and honoured to have been provided with these opportunities and recognize their pivotal importance in the success of this research. Finally, I would like to take this opportunity to extend my deepest gratitude to my family and friends for the wrath they likely endured these past five years. To my roller derby team, you have changed my life for the better. I love and admire each and every one of you and thank you for letting me get out my academic frustrations on the track. This dissertation belongs to all of you as much as it does to me. To my friends near and far who have never seemed to tire of keeping me motivated and laughing through all of the ups and downs of this academic career, you mean the world to me and I am thankful every day to have you in my life, KG, MS, TH, MT, AB, LS, TB, AG, JR, EB, DL, GML, DM, AK. To my family, I know since the beginning this career path has not always been clear or fathomable, but we have managed to make it happen. I say “we” because this journey we took together as a family. Thank you for always believing in me, I appreciate it more than words can say. Thank you to my grandparents, who without fail every week for the last five years have asked me “how is school going?” Your interest in my research has always been appreciated. Most importantly, to my parents, I love you and could not have done any of this without you, thank you beyond measure. iv For Ulla v TABLE OF CONTENTS PAGE Title Page i Abstract ii Acknowledgements iii Dedication v Table of Contents vi List of Tables x List of Figures xvii List of Appendices xx Copyrighted Material xxi Chapter 1: Introduction 1 1.1 Purpose and Significance 2 1.2 Research Objectives 5 Chapter 2: Literature Review – Historical Context 9 2.1 Introduction 9 2.2 The Dominican Order 9 2.3. The Odense Black Friars Monastery and Cemetery 10 2.4 Danish Historical Timeline 14 2.5 The Early and High Medieval Period (AD 1050 – 1536) 16 2.5.1 The Emerging Danish Nation during the Middle Ages 16 2.5.2 The Spread of Christianity and Population Growth 20 2.5.3 Emerging Agriculture and Diet during the Medieval Period 21 2.5.4 Climatic Shift between the Early and High Medieval Periods 24 2.5.5 The Black Death 26 2.5.6 Early and High Medieval Population Demography 28 2.5.7 Cattle Farming 30 2.6 The Post-medieval Period (AD 1536 – 1660) 31 2.6.1 Political Unrest and Shifting Borders 31 2.6.2 Post-medieval Population Demography 34 vi 2.6.3 Diet during the Post-medieval Period 35 2.7 Summary of the Medieval to Post-medieval Period in Denmark 37 Chapter 3: Literature Review – Osteological Context 39 3.1 Introduction 39 3.2 Skeletal Structure and Growth 39 3.2.1 Osteoblast Cells and the Creation of Bone 40 3.2.2 Osteoclast Cells and Bone Remodelling 41 3.2.3 Osteocyte Cells and Bone Maintenance 42 3.3 Bone Mineralization 43 3.3.1 Intramembranous Ossification 43 3.3.2 Endochondral Ossification 44 3.4 Bone Growth 44 3.4.1 Growth Plates and Linear Growth 44 3.4.2 Appositional Growth 45 3.4.3 Hormonal Influence on Skeletal Growth 45 3.5 The Stress Response 46 3.5.1 Skeletal Stress and Glucocorticoids 47 3.5.2 Osteocalcin 49 3.5.3 Archaeological Studies of Osteocalcin 51 3.6 Bioarchaeological Evidence of Skeletal Stress 53 3.7 Bringing the Stress Data Together 61 3.8 Theoretical Limitations in the Study of Skeletal Stress 68 3.8.1 The Osteological Paradox 68 3.8.2 Considerations of the Paradox 71 3.9 Methodological Limitations in the Study of Skeletal Stress 74 3.9.1 Lesion-based Assessments of Stress 74 3.9.2 Linear Growth Assessments of Stress 77 Chapter 4: Materials and Methods 80 4.1 Black Friars Skeletal Material 80 4.1.1 Selection Criteria 80 4.2 Archaeological Dating of the Black Friars Population 80 4.2.1 Arm Position 80 4.2.2 Burial Location 81 4.2.3 Relative Dating Techniques 82 vii 4.3 Sex Determination and Age Estimation 85 4.4 Pathological Analysis 88 4.5 Stress Lesion Data 89 4.5.1 Cribra Orbitalia (CO) and Porotic Hyperostosis (PH) 90 4.5.2 Enamel Hypoplastic Lesions (EHL) 91 4.5.3 Harris Lines (HL) 93 4.6 Growth and Development Data 95 4.6.1 Body Size Indicators (BSI) 95 4.6.2 Cortical Bone Thickness 102 4.7 Biochemical Data 102 4.7.1 Protein Extraction 104 4.7.2 MicroBCA Analysis 106 4.7.3 Enzyme-linked Immunoassay (ELISA) 107 Chapter 5: Results 109 5.1 Introduction 109 5.2 Intra-observer Error 109 5.3 Lesion-based Analyses 114 5.3.1 Pathology 114 5.3.2 Stress Indicators 128 5.4 Growth and Development-based Analyses 152 5.4.1 Body Size Indicators

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