University of Nevada, Reno

Periostitis: An Indicator of Stress and Health in Past Populations Explored

A thesis submitted in partial fulfillment of the requirements for the degree of

Bachelor of Arts in Anthropology and the Honors Program

by

Jordan N. Cramer

Dr. Marin Pilloud, Thesis Advisor

May, 2018

UNIVERSITY OF NEVADA RENO

THE HONORS PROGRAM

We recommend that the thesis prepared under our supervision by

Jordan N. Cramer

Entitled

Periostitis: An Indicator of Stress and Health in Past Populations Explored

be accepted in partial fulfillment of the requirements for the degree of

BACHELOR OF ARTS, Anthropology

______Marin Pilloud, Ph.D., Thesis Advisor

______Tamara Valentine, Ph. D., Director, Honors Program

May, 2018 i

Abstract

Periostitis is a skeletal condition where new bone is formed in response to many different insults to bone. The condition has been used as an indicator of stress in bioarchaeological research for decades and is key in research of disease in the past.

Despite the wide use of periostitis as an indicator of stress and disease there is little research into the condition itself and the process leading to the new bone formation. This thesis takes into consideration clinical and bioarchaeological literature in order to provide a better understanding of the process and causes of periostitis. In addition to analysis of literature a case study looking at prevalence of periostitis in an archaeological sample from central California is done to assess the use of periostitis as an indicator of stress and health in the past and the information that such analyses may give. A more rounded understanding of periostitis will allow research into stress in the past to better implement analysis of periostitis and as such create a better overall picture of health within the past.

ii

Acknowledgments

There are many people that I must thank for getting me where I am today.

First, I want to thank my mentor Marin Pilloud. Thank you for your patience and guidance through this long process. It is because of you that I have found an area of study that I truly love.

Secondly, I must thank my family for always supporting me and pushing me to be better and improve. I would also like to thank Bonnie for always reminding me to write and for her support. Special thanks to Robert for believing in me no matter what and putting up with me on those days that I was going crazy working on this project.

Lastly, I would like to thank the Honors Program. Thank you for believing in me and not letting me give up on myself.

Thank you to everyone who gave words of encouragement and supported me through this process. It would not have been possible without you.

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Table of Contents

Abstract ...... i Acknowledgments ...... ii List of Tables ...... v List of Figures ...... vi Introduction ...... 1 A Look at Stress ...... 7 What is Stress? Physiological Origins of Interpretation...... 7

Stress Research within Bioarchaeology ...... 9

Stress in Place of Health ...... 10

Skeletal Indicators of Stress ...... 11

The Periosteum and Periosteal Reactions...... 13 The Periosteum ...... 13

Periosteal Reactions ...... 17

Periostitis within Bioarchaeology ...... 22 Background and Methods ...... 26 Bio-Cultural Background of California Sample ...... 26

Data Collection Procedures ...... 27

Stage 1 ...... 28 Stage 2 ...... 29 Results and Analysis ...... 31 Full Sample and Temporal Analysis ...... 31

Early Period (3050-500 B.C.) ...... 34

Early Middle Period (500 B.C. – A.D. 420) ...... 36

Late Middle Period (A.D. 420-1010) ...... 38

Middle-Late Transition Period (A.D. 1010-1390)...... 40

Late Prehistoric (A.D. 1390-1720) ...... 42 iv

Protohistoric/Historic (A.D. 1720-1899) ...... 44

Overall Trends ...... 47

Conclusions ...... 49 Conclusions from Literature Review ...... 49

Conclusions from Case Study ...... 50

References ...... 54 Appendix A ...... 60 Figures ...... 60

Tables ...... 71

v

List of Tables

Table 1. Table of Prevalence by Temporal Period. 31

Table 2. Total Prevalence by Sex. 33

Table 3. Total Prevalence by Age 34

Table 4. Prevalence by Sex from 3050-500 B.C. 35

Table 5. Prevalence by Age from 3050-500 B.C. 36

Table 6. Prevalence by Age from 500 B.C. - A.D. 420 37

Table 7. Prevalence by Sex from 500 B.C. - A.D. 420 38

Table 8. Prevalence by Sex from A.D. 420-1010 39

Table 9. Prevalence by Age from A.D. 420-1010 40

Table 10. Prevalence by Sex from A.D. 1010-1390 41

Table 11. Prevalence by Age from A.D. 1010-1390 42

Table 12. Prevalence by Age from A.D. 1390-1720 43

Table 13. Prevalence by Sex from A.D. 1390-1720 44

Table 14. Prevalence by Sex from A.D. 1720-1899 45

Table 15. Prevalence by Age from A.D. 1720-1899 46

vi

List of Figures

Figure 1. Subtypes of Periosteal Reactions. 18

Figure 2.Central California Sites Included in CCBD (From Pilloud et al., 2014) 28

Figure 3. Prevalence of Periostitis per Temporal Period by Percentage 32

Figure 4. Total Percentage of Males and Females with Periostitis Present. 32

Figure 5. Total Percentage of Individuals with Periostitis Present Per Age 33

Figure 6. Percentage of Males and Females with Periostitis from 3050-500 B.C. 34

Figure 7. Percentage of Periostitis by Age from 3050-500 B.C. 35

Figure 8. Percentages of Periostitis by Age from 500 B.C. - A.D. 420 37

Figure 9. Percentage of Periostitis by Sex from 500 B.C. - A.D. 420 38

Figure 10. Percentage of Periostitis by Sex from A.D. 420-1010 39

Figure 11. Percentage of Periostitis by Age from A.D. 420-1010 40

Figure 12. Percentage of Periostitis by Sex from A.D. 1010-1390 41

Figure 13. Percentage of Periostitis by Age from A.D. 1010-1390 42

Figure 14. Percentage of Periostitis by Age from A.D. 1390-1720 43

Figure 15. Percentage of Periostitis by Sex from A.D. 1390-1720 44

Figure 16. Percentage of Periostitis by Sex from A.D. 1720-1899 45

Figure 17. Percentage of Periostitis by Age from A.D. 1720-1899 46

Figure 18. Change in Prevalence of Periostitis by Age over Time. 47

Figure 19. Prevalence of Periostitis Over Time by Sex 48

1

Introduction

Bioarchaeology works to answer questions about the archaeological past through analysis of human remains. Certain information about human lives may be observed on bone, which preserves for thousands, or even hundreds of thousands of years. Current research in the field of bioarchaeology includes analysis of basic elements such as stature, sex and age, diet, disease, and trauma. In addition to these more descriptive aspects, it is possible to study ideas such as population movement or migration, status, violence, health, and potentially much more. The diversity of subjects that may be studied under the field of bioarchaeology is made possible by the nature of bone.

One area of interest when studying the past through skeletal remains is disease, a phenomenon that continues to affect humans to this day. The study of disease in past populations, or paleopathology, is an interdisciplinary area of research with knowledge from many different disciplines, especially bioarchaeological and medical, that are needed for a full understanding as to how disease affected past populations (Ortner 2011).

From the medical field, knowledge of radiology and orthopedic pathology have importance, and bioarchaeology contributes osteological analysis as well as understanding of old, dry bone.

Orthopedic pathology, or the study of bone disease makes the study of paleopathology possible by providing understanding of disease processes observed in bone. Advances in radiology have given new ways to image bone and interpret the images obtained. The images obtained allow for a non-destructive way to view the inner workings of bone. Medical research gives new insights into bone and how it responds to disease, however; the focus is on living tissue. Osteological analysis, on the other hand, 2 is specialized in the appearance and study of bone material from people that are no longer living, whether from the distant past or more recent cases seen in forensic work.

Studying disease in the past is best done using all these methods and areas of knowledge together.

The study of paleopathology, the study of disease in the past, started in description of abnormalities observed within archaeological remains. Just as bioarchaeology as a field has developed beyond basic description, paleopathology has moved toward answering more broad picture questions. Research in the field includes disease patterning, disease evolution, temporal changes in disease frequencies, and relationships between diseases and humans. Insights as to disease in the past not only give a better understanding of conditions in the past but also allows for prediction of disease evolution or seeing broad patterns for different diseases (Ortner 2011). Through the study of disease an image of health in the past becomes possible.

However, “health” as it existed in the past proves problematic to study, health is defined by the World Health Organization (WHO) as, “a state of complete physical, mental and social well-being and not merely the absence of disease or infirmity” (World

Health Organization, 2017). When only the bones remain the mental and social well- being is impossible to know. Therefore, only the physical well-being of the individual can be studied which is influenced by many different factors, including environment, nutrition, genes, and pathogen exposure. Any deviation from physiological homeostasis can be caused by pathogens or stressors (Larsen 1999). Stress and disease exposure can be studied in relation to the “health” of an individual through analysis of skeletal material. 3

Since it is difficult, if not impossible, to determine a holistic understanding of health as it was in the past based on skeletal material, the use of the term stress instead of health has been of focus within bioarchaeology. Something to consider with the use of the term is that stress has come to have many different meanings depending upon the context. Within bioarchaeology stress is understood through biological understandings of the term, the understanding that the body physically responds to different conditions in predictable ways such as the hormone production that leads to the “fight or flight” response (Seyle, 1973). Stress has been of interest in bioarchaeology for many decades beginning with physiological interest into the term in the mid-20th century. It was with

Selye’s theory of biological stress that the concept of stress really took off within bioarchaeology (Klaus 2014). Stress is studied using different stress indicators that may be found on the skeleton.

The skeletal system’s main role is that of support and protection of the many organs in the body; however, these functions are not its sole purpose. Bone is also an important part of the musculoskeletal system that works to enable mobility and motion by being the anchor point for many different muscles, ligaments, and tendons as well as working as a lever system. The role of bone tissue within the body goes beyond even its role in mobility, as it is used to store fats, produce red blood cells and be a source of elements such as calcium within the body (White & Folkens, 2005). The many roles that bone tissue performs has made it adapt into a dynamic tissue within the body with a boundless range of variation observed and the ability to react and change throughout an individual’s lifetime. The changes seen to bone happen both normally during growth and development, and as a reaction to different conditions. 4

Bone originally develops in one of two ways depending on the type of bone. The majority of bone in the body is formed through endochondral bone formation. During this process a cartilage template is first formed that bone later replaces. This is a long process that allows for growth in early life, such as the lengthening of the bones in the arms and legs. The second form of bone development is intramembranous bone formation, where bone is formed within a membrane. Most cranial bones, or bones that make up the skull, form in this manner as well as other flat bones like the clavicles (collar bones). This form of bone development is seen in areas where bone must develop rapidly such as to protect the developing brain.

Once formed bone only reacts in two different ways to anything affecting it.

Bone can react by forming more bone, which is done by specialized cells called osteoblasts. The other way that bone reacts is by reabsorbing already existing bone. This reabsorption is done by multi-cellular units called osteoclasts. Reabsorption and formation of bone may occur at the same time in a tethered response or separately.

When osteoblasts and osteoclasts work together in a tethered response the process is called remodeling. Remodeling occurs throughout life and is used for bone maintenance and repair. Remodeling sites are thought to develop at random, but this process may be triggered through trauma to the bone (Clarke 2008). Modeling is the independent response of osteoclasts and osteoblasts. Though modeling may occur throughout the lifetime of an individual it is most prevalent in early growth. Modeling allows bone to adapt its shape to physiological and biomechanical conditions that it is experiencing (Clarke 2008). It is through these processes that bone may react to stressors such as trauma, disease, and environmental factors. 5

One skeletal indicator that is often observed in paleopathological studies is periosteal reaction or periostitis. In this work periosteal responses are referred to as periosteal reactions and periostitis interchangeably as both indicate the same condition, an inflammatory response in the periosteum because of insult to the periosteum or underlying cortical bone, with the only difference being the emphasis. Periostitis refers to the complete process while periosteal reactions is used to refer to the new bone formation. The periosteum is a tissue membrane that surrounds most bones in the body.

It is known to be involved in bone growth and repair (Dwek 2010). Due to this role, the periosteum is often the first to respond to an insult affecting bone tissue.

Insults that most often lead to a periosteal response are trauma, infection, and disease. The response of the periosteum to different stimuli is what defines the periosteal reaction. Periostitis may have many different appearances based on the intensity, duration, and aggressiveness of the insult. The reaction often starts with inflammation of the periosteum and results in new bone growth (Rana 2009). When analyzing periostitis within bioarchaeology, the literature often labels it as an unspecific periosteal reaction.

This categorization is done because of the variety of factors that may lead to the reaction

(e.g., traumatic fracture, infection, certain cancers).

When analyzed together with additional indicators such as location of the periostitis, a more specific diagnosis may be given with caution. Even without other indicators, periostitis may give insight into conditions and stress within the past when paired with demographic information, such as age and sex. Once the mechanisms behind the condition are understood a better diagnosis of different causes of insult may also be possible. 6

Analysis of data compiled from many different archaeological sites in California for the presence of periostitis within various demographic groups through time is considered in order to explore patterning of the condition and its use as a stress indicator.

In addition to analysis, literature in both the clinical and anthropological fields are considered to create a better understanding of periostitis as a condition and its etiology

(disease process) in the past. This work hopes to give a more robust understanding of the condition and the information periostitis gives in relation to stress. With this understanding further research into stress in the past may better implement analysis of periostitis and as such create a better overall picture of health within the past.

7

A Look at Stress

Though stress is a common word in the vocabulary of many people, this simple word has come to have many different meanings. It is used to describe physical forces, biological responses, as well as being a mental or physical state. Within bioarchaeology stress is interpreted through the physiological understanding of stress, adaptations that the body makes to changing conditions. Just as the field of bioarchaeology has changed and evolved over time, so has the use and study of stress within the field.

What is Stress? Physiological Origins of Interpretation.

The concept of stress has been considered since Roman times though its meaning changed and expanded throughout history (Robinson, 2018). Currently the term stress is most associated with the psychological perspective; however, it continues to hold importance across disciplines. The meaning that is of most interest within bioarchaeology and paleopathology comes from physiological understandings of the term, the understanding that the body responds to demands affecting it in a specific, predictable manner.

Physiological interest in the concept of stress began with Claude Bernard’s (1872) concept of the milieu intérieur, or internal environment. It is in Bernard’s work that the first understandings of homeostasis, or the process of the body to maintain a consistent environment, were developed (Robinson, 2018). In the beginning of the 19th century,

Walter Cannon furthered stress research, though at this time the term stress was not used.

Cannon’s research focused mainly on the concept of homeostasis and expanding the understanding of the concept and describing potential process for maintaining 8 homeostasis in the body. The concept of fight or flight, the adrenaline response to perceived danger, was the result of Cannon’s research into homeostasis combined with his later theory of emotion (Robinson, 2018). Without the consideration and understanding of homeostasis and internal regulating systems within the body further research into stress would be obsolete.

The research conducted in the late 19th century into the early 20th century lays the groundwork for the concept of stress as a subject of study. Stress as its own concept comes to be of interest in medical research with Hans Seyle. Seyle is often credited with originating the use of the term stress within medical research, though Seyle himself often stated otherwise (Robinson, 2018). Seyle developed the concept of the general stress response, which may be triggered by a wide variety of stressors and explored the ways the body reacts to stress.

The general stress response, as described by Seyle, has three different stages. The first stage is the alarm response. During this stage the body is put into a state of alarm due to the presence of a stressor, and the body prepares itself to react to the stressor.

After continued exposure to the stressor the alarm response is followed by the stage of resistance. The stage of resistance is characterized by adaptation of the body to the stressor. During this phase the body discovers a new sense of normal and improved function. The adaptation to the stressor does not last, and the final stage of the stress response is observed. During this stage of exhaustion, the body becomes worn down and loses that adaptation to the stressor that it gained. This stage of exhaustion occurs from chronic exposure to a stressor (Seyle, 1973). From this point on the stress concept has been further researched and developed in many different disciplines. These disciplines 9 include physiology of course, but also psychology, and bioarchaeology. It is from

Seyle’s research into stress and the stress response that bioarchaeological understandings of stress are derived.

Stress Research within Bioarchaeology

Within bioanthropology stress has been studied mostly in relation to adaptation, both in contemporary times and in the prehistoric past. The focus with stress research has been to link human adaptability and biology to social phenomenon (Goodman et al.,

1988). Bioarchaeology uses markers left on bone to understand biosocial conditions in the past. Stress becomes of particular interest in research in paleopathology, both in the study of disease and in cultural responses and treatment of disease.

Within bioarchaeology, stress and adaptation go hand in hand. The process of stress and how it affects an individual is not able to be studied within bioarchaeology as all that remains to study are the results of the process on bone. Due to this inability to study stress itself, skeletal indicators of stress have come to be used to study the presence of stress. The shift in focus in studying stress also leads to an altered meaning for the word in comparison to that used by physiology. This definition has come to be known as the physiological disruption as a result of environmental factors (Larsen, 1999). A model for the interpretation of stress indicators in skeletal populations was developed by

Goodman et al. (1988) with alterations and refiguring as new knowledge came to be. An important focus of the model is the ability of cultural systems to both be a means of buffering stressors as well as a cause of stressors. The model incorporates the Seylean concept that stressors may be either positive or negative in nature but see the same 10 physiological response. Within the model there are three important factors that lead to stress; these are environmental circumstances, host resistance, and cultural systems

(Goodman et al., 1988). Each has the ability to affect the other leading to stress on the population or within individuals. It is an endless cycle which highlights the importance of health and culture when considering human adaptation.

Stress in Place of Health

Health in popular language tends to be used to describe a lack of disease or adverse conditions. In truth health is a complex concept as it encompasses many different aspects of an individual’s wellbeing (Temple & Goodman, 2014). Many of these aspects, such as mental and social well-being, are not able to be studied from skeletal remains and arguably can only be speculated. In addition to the inability to study all aspects of health from skeletal remains, there is the osteological paradox (Wood et al., 1992).

The osteological paradox describes an innate dilemma with the study of health and disease in bioarchaeology: skeletal indicators of stress do not have a direct relationship with health (Temple & Goodman, 2014). Most disease processes that are known currently affect soft tissue, or non-bone tissues, almost exclusively. Bone reacts slowly, and generally any bone response to stress is the result of a chronic condition. The presence of an indicator of stress does not necessarily mean the individual was

‘unhealthy’ as they lived long enough to have evidence of the stress on the bone. The reverse is also true, that the lack of an indicator of stress indicates ‘good health’. It is possible that the individual died before the condition could affect the bone. The result of this paradox is that prevalence of an indicator or condition does not give an accurate 11 measure of health in the past. With health being a rather vague concept with shifting definitions and the difficulty of evaluating the health of people in the past, stress has become a more prevalent focus in paleopathological, and bioarchaeological research.

Skeletal Indicators of Stress

After consideration of what stress is as a biological response to a change in the body’s natural rhythms and how it is understood in the field of bioarchaeology, an understanding of how stress manifests within the human skeleton is needed. For this understanding to be possible, potential indicators of stress need to be known.

There are two main forms of skeletal stress indicators; these are growth disruption and disease. Growth disruption occurs mostly during adolescence when an impoverished environment hinders growth. This growth disruption may lead to stinted stature, misshapen bones, and changes in bone mass (Larsen, 1999). Another stress indicator that is popularly used to explore growth disruption is linear enamel hypoplasia. Linear enamel hypoplasia are defects in the enamel of teeth that are the result of stress during development. The popularity of linear enamel hypoplasia as a marker of stress has to do with the nature of enamel. Enamel, unlike bone, does not remodel, therefore any defects that develop are permanent on the tooth (Lewis & Roberts, 1997). In addition, there is strong experimental evidence that links linear enamel hypoplasia to stress events such as disease.

The second form of stress indicators is disease, which includes dental caries

(cavities) and antemortem tooth loss, non-specific infection such as periostitis, and infections with consistent, specific infection patterns (Larsen, 1999). Skeletal indicators 12 of disease vary greatly and as such an understanding of the disease process is crucial. In addition to a wide range of variation many different disease processes have the same or similar skeletal markers, for example periostitis is seen in many disease processes.

Careful consideration of all possible causes for the condition observed on the bone is important to ensure research interpretations are a true reflection of disease in the past.

Stress has come to have many different meanings that are often not well defined.

Within bioarchaeology it is often used in relation with health and adaptation. The study of stress has helped to move bioarchaeology beyond simple typology and toward more question-based research. Periostitis is one of many different indicators of stress used in the field especially in studies of health. An understanding of periostitis and the process causing the reaction is needed to better understand the use of periostitis as an indicator of stress.

13

The Periosteum and Periosteal Reactions

When studying stress and the idea of health in the past a common indicator that is analyzed is periosteal reaction. This bone response is commonly seen with many different diseases and stressors. As such, a complete understanding of the anatomy of the periosteum, the origin of periosteal reactions, is necessary. It is the unique anatomy of the periosteum that allows periosteal reactions to be observed. In addition to the anatomy of the periosteum, the role it fulfills needs to be explored. Once the specifics of the periosteum are understood a closer look at periostitis becomes possible.

The Periosteum

The periosteum is a fibrous membrane that surrounds the outer surface of the bone diaphysis, excluding the joint surfaces (where two bones interact). On long bones, such as the bones that make up the limbs, the periosteum is thickest in the central region of the diaphysis and the membrane is easily separated from the underlying bone. The periosteum thins as it works toward the ends of the bone where it is more strongly attached to the bone (Augustin, et al., 2007).

There are many different vessels and nerves found throughout the periosteum providing the underlying bone with blood as well as hormones and other nutrients. The intense pain that occurs with a periosteal injury, especially when a bone is broken, is due to the large amount of pain nerves located within the membrane (Augustin et al., 2007).

The periosteum becomes even more complex when observed microscopically.

The periosteum is made up of two different layers of cellular tissue. The outer layer is a fibrous layer that is mostly made up of a matrix of collagen fibers with a 14 sparsity of cells (Dwek, 2010). Collagen is a common protein in connective tissues that helps to give the tissues strength and elasticity. Collagen is also found in muscle and skin tissues. It is within this fibrous layer that most of the many vessels and nerves are found within the periosteum. This fibrous outer layer gives the periosteum structure and is important for blood supply to bone. The inner layer of the periosteum is what gives this membrane its ability to prompt bone growth. Unlike the outer layer the inner layer, also called the cambium layer, is opposite in construction as it has a high abundance of cells with sparse collagen matrix (Dwek, 2010). There are a variety of cells found within this layer including osteoblasts, fibroblasts (cells that form collagen and other fibers), as well as mesenchymal progenitor cells (stem cells) and osteogenic progenitor cells. The osteogenic progenitor cells are the precursors of osteoblasts. It is the presence of stem cells and osteogenic progenitor cells that allow for the regenerative ability of the periosteum (Chang & Knothe Tate, 2012).

The periosteum serves many different purposes in the body and is far more complex than first expected by researchers. Early research into the periosteum started back in the early 18th century when Duhamel discovered the osteogenic potential of the periosteum when it has been stimulated. Duhamel observed that when silver wire was placed underneath the periosteum of animal bone new bone growth was present several weeks later (Chang & Knothe Tate, 2012). This early discovery of the ability of the periosteum to form new bone research into the workings and potential of this tissue is still making new discoveries.

As was discovered in early research into the periosteum the main role that the tissue has in the body is in stimulating new bone growth. This bone growth occurs 15 throughout development of bone starting with embryonic bone development and continues as the bone further develops as the individual ages. With intramembranous ossification the periosteum develops from the margins of the mesenchymal model.

During endochondral ossification the beginnings of the periosteum develop early in the making of the cartilage model with cells collecting on the periphery. At this early stage in endochondral ossification, the precursor to the periosteum is known as the perichondrium. Once blood vessels make their way into the perichondrium the chondrocytes (cells that form cartilage) differentiate into osteoblasts. It is with the differentiation of chondrocytes into osteoblasts that marks the periosteum’s existence.

The periosteum fully develops at different times across the bone with the epiphyses, or ends being the last to develop at the end of puberty (Dwek, 2010). Once established the periosteum begins work on establishing bone on the periphery of the newly forming bone contributing to the appositional, or layered, growth of the bone. The appositional growth is what allows the bones to get larger as they get longer. The appositional growth of bone is largely variable, and it is the periosteum that mostly performs this process. The differences in bone growth between men and women has led some researchers to propose that steroids may be a regulator. Outside factors may also play a role as it has been observed that mechanical forces acting on the periosteum induces the expression of different genes within the tissue (Orwoll, 2003). Periosteal bone formation increasing the size of bone is most pronounced during puberty; however, the process continues through adulthood at a much slower rate.

In addition to bone formation during development of bone, the periosteum is key in fracture repair and healing. Previous research has observed that the periosteum is able 16 to heal large fractures in bone that other forms of bone repair are unable to bridge

(Chang, 2012). It is in the most common form of fracture repair, endochondral repair, that the periosteum has a major role. Endochondral fracture repair of broken bone starts with the formation of a hematoma with inflammation of the area surrounding the fracture and blood from torn vessels pooling. It is the formation of the hematoma that initiates the fracture repair through recruitment of different cells that work to mend the fracture

(Schell, 2017). If the fibrous layer of the periosteum is torn at the time of fracture surrounding soft tissue supports the hematoma until the tear is repaired. Once repaired, or if not torn to begin with the fibrous layer of the periosteum supports the hematoma keeping it in place around the fracture. Cells from the inner cambium layer differentiate and lead to the formation of a soft callus of cartilage surrounding the fracture. Along the periphery of the callus bone is deposited. This initial deposit of bone is known as periosteal reaction (Dwek, 2010). The soft cartilage callus then ossifies into a primary bony callus with little organization. From this point the callus is reorganized and then remodeled until the unorganized woven bone that is initially laid down is replaced by lamellar bone which is characteristic of normal healthy bone (Roberts & Manchester,

2007).

Finally, in addition to fracture repair and bone development the periosteum has been a focus for research into progenitor cells, also known as stem cells. Interest in the periosteum for this purpose is due to the regenerative potential that the periosteum exhibits. The periosteum has been used to stimulate bone growth in clinical applications such as facial reconstruction (Chang, 2012). The presence of both mesenchymal and osteogenic progenitor cells in abundance makes the periosteum a good candidate for stem 17 cell research. The periosteum provides a less invasive source for harvesting the progenitor cells for regenerative research and therapies, as well as exhibiting a much higher proliferation rate compared to other options (Chang, 2012). The periosteum continues to be the subject of study to better understand the different roles it has in the body and the potential it has in many different medical applications.

Periosteal Reactions

As discussed, the periosteum is a dynamic membrane that stimulates new bone growth in different conditions. The formation of reactive bone is the first response that bone has to many different insults. The reactive bone formed is known as periosteal reaction. Periostitis in turn refers to the process that results in the reactive bone development. Periostitis may be stimulated by insults that affect many different areas of bone tissue including the cortical surface of bone, just beneath the periosteum

(subperiosteal), within the periosteum (periosteal), or even soft tissues that are directly adjacent to the periosteum (parosteal) (Bisseret et al., 2015). The variety of insults that stimulate periostitis are different in origin such as affecting a specific location, the bone or surrounding tissue, or as a response to a generalized process. The conditions that stimulate periosteal reactions may be benign, malignant, or even systematic in nature

(Bisseret, 2015). Periosteal reactions have many different appearances based on the intensity, duration, and aggressiveness of the insult (Rana et al., 2009). These factors affecting the appearance of periosteal reactions are indicative of the nature of the insult that stimulated them. Despite the cause the reaction often starts with inflammation of the periosteum and results in new bone growth. 18

Periosteal reactions are placed into two main categories which are aggressive and non-aggressive (see Figure 1). In general, aggressive periosteal reactions occur in cases where new layers of bone are being put down very rapidly over a short period of time.

Figure 1. Subtypes of Periosteal Reactions.

A-D are non-aggressive reactions as follows: thin (A), solid (B), thick irregular (C), and septated (D). E-I are aggressive reactions as follows: laminated (onionskin) (E), perpendicular/hair-on-end (F), sunburst (G), disorganized (H), and Codman triangle (I) periosteal reactions. (From Rana et al., 2009)

These conditions tend to leave an uneven appearance and can lead to sunburst or perpendicular hair-on-end patterns where bone is laid down in spindles away from the bone (Rana, 2009). In comparison, non-aggressive reactions tend to occur with conditions where bone is deposited very slowly over a longer duration of time. The appearance tends to be much smoother and layers can often be seen (Rana, 2009). Even with these differences in appearance it may not be possible to distinguish the nature of 19 the cause of the condition as overlap is seen in the diseases and other stimuli that lead to each of the two main types.

Within the two main categories many different subtypes have been classified.

These subtypes are characterized by the distinctive appearance of the periosteal reaction.

However, many of these subtypes are similar in physical appearance on bone and radiological imaging is required to be able to determine the exact subtype that is present.

As stated the non-aggressive reactions are characteristically smoother in appearance. The bone development is typically more organized in comparison to aggressive periosteal reactions. The subtypes that fall under this category are thin, solid, thick irregular, and septated (Rana, 2009). Both thin and solid are smooth deposits of reactive bone the difference being the thickness of the reactive bone. Thick irregular periosteal reactions are basically as the name describes, thick deposits of reactive bone with an irregular surface. Finally, the last subtype in the non-aggressive category is septated. Septated reactions are similar in appearance to solid reactions on the outer surface; however, beneath the surface there are pockets in the reactive bone. It is thought that nonaggressive reactions are the result of uninterrupted bone formation.

In comparison aggressive reactions often indicate intermittent growth. The subtypes that fall under aggressive reactions are laminated (onion skin), perpendicular

(hair-on-end), sunburst, disorganized, and Codman’s triangle (Rana, 2009). With laminated reactions there are many different sheets of bone growth leaving a layered appearance. There are two subtypes that are considered spiculated. Spiculated reactions are very aggressive and lead to projections of bone rather than a smooth surface. The two spiculated subtypes are hair-on-end and sunburst. The difference between the two is the 20 orientation of the projections. In hair-on-end the projections form perpendicular to the cortical surface; with sunburst the projections radiate in all directions from the cortical surface. Disorganized reactions are again as the name would suggest, disorganized reactive bone that is not in any specified pattern. Lastly, there is the Codman’s triangle.

This subtype is different from the others in that it is not characterized by bone growth.

Codman’s triangles are formed when periosteum is lifted away from the bone surface by different causes. Soft tissue must be present for the Codman’s triangle to be observed and as such is more of a radiological indicator of periosteal reaction than a subtype on its own. Once the periosteal reaction has been identified and characterized differential diagnosis may be done to give potential causes.

The process of differential diagnosis proves difficult when dealing with periosteal reactions. Though the reaction may be distinctive in appearance the different subtypes are not always indicative of a specific cause let alone a specific disease. In addition, it is not possible currently to determine whether the cause behind a periosteal reaction is due to a benign or malignant condition through radiological means (Rana, 2009). Despite these drawbacks there are some conditions that are associated with a specific subtype within a clinical setting. Among these conditions are fractures which generally exhibit a solid appearance, and sunburst patterns of growth are linked to conventional osteosarcomas, malignant cancers of bone (Rana, 2009). In a clinical setting there is also potential to use periosteal reaction to diagnose specific conditions or diseases, but this diagnosis is reliant on additional information such as other symptoms and medical records; both of which are not available within an archaeological setting.

21

The periosteum is a complex tissue that is not fully understood. It is key in the development and maintenance of bone tissue. New bone formation, also known as periosteal reactions are often the first response of bone to an insult. As it is the first response there are many different causes to the condition. With an understanding of the anatomy of the periosteum and the process behind periosteal reactions, analysis of periostitis within the archaeological record may provide better and more nuanced information as to stress in the past.

22

Periostitis within Bioarchaeology

The analysis of periostitis is common throughout bioarchaeological literature. Analysis of periostitis becomes essential especially when studying disease in the past as the condition is a part of many different disease processes. Analysis of the intensity and location of periostitis proves useful in differential diagnosis of different conditions (Gladykowska-Rzeczycka, 1998). In addition to the study of specific diseases, periostitis has been used to explore a variety of research topics, ranging from migration patterns to overall health and survival in the past.

Periosteal reactions are a useful skeletal indicator for the study of stress and disease due to its relative abundance in the archaeological record (Weston, 2008). The presence of periostitis has come to be an indicator of non-specific infection in many studies as the condition is the initial response of bone to insult, whether the insult is a result of disease, trauma, or otherwise (Larsen, 1999). The nature of the insult may be determined through examination of the periosteal reaction.

In general, if periosteal reaction is widespread across many different skeletal elements it is considered systemic in nature. In contrast, if the reaction is observed in one specific area it is interpreted as being the result of a more focal or local stressor. In addition to the distribution of periosteal reactions the level of healing and remodeling gives insight into the process that may have caused the new bone formation. Periosteal reaction may be determined to be active or inactive based on the level of remodeling present (Marx, 2008). Periosteal reactions are not normally differentiated by active or nonactive in the literature as the focus has been the presence or absence of the condition, or the use of periosteal reactions in differential diagnosis. 23

Within bioarchaeology, periostitis has primarily been explored in two different manners. The first is with individual case studies. Though interesting, these studies give limited information as to conditions in the past. Despite this limitation, they may prove beneficial as a look at differential diagnosis. Individual case studies allow for a closer look at individual pathological conditions. An example of such a study is done by

Christensen et al. (2013). This particular study centers around a skeleton from South- west Hungary with skeletal indicators that could be indicative of several different diseases. By focusing on a single specimen, the processes of a condition may be better understood. In addition, it is in these case studies that differential diagnosis is explored.

This information may then be applied in later studies.

The second manner is studying prevalence in populations. Population studies look at a single condition and use statistical analysis in order to see overall patterns in populations. This approach allows for larger questions to be explored using the patterns in condition prevalence. The types of questions that may be answered vary greatly. One such question is migration patterns in the New world as assessed by Rothschild and

Rogers (2009). This study used prevalence patterns of periosteal reaction skeletal distribution in order to look at the potential migration routes from the Old world to the

New world. The data from this study raise questions about traditional thoughts of human migration to the New World as climatic restrictions of one of the forms of periosteal reaction observed goes against the migration routes as they were understood. Questions such as those on migration patterns are only a portion of questions that may be answered and is perhaps a less common line of questioning. More often studies using periostitis as an indicator of stress use it to explore questions of health. DeWitte (2014) explores this 24 idea in post-Black Death London. By exploring age patterns of periosteal reaction in post and pre-Black Death burials the effects of epidemic disease on population health were considered. This form of study gives understanding both on past conditions as well as giving insight to potential consequences of an epidemic for better planning and understanding for the future. There are many more questions that may be explored by looking at periosteal reaction and so a better understanding of the condition is key.

Working toward a better understanding of periostitis, a study by Weston (2008) looks at periostitis on its own by exploring the use of macroscopic features of the reaction and the potential specificity of periosteal reactions. To determine whether different diseases or processes left distinct periosteal reactions pathology museum specimens from two separate museums were used as they represent old bone samples with known diseases affecting the bone.

The specimens underwent both macroscopic and radiographic analysis of the periosteal reaction present. Once analysis was completed the results were divided by the disease categories described by the museum. The goal was to determine traits of the reaction that could be an indicator of specific conditions with older specimens. In the end it was concluded that it was not possible to link qualitative or quantitative traits to specific disease processes in older bone samples.

In the past, periostitis has mostly been used as an indicator of non-specific infection or as a method of differential diagnosis of different conditions. As such periosteal reactions are a key part of many paleopathological studies. In addition, the study of the prevalence of periostitis within populations has potential to give a better insight into past conditions. As periostitis becomes better understood it is applied to 25 research in new ways providing a more nuanced view of the past. How analysis of periostitis has been applied to research in the past influences how it may be used in the future.

26

Background and Methods

California is an environmentally diverse area that has seen human occupation for thousands of years. For this work, central California region is the focus of analysis. In this context central California is both a geographic and archaeologically determined area.

The archaeological record of the area is extensive with over 300 sites being excavated from the 1880s to present day (Schwitalla et al., 2014a). The continued occupation of the area through different environmental and cultural developments, as well as the extensive archaeological record, makes central California an ideal area for study of stress and disease.

Bio-Cultural Background of California Sample

The central California region was home to a number of different cultural groups.

For most of the prehistory of the area, organization is described as many small tribelets

(Jones & Klar, 2007). The people of the area had a hunter-gatherer lifestyle with some groups being more sedentary or more mobile than others. There is a great amount of evidence that suggests violence in the area throughout prehistory. Evidence of violence includes dismemberment and trophy taking (Schwitalla et al, 2014b), craniofacial fractures (Pilloud, et al., 2014), and sharp force/projectile trauma, trauma appearing on the bone in a narrow area from an outside material, such as a knife or blade (Schwitalla et al., 2014a).

The sample is divided into six different general time periods based on the Central

California Taxonomic System (CCTS) (Pilloud, et al., 2014). The first period of time stretches from about 3500 B.C. to 500 B.C. and is classified as the Early period. Just 27 before this time central California had a shift in climate leading to overall warmer and drier conditions. The warmer conditions led to new marshland development which could potentially support larger populations as well as the desiccation of lakes and rivers in the central valley area (Jones, 2007). This time period is followed by the Early Middle from

500 B.C. to A.D. 420, and the Late Middle from A.D. 420-A.D. 1010. The next time period, A.D. 1010-1390, is called the Middle-Late Transition. This time period corresponds with a significant change of climate known as the Medieval Climatic

Anomaly. During the Medieval Climatic Anomaly an increase in temperature has been recorded in many parts of the world leading to long lasting droughts (Pilloud, 2006). In addition to climatic change, this period also sees the introduction of the bow and arrow technology into the area. The period following the Medieval Climatic Anomaly is referred to as the Late Prehistoric. This period of time spans from roughly A.D. 1390 to

A.D. 1720. It is during this time period that Europeans first arrive in the area and

Spanish Mission spread through the area (Jones, 2007). The final time period is from

A.D. 1720 through A.D. 1899 and is designated as the Protohistoric/Historic. This time period is marked by European colonization of the area.

Data Collection Procedures

The analysis of periostitis by demographic groups for each period of time was done in order to assess the use of periostitis as an indicator of stress. For the analysis, information from the central California bioarchaeological database (CCBD) was used.

28

Stage 1 The CCBD has been the ongoing work of researcher Al Schwitalla and contains information on 16,820 individuals found from 329 archaeological sites across central

California (Schwitalla et al, 2014). The sites are spread across 30 different modern counties in California. The distribution of the sites is shown in Figure 1. The information contained within the database was compiled from many different sources including, published site reports, unpublished site reports, burial records, osteological appendices, and NAGPRA (Native American Graves and Repatriation Act) inventories. In many cases, the listed sources of information are the only information available for the burials as the individuals are most commonly re-buried according to the wishes of tribal descendants per state and federal law (Schwitalla et al, 2014). Important to note is that the database and the studies that are included in the database were not compiled for the purpose of this work and have been the focus of studies by other researchers in the past.

Figure 2.Central California Sites Included in CCBD (From Pilloud et al., 2014) 29

The CCBD contains a wide variety of information about the archaeological sites and the individuals found. For each burial the site designation, year of excavation, burial number, tribal association, temporal period, and demographic information was recorded.

In addition, information on a number of stress indicators was included. The stress indicators compiled include: Harris lines, enamel hypoplasia, porotic hyperostosis, cribra orbitalia, dental caries, osteomyelitis, periostitis, fractures, violence, sinusitis, ear exostosis, degenerative joint disease, osteoarthritis, meningitis, treponematosis, cancerous tumors, and mass burials (Schwitalla, 2013). When available additional information on disease was provided such as element affected, the side of the body, and additional contextual comments. Since many of the original records used to compile the database had incomplete recording there are many instances where information is absent for one or more categories of information for a burial. When information was not present the category was labelled as not reported by the author and/or excavator (NRBAE)

(Schwitalla, 2013). As the focus of this work is periostitis only burials with recorded presence or absence of disease were included for study. Access to the CCDB was given to Dr. Pilloud who then compiled the burials with information on periostitis into a spreadsheet that was then provided for this work. Only burials that had the presence or absence of periostitis recorded in the data base was included in the spreadsheet.

Stage 2 A total of 7,478 burials had information on analysis for disease. I organized the burials by temporal period for statistical analysis. Statistical analysis was done for each 30 temporal period individually for presence of periostitis by age groups and sex. In addition, statistical analysis was done on the data set as a whole for overall trends over time. All statistical analysis was performed using SSPS Statistics software version 25. In the software Chi square tests were done to determine whether differences observed between different demographic groups were statistically significant at an alpha level of

0.05.

31

Results and Analysis

This chapter is divided into two main sections. The first is analysis of temporal changes in the prevalence of periostitis across the entire sample. In addition to the temporal changes the overall trends for presence of periostitis among different demographic groups in the sample are analyzed. Following examination of overall trends in the sample each time period is analyzed individually.

Full Sample and Temporal Analysis

Among the sample of 7,478 burials from the database, 1,001 individuals had periostitis present per the database. The distribution of individuals by temporal period is shown in Table 1. The result of Chi Squared analysis was a value of 24.302 with a p- value of 0.000. This result indicates that there is statistical significance in the difference in prevalence per time period and that the results are not just due to random chance.

Table 1. Table of Prevalence by Temporal Period.

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Temporal Period Count Percentage 3050-500 B.C. 106/690 15.4 500 B.C.- A.D. 420 215/1584 13.6 A.D. 420-1010 315/2025 15.6 A.D. 1010-1390 222/1797 12.4 A.D. 1390-1720 107/977 11 A.D. 1720-1899 36/405 8.9

32

50 45 40 35 30 25 Percentage of Individuals with 20 Periostitis 15 Present 10 5 0 3050-500 500 B.C.- A.D. 420- A.D. 1010- A.D. 1390- A.D. 1720- B.C. A.D. 420 1010 1390 1720 1899 Temporal Period

Figure 3. Prevalence of Periostitis per Temporal Period by Percentage There is an overall decrease in prevalence through time with a peak during A.D. 420-

1010 (See Figure 3). The prevalence difference between sexes was statistically significant with a chi squared value of 4.471 and a p-value of 0.034 showing a relationship between sex and prevalence of periostitis. Though the prevalence is similar between sexes (See Figure 4). Females had 15.5% prevalence of periostitis while males had 17.7% prevalence.

50 45 40 35 30 Percentage of 25 Individuals 20 with Periostitis 15 Present 10 5 0 Female Male Sex

Figure 4. Total Percentage of Males and Females with Periostitis Present 33

Table 2. Total Prevalence by Sex.

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Sex Count Percentage

Female 394/2547 15.5 Male 452/2558 17.7

Table 2 shows the exact counts for each sex as well as the percentage with periostitis.

Finally, the chi squared value between age groups was found to be 114.258 with a p- value of 0.000, indicating a statistically significant difference between age groups. The count per age group is outlined in Table 3. Figure 5 shows that overall prevalence increased with age with a peak in Juveniles and in Old Adults. In addition, across all time periods periostitis was absent in neonates.

50 45 40 35 30 Percentage of 25 Individuals with 20 Periostitis 15 Present 10 16.7 15.4 18.3 11.5 13.3 5 0 5.7 4.6 0

Age Category

Figure 5. Total Percentage of Individuals with Periostitis Present Per Age

34

Table 3. Total Prevalence by Age

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Age Count Percentage Neonate 0/58 0 Infant 25/439 5.7 Child 35/756 4.6 Juvenile 3/18 16.7 Adolescent 41/355 11.5 Young Adult 143/1073 13.3 Adult 582/3782 15.4 Old Adult 171/935 18.3

Early Period (3050-500 B.C.)

Within the Early period there was a 15.4% prevalence of periostitis with 106 individuals out of 690 being affected. Differences between sexes are shown in Figure 6, with the chi squared value being 1.406 and the p-value 0.236.

50 45 40 35 30 Percentage of 25 Individuals 20 with 15 Periostitis 10 Present 5 0 Female Male Sex

Figure 6. Percentage of Males and Females with Periostitis from 3050-500 B.C. 35

Table 4. Prevalence by Sex from 3050-500 B.C.

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Sex Count Percentage

Female 33/239 13.8

Male 48/272 17.6

There is a greater difference between sexes than seen in all temporal periods together with females having 13.8% prevalence and males 17.6% (See Table 4); however, the difference is not statistically significant. The distribution of periostitis increases with age during this period as seen in Figure 7.

120

100

80

Percentage of 60 Individuals with 40 Periostitis Present 20

0

Age Categories

Figure 7. Percentage of Periostitis by Age from 3050-500 B.C.

36

Table 5. Prevalence by Age from 3050-500 B.C.

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Age Count Percentage

Neonate 0/3 0

Infant 0/8 0

Child 1/47 2.1

Juvenile 1/1 100

Adolescent 4/33 12.1

Young Adult 11/111 9.9

Adult 49/325 15.1

Old Adult 40/159 25.2

Something of note is the 100% prevalence in juveniles is due to only a single individual of this age being in the sample (See Table 5). The chi squared value by age was 28.291 with a p-value of 0.000, making the results statistically significant. Though this result may be skewed by the sample size of some age groups, such as juveniles.

Early Middle Period (500 B.C. – A.D. 420)

The Early Middle Period had a prevalence of periostitis at 13.6% with 215 out of

1584 individuals having the condition. This period of time saw an increase in prevalence with age as seen in Figure 8. The chi squared value for the temporal period by age was

51.299 with a p-value of 0.000. 37

50 45 40 35 30 25 Percentage of 20 Individuals with Periostitis 15 Present 10 5 0

Age Category

Figure 8. Percentages of Periostitis by Age from 500 B.C. - A.D. 420

Table 6. Prevalence by Age from 500 B.C. - A.D. 420

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Age Count Percentage Neonate 0/8 0 Infant 2/114 1.8 Child 6/182 3.3 Juvenile 0/2 0 Adolescent 11/96 11.5 Young Adult 22/209 10.5 Adult 138/784 17.6 Old Adult 36/175 20.6

Again, something of note is the small sample size for juveniles (see Table 6). The distribution of periostitis by sex was similar to those seen in previous samples with 38 females at 17.3% and males at 17.6% (See Table 7). The prevalence was closer during this period than in previous ones, as shown in Figure 9 with the chi squared value being

0.017 with a p-value of 0.897. This high p-value indicates no relationship between prevalence of periostitis and sex.

50 45 Percentage of 40 Individuals 35 30 with 25 Periostitis 20 Present 15 10 5 0 Female Male Sex

Figure 9. Percentage of Periostitis by Sex from 500 B.C. - A.D. 420

Table 7. Prevalence by Sex from 500 B.C. - A.D. 420

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Sex Count Percentage

Female 84/486 17.3

Male 95/540 17.6

Late Middle Period (A.D. 420-1010)

The Late Middle Period saw a peak in prevalence of periostitis at 15.6%, with 315 of 2025 individuals being affected. When analyzed by sex a chi squared value of 1.459 with a p-value of 0.227 was found. Females had a prevalence of 18.0% and males a 39 prevalence of 20.5% (See Table 8). Once again there was little difference between sexes with males having a slightly higher prevalence (Figure 10).

50 45 40 35 30 Percentage of 25 Individuals 20 with Periostitis 15 Present 10 5 0 Female Male Sex

Figure 10. Percentage of Periostitis by Sex from A.D. 420-1010

Table 8. Prevalence by Sex from A.D. 420-1010

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Sex Count Percentage Female 131/729 18 Male 145/708 20.5

This temporal period saw the same increase of prevalence with age as seen before (See

Figure 11). The chi squared value was 28.475 with a p-value of 0.000, making the results statistically significant (See Table 9). 40

50 45 40 35 30 25 Percentage of 20 15 Individuals 10 with Periostitis 5 Present 0

Age Category

Figure 11. Percentage of Periostitis by Age from A.D. 420-1010

Table 9. Prevalence by Age from A.D. 420-1010

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Age Count Percentage Neonate 0/9 0 Infant 2/75 2.7 Child 14/205 6.8 Juvenile 0/4 0 Adolescent 17/100 17 Young Adult 51/297 17.2 Adult 188/1094 17.2 Old Adult 42/222 18.9

Middle-Late Transition Period (A.D. 1010-1390)

The prevalence of periostitis decreased from that seen in the Late Middle Period with 12.4%, 222 of 1797 individuals, affected. In comparison to the period before prevalence between sexes saw a greater difference with 13.5% of females, and 17.5% of 41 males having periostitis present (See Figure 12).

50 45 40 Percentage of 35 Individuals 30 with 25 Periostitis 20 Present 15 10 5 0 Female Male Sex

Figure 12. Percentage of Periostitis by Sex from A.D. 1010-1390

Table 10. Prevalence by Sex from A.D. 1010-1390

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Sex Count Percentage

Female 85/630 13.5

Male 105/600 17.5

The chi squared value found was 3.780 with a p-value of 0.052. A p-value of 0.052 indicates no relationship between sex and periostitis exist during this temporal period

(See Table 10). Analysis of age indicated similar results as other time periods with prevalence increasing with age (See Figure 13). The chi squared value for age was

18.512 with a p-value of 0.018, indicating a relationship between periostitis and age during the time period (See Table 11). 42

50 45 40 35 30 Percentage of Individuals with 25 Periostitis 20 Present 15 10 5 0

Age Category

Figure 13. Percentage of Periostitis by Age from A.D. 1010-1390

Table 11. Prevalence by Age from A.D. 1010-1390

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Age Count Percentage Neonate 0/9 0 Infant 16/121 13.2 Child 7/170 4.1 Juvenile 0/6 0 Adolescent 5/73 6.8 Young Adult 32/262 12.2 Adult 130/921 14.1 Old Adult 32/220 14.5

Late Prehistoric (A.D. 1390-1720)

The Late Prehistoric period saw a continued decrease in the prevalence of 43 periostitis at 11% prevalence with 107 of 977 individuals having periostitis present.

Analysis of prevalence by age showed a general trend of prevalence increasing with age, though a peak is seen in Juveniles (See Figure 14).

50 45 40 35 Percentage of 30 Individuals 25 with Periostitis 20 Present 15 10 5 0

Age Category

Figure 14. Percentage of Periostitis by Age from A.D. 1390-1720

Table 12. Prevalence by Age from A.D. 1390-1720

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Age Count Percentage

Neonate 0/14 0

Infant 3/77 3.9

Child 4/89 4.5

Juvenile 1/3 33.3

Adolescent 4/45 8.9

Young Adult 14/118 11.9

Adult 66/508 13

Old Adult 15/114 13.2 44

The chi squared value found was 13.910 with a p-value of 0.053. This p-value puts the results (See Table 12) just out of the 95% confidence level and so no relationship between age and prevalence is seen during the temporal period. With sex the chi squared value found was 2.233 with a p-value of 0.135. Males had the higher prevalence at

16.3% while females had a prevalence of 12.2% (See Figure 15 and Table 13).

50 45 40 Percentage of 35 Individuals 30 with 25 Periostitis 20 Present 15 10 5 0 Female Male Sex

Figure 15. Percentage of Periostitis by Sex from A.D. 1390-1720

Table 13. Prevalence by Sex from A.D. 1390-1720

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Sex Count Percentage

Female 41/336 12.2 Male 53/326 16.3

Protohistoric/Historic (A.D. 1720-1899)

During the Protohistoric/Historic period a prevalence of 8.9% is observed with 36 of 405 individuals having periostitis present. During this period a reverse of previous 45 trends is seen with prevalence by sex (See Table 14). Females show a greater prevalence of periostitis at 15.7% while males are at only 5.4% prevalence, making for a greater difference by sex (See Figure 16).

50 45 40 35 Percentage of 30 Individuals 25 with Periostitis Present 20 15 10 5 0 Female Male Sex

Figure 16. Percentage of Periostitis by Sex from A.D. 1720-1899

Table 14. Prevalence by Sex from A.D. 1720-1899

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Sex Count Percentage

Female 20/127 15.7

Male 6/112 5.4

The chi squared value for sex was 6.628 with a p-value of 0.01 indicating a relationship between sex and periostitis during this time period. Prevalence by age also saw a shift with periostitis being most prevalent in young adults (See Figure 17). The chi squared value was 16.573 with a p-value of 0.020. This temporal period sees a change in the 46 pattern of prevalence with young adults showing the most prevalence (See Table 15).

This is discarding the juveniles which are likely skewed by sample size again.

100 90 80 70 60 50 40 Percentage of 30 Individuals with 20 Periostitis Present 10 0

Age Category

Figure 17. Percentage of Periostitis by Age from A.D. 1720-1899

Table 15. Prevalence by Age from A.D. 1720-1899

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Age Count Percentage Neonate 0/15 0 Infant 2/44 4.5 Child 3/63 4.8 Juvenile 1/2 50 Adolescent 0/8 0 Young Adult 13/76 17.1 Adult 11/150 7.3 Old Adult 6/45 13.3

47

Overall Trends

When comparing percentages of periostitis by age over time the majority of the different age groups see little change between temporal periods (Figure 18).

120

100

80 Neonate 60 Infant Child Percentage of 40 Juvenile Individuals with Adolescent Periostitis Present 20 Young Adult Adult

0 Old Adult

Temporal Period

Figure 18. Change in Prevalence of Periostitis by Age over Time.

The exception to this trend is seen in the juvenile age group. The large changes in this particular category are probably due to the small sample sizes of this age group throughout the sample. The prevalence of periostitis by sex had some variation over time

(Figure 19). The prevalence in females stays fairly consistent through time while the prevalence in males sees an increase peaking in A.D. 420-1010, followed by continued decreases after this period. 48

50 45 40 35 30 Percentage of 25 Individuals with 20 Periostitis Present 15 10 5 0 3050-500 500 B.C.- A.D. 420- A.D. 1010- A.D. 1390- A.D. 1720- B.C. A.D. 420 1010 1390 1720 1899 Temporal Period

Female Male

Figure 19. Prevalence of Periostitis Over Time by Sex

Over the span of time studied prevalence of periostitis decreased in central

California. Between time periods there was little change in the prevalence by age and by sex. Differences in prevalence by sex were statistically insignificant and were likely due to sample size or random chance. A significant difference was seen in sex during the final time period A.D. 1720-1899. In this time period females showed a higher prevalence than males which is the reverse of what is seen in every other temporal period. A relationship between prevalence of periostitis and age was seen with the prevalence increasing with age. This pattern was seen in all time periods except the final temporal period A.D. 1720-1899. The juvenile age category was likely skewed due to sample size which would account for the large changes in prevalence between temporal periods. 49

Conclusions

Periostitis as an indicator of stress and disease has seen a lot of use within the field of bioarchaeology. Most often periostitis is used as an indicator of unspecific infection or unspecific disease as the condition is observed with many different disease processes. Despite the wide use of analysis of periostitis within the field of bioarchaeology little research has been done into the condition itself. By looking to clinical research of the condition and the process behind the condition a better understanding of periostitis becomes possible, which in turn leads to better use of analysis of periostitis in the future. In addition to analysis of literature on periostitis a case study of the prevalence of periostitis in a sample from central California was done to assess the use of periostitis as an indicator of stress in the past.

Conclusions from Literature Review

The ability of the periosteum to instigate new bone growth has been known since the 19th century; however, there is still much about the process that is not well understood. There are structures within the periosteum that have not seen a lot of research, such as Sharpey’s fibers that help anchor the periosteum to the underlying bone

(Chang &Knothe Tate, 2012). In addition to uncertainties in the anatomy of the periosteum, the exact process to instigate new growth is not fully understood. The purpose of each of the different cell types and what is used to trigger these cells is one area for further research.

In addition, there are a great number of causes that have been documented for periosteal reactions including: trauma, infection, chronic diseases, and even certain drug 50 therapies. Clinical research into diagnosis of different conditions through radiographs of periostitis has seen some success. This is done mostly through recognition of patterning of periosteal reactions within the body (Chen et al., 2012). Even this remains difficult as different conditions may exhibit similar patterning of periostitis. Attempts by Weston

(2008) to identify characteristics of periosteal reactions that were specific to different diseases within museum specimens showed no discerning characteristics for different disease processes.

From analysis of both clinical and bioarchaeological literature of periostitis it is apparent that there is still much to learn about this seemingly simple process. The use of periostitis as an indication of unspecific infection within bioarchaeological research does not take into full consideration the nature of the condition. With so many different potential causes it is simply not possible to be sure that the cause was infection, especially as there is no way to differentiate periosteal reactions by cause at this time.

The study of stress within bioarchaeology is complex with new perspectives and developments being seen to this day. Skeletal indicators of stress are conditions that represent times of growth disruption such as linear enamel hypoplasia and indicators of disease. Periostitis is a common indicator of disease that is used in research, but there is debate on whether it should be considered an indicator of stress at all (Klaus, 2014).

Conclusions from Case Study

The analysis of periostitis within a sample from central California had some unexpected results. The peak in prevalence was earlier than expected, being in A.D. 420-

1010. With the Medieval Climatic Anomaly leading to long droughts in the area, an 51 increase in presence of periostitis was expected from A.D. 1010-1390. In addition, the continued decrease in prevalence going into the time of contact with Europeans was also of surprise. Perhaps these results are due to the osteological paradox, being that people were dying before any sign of stress or disease could manifest in the skeleton.

Looking at prevalence of periostitis between sexes showed that there was little difference between males and females. Both sexes had percentages close to each other repeatedly with no statistical difference between the two in most temporal periods. An interesting finding was the switch seen in the Protohistoric/Historic period where females exhibited a higher prevalence than males. This difference was also shown to be statistically significant with a p-value of 0.01. This time period is marked by conflict between Native American groups and Europeans coming to the area. The difference may be seen as men were in combat more frequently and so were dying from trauma before periostitis could develop. This, however, is mere speculation as there is no way to confirm this theory.

With analysis of age, a pattern was seen that repeated in each temporal period.

The pattern illustrated that the prevalence of periostitis increased with age. This increase is expected as older individuals would have more time for chronic conditions to affect the skeleton. In addition to a greater amount of time, there is a natural decrease in the strength of bone with age. With the decrease in strength, the bone is more susceptible to breakage and trauma, leading to more instances of periostitis. The prevalent changes observed in juveniles is likely due to the low sample sizes as this age group repeatedly had only a few individuals, in some temporal periods only one, which could skew the prevalence. 52

The prevalence patterns of periostitis gives some insight into conditions in the past within the area. The combination of the overall decrease in prevalence of periostitis over time and the increase in periostitis with age is very interesting. As time goes on it would be expected that prevalence of periostitis increases as people survive stressors and diseases long enough to have the condition develop, which is not what is observed in the sample from central California. The decrease in prevalence of periostitis that is observed in central California is most likely the result of the osteological paradox, that is that people are dying before any indicators of stress, such as periostitis can manifest on the skeleton. If this is the case, then the presence of periostitis would be an indication of adaptation. Those individuals that have periostitis represent the healthiest individuals as they survived long enough to develop a bony response.

Further research that may be done to improve this work would be to investigate the reason for the peak in prevalence of periostitis during the late middle period (A.D.

420-1010). The peak is unexpected in this temporal period as I would have expected it to occur from A.D. 1010-1390 when the medieval climatic anomaly occurred. Another instance that I would like to investigate is the switch that is seen in prevalence between sexes during the final temporal period A.D. 1720-1899. There is a significant relationship between prevalence in periostitis and sex during this period, but the cause is not known for sure. One final area of research that could prove interesting would be to investigate the state of periostitis observed, such as whether they were active or healed, in the individuals within the sample used for this work. This area of research is most likely not possible as the majority of the individuals from the sample were reburied or are otherwise inaccessible for further research. 53

Overall, though the trends of periostitis are interesting they do not give much information as to stress in past populations. With there being so many potential causes for the condition it is difficult to be sure exactly what the cause of the periosteal reaction was. In many cases periosteal reactions occur due to everyday occurrences. On its own analysis of periostitis just does not give enough information to obtain a clear picture of conditions in the past. If information was available as to whether the periosteal lesions were active or healing and their location a better conclusion and understanding of stress in the past of central California would be possible. More research is needed into the process of periosteal reactions so that the analysis of periosteal reactions as an indicator of stress can be more nuanced rather than just being able to say that something upset the periosteum, but no further information is known. 54

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60

Appendix A

Figures

Figure 1. Subtypes of Periosteal Reactions.

A-D are non-aggressive reactions as follows: thin (A), solid (B), thick irregular (C), and septated (D). E-I are aggressive reactions as follows: laminated (onionskin) (E), perpendicular/hair-on-end (F), sunburst (G), disorganized (H), and Codman triangle (I) periosteal reactions. (From Rana et al., 2009) 61

Figure 2. Central California Sites Included in CCBD (Pilloud et al., 2014) 62

50

45

40

35

30

Percentage of 25 Individuals with Periostitis Present 20

15

10

5

0 3050-500 B.C. 500 B.C.- A.D. A.D. 420-1010 A.D. 1010-1390 A.D. 1390-1720 A.D. 1720-1899 420 Temporal Period

Figure 3. Prevalence of Periostitis per Temporal Period by Percentage

63

50 45 40 35 30 Percentage of 25 Individuals with Periostitis Present 20 15 10 5 0 Female Male Sex

Figure 4. Total Percentage of Males and Females with Periostitis Present.

50 45 40 35 30 Percentage of 25 Individuals with 20 Periostitis Present 15 10 16.7 15.4 18.3 5 11.5 13.3 0 5.7 4.6 0

Age Category

Figure 5. Total Percentage of Individuals with Periostitis Present Per Age 64

50 45 40 35 30 Percentage of Individuals with 25 Periostitis Present 20 15 10 5 0 Female Male Sex

Figure 6. Percentage of Males and Females with Periostitis from 3050-500 B.C.

120

100

80

Percentage of 60 Individuals with Periostitis 40 Present 20

0

Age Categories

Figure 7. Percentage of Periostitis by Age from 3050-500 B.C 65

50

45

40

35

30

Percentage of 25 Individuals with Periostitis 20 Present 15

10

5

0

Age Category

Figure 8. Percentages of Periostitis by Age from 500 B.C. - A.D. 420

50 45 40 Percentage of 35 Individuals with 30 Periostitis 25 Present 20 15 10 5 0 Female Male Sex

Figure 9. Percentage of Periostitis by Sex from 500 B.C. - A.D. 420 66

50 45 40 35 Percentage of 30 Individuals with 25 Periostitis 20 Present 15 10 5 0 Female Male Sex

Figure 10. Percentage of Periostitis by Sex from A.D. 420-1010

50 45 40 35 30 Percentage of 25 Individuals with Periostitis 20 Present 15 10 5 0

Age Category

Figure 11. Percentage of Periostitis by Age from A.D. 420-1010

67

50 45 40 35 Percentage of 30 Individuals with 25 Periostitis Present 20 15 10 5 0 Female Male Sex

Figure 12. Percentage of Periostitis by Sex from A.D. 1010-1390

50

45

40

35

Percentage of 30 Individuals with 25 Periostitis Present 20 15

10

5

0

Age Category

Figure 13. Percentage of Periostitis by Age from A.D. 1010-1390 68

50 45 40 35 30 Percentage of 25 Individuals with 20 Periostitis Present 15 10 5 0

Age Category

Figure 14. Percentage of Periostitis by Age from A.D. 1390-1720

50

45

40

35

30 Percentage of Individuals with 25 Periostitis Present 20

15

10

5

0 Female Male Sex

Figure 15. Percentage of Periostitis by Sex from A.D. 1390-1720 69

50 45 40 35

Percentage of 30 Individuals with 25 Periostitis Present 20 15 10 5 0 Female Male Sex

Figure 16. Percentage of Periostitis by Sex from A.D. 1720-1899

100 90 80 70 60 50

Percentage of 40 Individuals with 30 Periostitis Present 20 10 0

Age Category

Figure 17. Percentage of Periostitis by Age from A.D. 1720-1899 70

120

100

80 Neonate 60 Infant Child 40 Percentage of Juvenile Individuals with Adolescent Periostitis Present 20 Young Adult

0 Adult Old Adult

Temporal Period

Figure 18. Change in Prevalence of Periostitis by Age over Time.

50 45 40 35 30 Percentage of 25 Individuals with 20 Periostitis Present 15 10 5 0 3050-500 500 B.C.- A.D. 420- A.D. 1010- A.D. 1390- A.D. 1720- B.C. A.D. 420 1010 1390 1720 1899 Temporal Period

Female Male

Figure 19. Prevalence of Periostitis Over Time by Sex 71

Tables

Table 1. Table of Prevalence by Temporal Period.

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Temporal Period Count Percentage

3050-500 B.C. 106/690 15.4

500 B.C.- A.D. 420 215/1584 13.6

A.D. 420-1010 315/2025 15.6

A.D. 1010-1390 222/1797 12.4

A.D. 1390-1720 107/977 11

A.D. 1720-1899 36/405 8.9

72

Table 2. Total Prevalence by Sex.

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Sex Count Percentage

Female 394/2547 15.5

Male 452/2558 17.7

Table 3. Total Prevalence by Age

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Age Count Percentage

Neonate 0/58 0

Infant 25/439 5.7

Child 35/756 4.6

Juvenile 3/18 16.7

Adolescent 41/355 11.5

Young Adult 143/1073 13.3

Adult 582/3782 15.4

Old Adult 171/935 18.3

73

Table 4. Prevalence by Sex from 3050-500 B.C.

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Sex Count Percentage

Female 33/239 13.8

Male 48/272 17.6

Table 5. Prevalence by Age from 3050-500 B.C.

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Age Count Percentage

Neonate 0/3 0

Infant 0/8 0

Child 1/47 2.1

Juvenile 1/1 100

Adolescent 4/33 12.1

Young Adult 11/111 9.9

Adult 49/325 15.1

Old Adult 40/159 25.2

74

Table 6. Prevalence by Age from 500 B.C. - A.D. 420

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Age Count Percentage

Neonate 0/8 0

Infant 2/114 1.8

Child 6/182 3.3

Juvenile 0/2 0

Adolescent 11/96 11.5

Young Adult 22/209 10.5

Adult 138/784 17.6

Old Adult 36/175 20.6

Table 7. Prevalence by Sex from 500 B.C. - A.D. 420

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Sex Count Percentage

Female 84/486 17.3

Male 95/540 17.6

75

Table 8. Prevalence by Sex from A.D. 420-1010

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Sex Count Percentage

Female 131/729 18

Male 145/708 20.5

Table 9. Prevalence by Age from A.D. 420-1010

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Age Count Percentage

0 Neonate 0/9 2.7 Infant 2/75 6.8 Child 14/205 0 Juvenile 0/4 17 Adolescent 17/100 17.2 Young Adult 51/297 17.2 Adult 188/1094

18.9 Old Adult 42/222 76

Table 10. Prevalence by Sex from A.D. 1010-1390

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Sex Count Percentage

Female 85/630 13.5

Male 105/600 17.5

Table 11. Prevalence by Age from A.D. 1010-1390

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Age Count Percentage

Neonate 0/9 0

Infant 16/121 13.2

Child 7/170 4.1

Juvenile 0/6 0

Adolescent 5/73 6.8

Young Adult 32/262 12.2

Adult 130/921 14.1

Old Adult 32/220 14.5 77

Table 12. Prevalence by Age from A.D. 1390-1720

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Age Count Percentage

Neonate 0/14 0

Infant 3/77 3.9

Child 4/89 4.5

Juvenile 1/3 33.3

Adolescent 4/45 8.9

Young Adult 14/118 11.9

Adult 66/508 13

Old Adult 15/114 13.2

Table 13. Prevalence by Sex from A.D. 1390-1720

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Sex Count Percentage

Female 41/336 12.2

Male 53/326 16.3

78

Table 14. Prevalence by Sex from A.D. 1720-1899

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Sex Count Percentage

Female 20/127 15.7

Male 6/112 5.4

Table 15. Prevalence by Age from A.D. 1720-1899

*Count is the number of individuals with periostitis out of the total burials for the temporal period

Age Count Percentage

Neonate 0/15 0

Infant 2/44 4.5

Child 3/63 4.8

Juvenile 1/2 50

Adolescent 0/8 0

Young Adult 13/76 17.1

Adult 11/150 7.3

Old Adult 6/45 13.3