Bone Development and Niche Construction Theory

Catherine Roberts, Penn State University, Dept of Anthropology

Abstract

Niche Construction Theory provides the ideal framework for understanding

human development. Development occurs within human-constructed reality and is

guided by both biological and environmental processes. This paper explores the

complexities of bone development through explanations of epigenetics, plasticity, and

cell signaling and places it within cultural contexts. Development of the musculoskeletal

system exists in a feedback loop within biological and cultural coevolution on both the

molecular and macro scales. Evidence from instances of bone remodeling and

morphological changes in hominid evolution exemplify that the genetic processes of

bone development are inseparable from external factors. Actions and access to

resources ingrained in cultural practices influence the developing bone throughout a

persons lifetime. This connection between culture and biology has shaped the

morphology and variation of past communities and continues to drive the evolution of

the human species.

Introduction

The question of nature vs. nurture has been asked by researchers and the

mainstream public for generations. However, only recently have we been able to

observe the complexity of these processes and the ways in which they are intertwined.

Both environmental and genetic factors come together in developmental processes

such that changes in either will influence not only the phenotype of the individual but
also the evolution of their species. On the borders between these factors lie complex

processes, such as changes in plasticity and epigenetic influences. Another

complicating aspect of evolution is niche construction, the influence of organisms on

their environment and the resulting pushback. Niche construction theory places

organism development in an ecological context and explores the cycle of organisms and

environments interacting to shape one another.

The aim of this paper is to explore the processes of bone development within the

cycle of niche construction. As humans, we live in a designed world, shaped by social

interactions, technological development, and other cultural influences. Additionally,

developmental concepts, such as bipedal walking, are central to our identification as

humans. Combining social and biological networks through niche construction theory

creates a larger picture of human ecology and evolution. Through exploring the

dynamics of niche construction, both cultural and biological evolution, and human

development, it becomes clear that he niche construction framework provides the most

complete image of the complexities of bone morphology, development, and evolution

within environmental context.

Niche Construction

Nothing about culture makes sense except in the light of evolution (Richerson

and Boyd 2005). This is a play on the quote from Theodosius Dobzhansky in which he

remarks that evolution is the only way to make sense of the discoveries of biology.

Through this statement, Richerson and Boyd argue that the same is true for culture.

Evolutionary theory allows the dynamics of cultures to combine with biology to form a
broader, clearer picture, rather than just a collection of facts. Culture and genetics are

both subject to competition and selection. Every question asked about human culture

must be answered with both cultural and biological evidence (2005).

Barbara Rogoff opens her book, The Cultural Nature of Human Development by

stating, People develop as participants in cultural communities. Their development can

be understood only in light of the cultural practices and circumstances of their

communities, which also change (2003). She argues for a nonlinear understanding of

development. While earlier views focus on a specific goal of ideal human development,

models such as niche construction more accurately reflect the reality of evolution. Just

as Darwins theory involves evolution of species in response to changing environments,

without any progress necessarily of which to speak, culture also follows this trend of

continuous change without a specific set of practices to eventually attain.

Niche construction is the process by which organisms interact with and are

affected by their own and each others environments. This continuous feedback loop

propels ecology as well as the changes that occur over time (Laland and OBrian 2011).

Many discussions of organism evolution focus on natural selection. Natural selection is

the concept that the conditions of the environment shape the direction and result of

evolutionary processes. Nature selects the most adept individuals who then survive to

pass on their traits to the next generation. This, however, is only one of the many

processes that occur within an ecosystem to drive evolution. Niche construction theory

takes into account a species ability to transform their environment and the ways in

which these modifications go on to affect the species. With niche construction theory,

not only are humans shaped by their environment, but they are recognized to have the
ability to influence selective factors. Laland and OBrian (2011) describe niche

construction theory as a developmental process rather than merely a product,

emphasizing this theory as a conceptual framework within which researchers could

approach problems with a different mindset and explore concepts that are left out of the

usual conversation of evolution through natural selection.

Niche construction theory provides an illustrative framework for both the

biological and the social sciences. It blurs common assumptions about which factor, the

individual or the environment, is the instigator of evolutionary change. Just as biologists

focus on organism adaptations to the environment, social scientists focus on the

creation of society through human agancy. Niche construction does not necessarily

begin with genetic traits, nor does genetic variation necessarily stem from variations in

the environment. While niche construction is recognizably the product of learned

knowledge and cultural practices, this transmission of information dependent on existing

biological traits and processes (Laland and OBrian 2011).

For example, George A. Miller (1972) describes linguistic communication as a

combination of biological, psychological, and social processes. It is not as though the

complexities of brain processing developed first, followed later by language. Small

changes to the brain that aid with language processing and increases in complexity of

verbal communication in human society both evolved together over time, one

influencing the other, to develop the mechanisms we see in humans today. Additionally,

the vocal tracts of humans are more complex than their other modern primates,

indicating that these mechanisms, too, played a role in the networks of language

evolution. If we considered any of these factors on their own, we would be confounded

by the adaptation of language-processing brains that have no language to speak, for

example, or the development of complex pharynges with no ability to process the

dynamics of language.

Central to niche construction theory is the coevolution of genes and cultures. Just

as genes are selected for based on specific environmental and lifestyle conditions, so

too are cultures shaped by the traits of their constituents. Cultural traits and practices

modify selection factors and shape other aspects of evolutionary mechanics (Feldman

and Laland 1996). Like genes, cultural traits are passed on from one generation to the

next. If particular practices continue to be common in a society, genes that are

detrimental in cultural context will disappear and advantageous genes will increase in

prevalence. Additionally, the genotype of a society has the ability to shape the

inheritance and evolution of culture. Common examples include agricultural methods,

as related to energy and digestion, and tool use, as related to adaptations in anatomy

and physiology. For example, accessibility to milk favors the ability to digest lactose into

adulthood. The resulting trait of lactase persistence allows for the addition of milk as an

important aspect of ones diet. The practice of animal domestication (through which milk

can easily be attained) is passed on between generations, thereby increasing selection

for lactose digestion (Feldman and Laland 1996). Similarly, in the aforementioned

language example, societal and biological traits coevolved to form language. An

increased need to communicate favors improved language processing, the devotion of

brain function to language encourages the inheritance of linguistic communication

methods within a society and so on (Miller 1972).

 Humans interact with the world through their behavior. Behavior is the means

through which personal autonomy can be achieved, allowing humans to interact with

society. Behavior also functions as a response mechanism, which is affected by

environmental stimuli, and a means of passing on cultural practices through learned

response. Within the framework of niche construction theory, the evolution of behavior

can be explained. Niche construction models allow for both the biological and social

aspects of behavior to be inherited together and coevolve through the relationship

between humans and ecological contexts (Laland Kendal and Brown 2007).

Genetics and environment intersect at the process of development. Cells

respond to signals from one another as well as external stimuli. Developmental biology

primarily focuses on cell differentiation and patterning to bring about particular

phenotypes from an individuals genotype. However, this is only part of the picture. The

field of ecological developmental biology focuses on the external factors involved with

organism development. Many environmental factors present during development

influence the actions of the cells. Examples include nutrient deficiencies, temperature,

and electromagnetic radiation. Additionally, epigenetic factors, especially the presence

of signaling proteins, modify developmental patterning (Gilbert and Epel 2009).

Plasticity and Epigenetics

Selection for advantageous traits tends to be even more complex than a two-way

feedback loop. Plasticity is the ability to change over time as the result of modifying

effects. From the creation of gametes by their parents and throughout an individuals

lifetime, one is subject to influence by both genetic and environmental factors (Sinha
2006). Epigenetics involves the processes through which genes are expressed. Rather

than gene sequence changes being inherited, the genes can remain unaltered while

changes to their expression patterns occur. For example, patterns of hormone presence

control the development of sexual characteristics. The activity of the hormone

aromatase during embryo development is responsible for sex determination in turtles

and other reptiles through its ability to convert testosterone to estradiol. High

temperatures increase aromatase activity, reducing the testosterone concentration while

increasing the estrogen concentration. Low amounts of testosterone cause ovaries to

form. Turtle eggs exposed to low temperatures will have high testosterone levels, and

the embryos will develop testes. Therefore, the sex ratio of a clutch is determined by the

temperatures to which each embryo was exposed during the critical period of sex

determination.

Not only is plasticity relevant on the cellular level, but the trait of plasticity itself is

also under the influence of selection. Rather than selecting for traits that are ideal for

one environment or another, organism evolution can move in the direction of an

increase in an individuals ability to adapt to an ever-changing world. In the aromatase

example, the aromatase gene was selected for and became fixed in the turtle lineage,

allowing for increased regulation of sex during development. Selection for plasticity is an

essential component of evolutionary change (Gilbert and Epel 2009). Through niche

construction, organisms can modify their environment to suit their current needs rather

than always having to change genetically, a much slower route, every time

environmental changes occur. Because of the convenience of this method and its
contribution to the survival of a species, plasticity is usually a highly adaptive trait

(Laland and Brown 2006).

Humans have developed high plasticity in many aspects, especially in the brain

and musculoskeletal system. Studies in visual perception reveal that the manufactured

world humans are born into shape the way their brains process visual information. The

interaction of the brain with the environment through visual perception functions in a

feedback loop similar to the cycle of niche construction theory (Gibson 1950).

Perception relies on both environmental stimuli and brain function in working

conjunction. Neither is the designated cause, paired with the other as an effect.

Activity of neuron and connections between them are altered by behaviors and cognitive

response to changing stimuli. The plasticity of neurons allow humans to improve their

perception and ultimately their behavior in response to the characteristics of the

environment in which they live (Stiles 2000).

Many epigenetic effects are studied through behavior and brain processing.

Cross-cultural psychological studies in response to various stimuli and human bonding

are conducted to observe the differences in biological function between individuals as

the result of cultural influence. Mathematical models of ethnographic curves and

epigenetic traits show that ethnographic patterns are highly influenced by epigenetic

actions (Lumsden and Wilson 1980).

Bone Development

Bone formation begins with the creation of a cartilage matrix, which is then

replaced with bone. From centers of ossification either the cartilaginous matrix is directly
ossified or growth plates form, allowing for continued cell formation, differentiation, and

calcification. Three types of cells are involved in both the formation and remodeling of

bone: osteoblasts, osteoclasts, and osteocytes. Osteoblasts are responsible for the

formation of new bone while osteoclasts resorb existing bone. Osteocytes exist within

the bone matrix and synthesize type I collagen and several proteins and deposit

minerals. The processes of forming, maintaining, and removing bone continue

throughout an organisms lifetime, though they are most active during initial

development from embryo to adult (Hall 2005).

Cell activity is controlled through patterns of signaling between cells. Groups of

cells receive and transmit signals to nearby cells. These signals control the proliferation,

differentiation, organization, etc. of cells, through regulating the expression of

transcription factors. There are not a large number of such signaling pathways, but the

patterns of their transmission through or between cells during development are

responsible for the differences in morphology observed between species. Morphogen

gradients establish spacial planes which provide information as to how particular cells,

based on their relative location, ought to behave (Perrimon, Pitsouli, and Shilo 2012).

Development of the musculoskeletal system occurs in this way, allowing for unique

morphologies to evolve in different vertebrate lineages. The cells of long bones are

aware of their positions and can therefore differentiate to form different structures. In

humans, some of the ends of long bones develop secondary centers from which bone is

ossified. Between the original, primary center and secondary center, a growth plate is

formed, allowing for the bone to continue to grow in length. If the signal patterns or ways
in which cells respond to particular signals are altered in any way, the development of

the organism will be affected.

The appearance of secondary centers of ossification may be an example of

epigenetic effects on patterning. While all vertebrates develop bone, only some species,

such as mammals and reptiles, develop growth plates during long bone formation.

However, it is likely that birds have the genetic ability to form secondary centers but that

their cells are not exposed to the same constraints mammalian cells experience during

development. Difference in rate and rate change over time of perichondrial and

endochondrial ossification put stress on the dividing cells. Joint stress by muscles and

hydrostatic forces result in the formation pattern of the chondroepiphysis seen in

mammals, in which a secondary center ossifies before cells in the growth plate

formation region. The cells in bird long bone are not subject to these forces and bone

formation continues to occur, directly calcifying nearby cartilage, outwards from the

primary center. (Carter, Mikic, and Padeon 1998).

Environmental Influences

Bone development does not stop once an individual reaches adulthood;

osteoblasts, osteoclasts, and osteocytes continue act on the bone, remodeling it

throughout an individuals lifetime. Throughout a persons lifetime, their bones are

constantly remodeling and responding to various stresses placed upon them. The use

of models allows researchers to replicate characteristics of bone and observe the

effects of varying degrees of mechanical stress. Both overloading and disuse cause

changes to osteoblast and osteoclast activity, the cells driving bone remodeling
(Hazelwood, et al. 2001). Oftentimes, what we would consider functional, ordinary

human bone morphology cannot come about without external forces guiding

development. Femur shaft angle and other features are a crucial aspect of bipedal

movement. Fetal activity and muscle contractions in the womb and into childhood give

the human femur its characteristic shape. Computer models were able to predict the

effect patterns of reduced loading would have on femur shaft radius and relative

endosteum and periosteum thickness (Carter, Meulin, and Beaupre 1996).

In a study which involved the removal of sheep ulnae, researchers observed the

remodeling efforts of the radius in response to increased strain. The new bone

deposited was not random, but occurred mainly on the caudal cortex, suggesting that

bone growth is not a simple response to increased stress, but a deliberate action to

increase the integrity of the bone through changes in shape and structure (Lanyon, et

al. 1982). Levels and types of stress placed on the skeleton throughout an individuals

lifetime allow for changes in function through remodeling. This plasticity of bone is an

adaptation of vertebrates which also acts to shape the abilities and actions of a species.

Metabolic bone disease is caused by nutrient deficits in the body affecting bone

function. Vitamin D deficiency is well-known as the cause of rickets, while malnutrition

and illness also has an effect on bone integrity. Differences in subsistence practices

vary the nutrients consumed by people in different communities. Additionally, natural

forces can inhibit crop growth and cause group displacement. Health problems caused

by food shortage, overcrowding, and lack of hygiene may also have an effect on bone

metabolism directly or through increased risk of infection. Breastfeeding/weaning culture

strongly affects infant nutrition, and nutrient deficits in mothers can cause congenital
bone disease in their children (Brickley and Ives 2008). Populations exposed to these

problems adapt over time by conserving energy, storing nutrients in the body, and

modifying bodily processes to survive more easily under these conditions. Humans

living at higher latitudes have a polymorphism in the vitamin D binding protein gene.

These individuals are exposed to less sunlight on a daily basis. UV rays activate vitamin

D3 production, so this polymorphism is likely related to decreased levels of vitamin D in

the body (Bouillon 2010).

Disease exposure and transmission are another aspect of human society that is

both affected by and reliant on both genetic and social factors. In studies of HIV

symptomatology, researchers have found a variety of bone disorders, especially

osteopenia and osteoporosis, caused by the virus (Mondy, et al. 2003). The virus

appears to cause a switch in cell activity to the production of osteoclastogenic

cytokines. These proteins signal an increase in osteoclast activity, causing the observed

bone loss (Vikulina, et al. 2010). In addition to increasing the chance of fracture, this

effect also impacts an individuals ability to heal when a fracture occurs. It is not the

virus itself that slows bone healing, but the effects it has on bone density (Richardson,

et al. 2008). Multiple studies also hypothesize that some anti-viral medications may also

cause loss of bone mass, though their effects are highly variable and mechanisms not

fully understood (Mondy, et al. 2003, Vikulina, et al. 2010). Access to medication is

another aspect of human society heavily intertwined with both biology and culture,

shaping both the evolution of the human species and that of the microbes directly

affected by the drugs.

 Corruccini and Beecher (1982) observed variation in occlusal features of squirrel

monkeys based on diet. Monkeys raised on soft diets developed narrower arches and

rotated, displaced, and crowded teeth. Those raised on hard foods developed

morphologies that were more accommodating to their tooth eruption. In human society,

the transition to agriculture may have had the same effect. Comparisons of hunter-

gatherer and farmer remains from approximately 2000 years BP show differences in

craniofacial morphology. Farming populations from the Argentine Center-West generally

have smaller masticulatory and posteoneural facial features, such as changes to the

palate and shorter maxillary arches, than their hunter-gatherer counterparts (Sardi,

Novellino, and Pucciarelli 2006). In comparing skulls from the 16th and 17th centuries

with modern, living individuals, the appearance of differences, such as anterior rotation

of the mandible and translocation of the maxilla, supports the hypothesis that

masticatory activity affects craniofacial development. Foods, especially for infants, have

become significantly softer over the past couple centuries, resulting in less activity and

effort needed by the jaws of modern humans in order to chew food (Varrela 1992). The

autonomy of humans constructed this modern lifestyle, while simultaneously, the diets

of individuals perpetuated by this lifestyle have affected the mechanical digestive ability

of human communities.

Just as reduction in use of the masticatory muscles has the ability to affect facial

morphology, the disuse of other muscles has an effect on the structure of the rest of the

skeleton. Certain types of nerve damage cause decreased ability to control particular

muscles. Changes to muscle and bone occur in response to this decrease in activity.

While the vascular system continues to reach the affected area, maintaining the quality
of bone, the overall bone mass decreases (Gillespie 1954). Since forces are no longer

being activity exerted on the bone, the body has no need to continuously build new

bone.

Human Walking

Changes in locomotive posture over thousands of years as well as the actions of

current primates allow them to perform the actions involved with their varying modes of

locomotion. Increased loads on particular joints result in selection for broader bone

surfaces to allow for more muscle attachments (Swartz 1989). In this way, locomotor

behaviors both affect and are constrained by musculoskeletal characteristics. For

example, the broad surface of the human pelvis allows attachments of the large human

gluteal muscles to propel bipedal movement. The shortened ilium, as compared to

chimpanzees, shifts the center of mass back, over the hips, reflecting a history of

selection for mechanical features that improve bipedal movement (Lovejoy 1988). The

evolution of bipedal walking was simulated by Hase and Yamazaki (1998). Their model

took a population with chimp-like features over a series of 5000 step changes with the

result of modern human-like morphology. This program incorporated algorithms that

simulated evolution towards decreased energy use and lessened strain on the skeleton

and muscles (Hase and Yamasaki 1998). Bipedality was not simply a decision by

chimp-like ancestors to walk upright and morphological changes followed suit, but each

stage of change involved optimal adaptation for whatever niche the communities

occupied at that stage.

 The use and manufacturing of stone tools by early Hominid species created

another means through which humans could interact with their environments. The ability

to process carcass meat and bone marrow changed diet and nutrition as well as

hominid behavior. Early Homo species at the beginnings of widespread tool use were

able to move away from harder, grittier foods, changing the digestive actions involved

with procuring sustenance. Because of these lifestyle changes, differences in dental

morphology are seen between the first Homo species and earlier hominid species. Gut

size also changed with a reduction of stomach and colon size in response to a more

meat-based diet, involving less processing of plant fibers. The small intestine developed

further to allow for increased fat digestion. Scavenging activities of early Homo

individuals required changes to social structure and interactions. Communities

developed centers for scavenging endeavors to where scavengers could return to

process their carcasses (Cameron and Groves 2004).

Early Homo fossils are classified based on differences in skeletal morphology.

The variation in behavior and selection that led to these differences cannot be directly

observed. Hypotheses about some traits can be made based on information about the

environment and hominid interactions with it. Drastic climactic changes let to many

hominin speciation events. Habitats in the Pleistocene were frequently changing, and as

hominins moved out of Africa, new resources became available. Dmanisi, Georgia was

rich in food resources and tool materials and the appearance of land bridges allowed for

easy movement from East Africa to Eurasia. With the emergence of Neanderthals came

larger brain case size and more rounded cranial features. Neanderthals are associated
with fine crafted tools and are believed to have interacted in complex social systems

(Cameron and Groves 2004).

Within Cultural Context

Bone plasticity increases the possible abilities of an individual. Increases in bone

and muscle mass make more strain-heavy activities possible. This has a subsequent

effect on the actions of a species. Because the morphology of the bone is the result of

the musculoskeletal system responding to environmental strains, bioarchaeologists are

able to use this data to make conclusions about the activities of ancient peoples.

Archaic humans appear to have broader surfaces on their scapulae and more curved

radii, as the result of more strain being placed on the shoulder and the corresponding

development of more musculature. Additionally, early modern humans have differently

shaped femur bones, suggesting that archaic humans needed more robust leg bones to

allow for the greater stress placed upon them (Lieberman and Shea 1994).

While archaic humans were able to respond to stressful environments, humans

also have the ability to modify their lifestyles such that these strains are no longer

necessary, as seen in more modern populations. Children living in different societies

have differing expectations placed upon them at varying stages of their lives. These

roles are never static either. Some 6 year olds, in Mayan societies, may act as

caregivers for younger children, while others, in North America and Europe, may be

expected to sit still in a classroom all day. All communities involve a degree of variation,

and practices are always changing as the knowledge and ideologies of the people

change. In the US, it is difficult for us to to imagine times before child labor laws and
standardized education. American children in the past grew up under very different

circumstances than children today. Cultural practices involve a mixture of inherited

behaviors and responses to forces placed on them by current peoples (Rogoff 2003).

It is difficult to sort out which structures stem from genetic history and which are the

result of physical activity. Cultural and biological evolution are so interconnected that

trying to separate the two is, in some ways, a useless endeavor (as seen earlier with the

evolution of plasticity). Trying to understand one or the other in isolation separates them

from the context in which they occur. Human morphology and variation can only be

understood within context of all current and historical factors (Ruff, Holt, and Trinkaus

2006).

The global incidence of musculoskeletal disorders has attracted the focus of

international healthcare organizations. Villages in Botswana are no exception.

Demanding physical labor caring for crops, maintaining yards, and moulding bricks are

an essential part of villagers lives, holding high sociocultural value to them. These

activities involve maintaining specific postures that may be associated with the lower

back pain experienced by some of the villagers. It is common for villagers to walk long

distances to collect water and firewood. Chronic pain prevents villagers from performing

these culturally and biologically important tasks. Emigration of middle-generation

individuals and the spread of morbid diseases, like AIDS, has shifted the social roles for

Batswana, forcing individuals with musculoskeletal disorders to struggle with more

responsibilities and larger work loads (Hondras, et al. 2016).

Examining the remains of past societies can reveal information about the health

and cultural practices of past peoples. The poor living conditions in late Roman
Dorchester are reflected in the bones of children buried in its cemeteries. Rickets and

anemia were common in the observed population. Children buried in Christian-style

graves suffered rib fractures, thought to be caused by rickets and scurvy, and

pathological conditions associated with anemia. Poor nutrition and lack of exposure to

sunlight as an infant likely caused these conditions. Practices adopted by some

communities, like fasting and poor quality diets of newborns and swaddling would cause

these effects (Lewis 2009). In developing these cultural norms, these communities

played a role in shaping the survival rates of their children and the practices inherited by

future generations.

Recently, humans have started to develop their own, synthetic organs. Scientists

continue to experiment with both organic and inorganic materials to replicate the

features and functions of human tissues. Scaffolds comprised of calcium phosphate

ceramics are engineered and tested under mechanical stress in efforts to recreate bone

tissue. The goal is to develop a structure with the porosity of bone and equivalent

compressive strength (Ramay and Zhang 2003). Niche construction at this level is the

direct manipulation of human biology. Advancement of technology over time and need

for therapeutic bone reconstruction, both the result of human society and behaviors,

have driven science to this point and these actions will continue to shape the human

species and the world.

Conclusion

The combination of biological and social factors that contribute to bone

development are best studied within the framework of niche construction theory. The
role of the skeleton in human behavior and its plasticity in response to the environment

make it an easy candidate for the incorporation of cultural anthropological theory into

the field of biology. Margaret Lock has expressed concern that the increased study of

epigenetics by molecular biologists will drive evolutionary studies back to reductionism,

with a focus on cell function and gene expression inheritance. She urges

anthropologists to pursue research that will explore the contexts surrounding epigenetic

discoveries (Lock 2012). Niche construction theory provides a clear framework but

takes into account the necessary dynamics of lifes processes. It recognizes that

humans both impact and are influenced by their biological, environmental, and

sociocultural circumstances and combines these networks to allow researchers to study

the complexities of human life while keeping their research subject within its real-life

context.. Understanding these complex interactions between humans and their

environments are necessary for understanding the development of the human skeletal

system. Placing bone formation and remodeling within the niche construction

framework encourages further research into the intertwined processes of biological and

cultural coevolution.
 References

Bouillon, Roger 2010Genetic And Environmental Determinants Of Vitamin D Status. The

Lancet 376(9736). Elsevier BV: 148-149.

Brickley, Megan and Rachel Ives 2008The Bioarchaeology Of Metabolic Bone Disease.

1st edition. Amsterdam: Academic Press.

Cameron, David W and Colin P Groves 2004Bones, Stones, And Molecules. 1st edition.

Burlington, MA: Elsevier Academic Press.

Carter, D. R., B. Mikic, and K. Padian 1998Epigenetic Mechanical Factors In The

Evolution Of Long Bone Epiphyses. Zoological Journal Of The Linnean Society

123(2). Wiley-Blackwell: 163-178.

Carter, D.R., M.C.H. Van der Meulen, and G.S. Beaupr 1996Mechanical Factors In

Bone Growth And Development. Bone 18(1). Elsevier BV: S5-S10.

Corruccini, R. and R. Beecher 1982Occlusal Variation Related To Soft Diet In A

Nonhuman Primate. Science 218(4567). American Association for the

Advancement of Science (AAAS): 74-76.

Day, Rachel L., Kevin N. Laland, and F. John Odling-Smee 2003Rethinking Adaptation:

The Niche-Construction Perspective. Perspectives In Biology And Medicine

46(1). Johns Hopkins University Press: 80-95.

Gibson, James J 1950The Perception Of The Visual World. 1st edition. Boston:

Houghton Mifflin.

Gilbert, Scott F and David Epel 2009Ecological Developmental Biology. 1st edition.

Sunderland, Mass.: Sinauer Associates.

Hall, Brian K 2005Bones And Cartilage. 1st edition. San Diego, Calif.: Elsevier

Academic Press.

Hase, Kazunori and Nobutoshi Yamazaki 1999Computational Evolution Of Human

Bipedal Walking By A Neuro-Musculo-Skeletal Model. Artificial Life And Robotics

3(3). Springer Nature: 133-138.

Hazelwood, Scott J, R Bruce Martin, Mark M Rashid, and Juan J Rodrigo 2001A

Mechanistic Model For Internal Bone Remodeling Exhibits Different Dynamic

Responses In Disuse And Overload. Journal Of Biomechanics 34(3). Elsevier

BV: 299-308.

Hondras, M, J Hartvigsen, C Myburgh, and H Johannessen 2016Everyday Burden Of

Musculoskeletal Conditions Among Villagers In Rural Botswana: A Focused

Ethnography. Journal Of Rehabilitation Medicine 48(5). Acta Dermato-

Venereologica: 449-455.

Laland, K. N., J. Odling-Smee, and M. W. Feldman 2001Cultural Niche Construction

And Human Evolution. Journal Of Evolutionary Biology 14(1). Wiley-Blackwell:

22-33.

Laland, Kevin N. and Michael J. OBrien 2011Cultural Niche Construction: An

Introduction. Biological Theory 6(3). Springer Nature: 191-202.

Laland, Kevin N., Jeremy R. Kendal, and Gillian R. Brown 2007The Niche Construction

Perspective. Journal Of Evolutionary Psychology 5(1). Akademiai Kiado Zrt.:

51-66.

Lanyon, L.E., A.E. Goodship, C.J. Pye, and J.H. MacFie 1982Mechanically Adaptive

Bone Remodelling. Journal Of Biomechanics 15(3). Elsevier BV: 141-154.

Lewis, Mary E. 2009Life And Death In A Civitas Capital: Metabolic Disease And Trauma

In The Children From Late Roman Dorchester, Dorset. American Journal Of

Physical Anthropology 142(3). Wiley-Blackwell: 405-416.

Lieberman, Daniel E. and John J. Shea 1994Behavioral Differences Between Archaic

And Modern Humans In The Levantine Mousterian. American Anthropologist

96(2). Wiley-Blackwell: 300-332.

Lock, Margaret 2013The Epigenome And Nature/Nurture Reunification: A Challenge For

Anthropology. Medical Anthropology 32(4). Informa UK Limited: 291-308.

Lovejoy, C. Owen 1988Evolution Of Human Walking. Scientific American 259(5).

Springer Nature: 118-125.

Lumsden, C. J. and E. O. Wilson 1980Translation Of Epigenetic Rules Of Individual

Behavior Into Ethnographic Patterns. Proceedings Of The National Academy Of

Sciences 77(7). Proceedings of the National Academy of Sciences: 4382-4386.

Miller, George A. 2016Linguistic Communication As A Biological Process. In Biology And

The Human Sciences. 1st edition. J. W. S. Pringle, ed. London: Oxford University

Press.

Mondy, Kristin, Kevin Yarasheski, and William G. Powderly et al. 2003Longitudinal

Evolution Of Bone Mineral Density And Bone Markers In Human

Immunodeficiency VirusInfected Individuals. Clinical Infectious Diseases 36(4).

Oxford University Press (OUP): 482-490.

Perrimon, N., C. Pitsouli, and B.-Z. Shilo 2012Signaling Mechanisms Controlling Cell

Fate And Embryonic Patterning. Cold Spring Harbor Perspectives In Biology 4(8).

Cold Spring Harbor Laboratory Press: a005975-a005975.

Ramay, Hassna R.R and M Zhang 2004Biphasic Calcium Phosphate Nanocomposite

Porous Scaffolds For Load-Bearing Bone Tissue Engineering. Biomaterials

25(21). Elsevier BV: 5171-5180.

Richardson, J., A. M. Hill, and C. J. C. Johnston et al. 2008Fracture Healing In HIV-

Positive Populations. Journal Of Bone And Joint Surgery - British Volume 90-

B(8). British Editorial Society of Bone & Joint Surgery: 988-994.

Richerson, Peter J and Robert Boyd 2005Not By Genes Alone. 1st edition. Chicago:

University of Chicago Press.

Rogoff, Barbara 2003The Cultural Nature Of Human Development. 1st edition. Oxford

[UK]: Oxford University Press.

Ruff, Christopher, Brigitte Holt, and Erik Trinkaus 2006WhoS Afraid Of The Big Bad

Wolff?: WolffS LawAnd Bone Functional Adaptation. American Journal Of

Physical Anthropology(129). Wiley Online Library: 484-498.

Sardi, Marina L., Paula S. Novellino, and Hctor M. Pucciarelli 2006Craniofacial

Morphology In The Argentine Center-West: Consequences Of The Transition To

Food Production. American Journal Of Physical Anthropology 130(3). Wiley-

Blackwell: 333-343.

Sardi, Marina L., Paula S. Novellino, and Hctor M. Pucciarelli 2006Craniofacial

Morphology In The Argentine Center-West: Consequences Of The Transition To

Food Production. American Journal Of Physical Anthropology 130(3). Wiley-

Blackwell: 333-343.

Sinha, Chris 2006Epigenetics, Semiotics, And The Mysteries Of The Organism.

Biological Theory 1(2). Springer Nature: 112-115.

Stiles, Joan 2000Neural Plasticity And Cognitive Development. Developmental

Neuropsychology 18(2). Informa UK Limited: 237-272.

Swartz, Sharon M. 1989The Functional Morphology Of Weight Bearing: Limb Joint

Surface Area Allometry In Anthropoid Primates. Journal Of Zoology 218(3).

Wiley-Blackwell: 441-460.

Varrela, J. 1992Dimensional Variation Of Craniofacial Structures In Relation To

Changing Masticatory-Functional Demands. The European Journal Of

Orthodontics 14(1). Oxford University Press (OUP): 31-36.

Vikulina, T., X. Fan, and M. Yamaguchi et al. 2010Alterations In The Immuno-Skeletal

Interface Drive Bone Destruction In HIV-1 Transgenic Rats. Proceedings Of The

National Academy Of Sciences 107(31). Proceedings of the National Academy of

Sciences: 13848-13853.