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BIOSOCIAL INHERITANCE: A FRAMEWORK FOR THE
STUDY OF THE INTERGENERATIONAL
TRANSMISSION OF HEALTH DISPARITIES
Morgan K. Hoke
Northwestern University
Thomas McDade
Northwestern University
Proponents of global market expansion claim that economic growth brings prosperity and
creates a more equitable global society, yet these efforts may result in the exacerbation of eco-
nomic disparities that translates into significant disparities in health. These trends underscore
the importance of research examining the social determinants of health in an increasingly
unequal world. This article brings together concepts from biocultural anthropology, develop-
mental origins of health and disease, and research examining the long-term effects of early
environments on social capital to formulate the notion of biosocial inheritance. Biosocial
inheritance is the process whereby social adversity in one generation is transmitted to the
next through reinforcing biological and social mechanisms that impair health, exacerbating
social and health disparities. Such a theoretical framework considers contemporary and his-
toric political-economic forces that shape inequalities in human health, across generations.
We examine the biosocial determinants of growth in the Andes as a case study of biosocial
inheritance, developing a model that will illuminate the contemporary and historic sources of
persistent poverty in low-income communities around the world. By highlighting the biosocial
mechanisms underpinning the intergenerational transmission of poverty, this model allows
for the formulation of interventions aimed at breaking the cycle of poverty and a reconsider-
ation of the way we think about poverty and social mobility. [inequality, health disparities,
intergenerational, biosocial inheritance, biocultural anthropology]
Advances in science, technology, and public health have led to a shift in global life
expectancy from an average of 30 to 67 years between 1800 and 2000 (Riley 2001).
Countries such as Japan and Switzerland now boast life expectancies of up to 82 years
(Gapminder 2008). However, significant inequalities remain between nations, with life
expectancies of just 45 and 47 years in Sierra Leone and Botswana respectively (Gapminder
2008). Further, beyond the obvious macrolevel disparities, rapidly growing inequities in
health due to social and economic inequalities also exist “within” countries. While we
have taken great steps forward in health and health technologies, access to these benefits
has not been disseminated equally, and in many places, unequal access contributes to
ANNALS OF ANTHROPOLOGICAL PRACTICE 38.2,pp.187–213. ISSN: 2153-957X. C2015 by the American
Anthropological Association. DOI:10.1111/napa.12052
Annals of Anthropological Practice 38.2/Biosocial Inheritance 187
TAB L E 1. Generational Effect Definitions
Cross-generational: Refers to characteristics in one generation that affect outcomes in the next
generation. The effects themselves may not be shared across generations. Example: The
nonepigenetic effects incurred through the prenatal environment can largely be considered
intergenerational. A mother’s consumption of mercury-laden fish can lead to a higher mercury
concentration in the developing fetus and thus reduce cognitive function. This process is the result
of intergenerational effects, an act in one generation that significantly affects another. The exposure
may have distinct effects on the two generations involved. In the case of mercury exposure, there
may be little consequence for the mother but significant effects on the fetus.
Multigenerational: Refers to direct exposures that result in significant effects on more than two
generations, simultaneously. Example: A gestating woman is exposed to an endocrine disruptor that
affects herself, her gestating female fetus, and the germ line of the third generation whose
primordial germ cells are present after two weeks of gestation. All three generations experience
altered outcomes as a consequence of direct exposure to the endocrine disruptor.
Transgenerational: Refers to effects that are transmitted through the germ line or other mechanisms
leading to affects in multiple generations without direct exposure and thereby may affect more than
three generations. Example: Kuzawa’s proposed model of “phenotypic inertia” (2005) wherein the
metabolic phenotype of a given individual is understood as the product of a long-term, integrated
signal of the nutritional environment over multiple matrilineal generations. Epigenetic changes
made in one generation are passed onto the next without the subsequent generation experiencing
the precipitating conditions directly, as would be the case in the fourth generation (no direct
exposure, only epigenetic transmission) of the above scenario with exposure to an endocrine
disruptor (Skinner 2008).
growing health disparities across different segments of the population (Olshansky et al.
2012).
The persistence of poverty in the face of market expansion and improving conditions
around the world has led scholars to examine the pathways through which inequality
perpetuates itself across generations (e.g., Harper et al. 2003). However, only recently has
this cross-, multi-, or transgenerational work been extended to the examination of health
disparities. With the proliferation of this work has come debate over the meanings of
terms such as inter-, cross-, multi-, and transgenerational. In order to avoid confusion
throughout this article, we utilize these terms in accordance with the definitions found
in Table 1.
Research on population health emphasizes the role of early-life environments in
shaping trajectories of health, thereby leading to a proliferation of work linking early-life
experiences with later-life outcomes. While some of this work emerged out of research
on health inequalities (Blane et al. 2007), it has yet to fully engage with research and
methods from other disciplines (Richter and Blane 2013). Such engagement may allow for
a better understanding of the ways in which macrosocial and political-economic forces
structure the formation and perpetuation of disparities through early-life environments.
Additionally, such a perspective has rarely been applied in the realm of global health,
where it may be particularly useful in understanding and intervening in the cycle of
poverty.
We argue in this article that it is necessary to conceptualize early-life environments
as the product of intergenerational processes and contemporary political-economic
188 Annals of Anthropological Practice 38.2/Biosocial Inheritance
contexts. Adult health is shaped by early life, which is in turn a product of early en-
vironments influenced by ongoing political and economic and intergenerational pro-
cesses. By explicitly understanding early-life environments—and their role in shaping
adult health—as the result of intergenerational or transgenerational forces, scholars can
better situate contemporary patterns of health and disease in their political-economic
and historic contexts and therefore intercede more effectively.
The purpose of this article is to put forth a coherent framework for examining the
cross-, multi-, or transgenerational processes that lead to social disparities in health.
We begin with a brief review of the relevant literature related to health disparities and
transgenerational effects in the fields of biocultural anthropology, life-course epidemiol-
ogy, and research examining the long-term effects of early environments and health on
social capital. Next, we present the concept of biosocial inheritance, a framework that
integrates the study of contemporary and past political-economic forces shaping inequal-
ities in health with insights from the study of early life and transgenerational influences
on health and disease across the life course. The biological mechanisms through which
biosocial inheritance can occur will be described along with a brief discussion of their
incorporation into anthropological practice. To demonstrate the utility of this concept,
we present a case study in which the notion of biosocial inheritance is utilized to examine
the influences of height among the high-altitude population of Nu˜
noa, Peru. Finally, a
discussion of the implications of this framework and directions for future research will
be presented.
BACKGR OUN D
Biocultural Anthropology and Health
The biocultural perspective in anthropology is one that acknowledges the myriad ways in
which culture and biology are inextricably united in “a continuous feedback relationship
of ongoing exchange” (Lock 1998:410). The holistic perspective of biocultural inquiry
allows anthropologists to examine a diverse array of topics across time with human
health often serving as a primary interest. It incorporates both evolutionary and social
perspectives, with biological and social variables serving as cause and effect (Wiley 1992).
Since the late 1990s, the “new” biocultural synthesis has sought to foreground political-
economic issues in the study of human biology. These scholars employed political-
economic analysis in the study of “local biologies” (Lock and Kaufert 2001), thereby
tying them to larger global and historical contexts (Goodman and Leatherman 1998a).
In particular, these scholars called for a shift in the focus of biocultural anthropology and
notions of health from evolutionary theory and adaptability to the inclusion of political
economic analysis in the study of human biology both past and present. This increased
consideration of historicized power and its dynamics (Roseberry 1998;Wolf1982) allows
for a more nuanced understanding of both social and cultural variables in biocultural
analyses of human health. Prior to these changes, the exclusion of political-economic
contexts led to discussions of human variation that relied too heavily on adaptationist
explanations and that now appear somewhat simplistic.
Annals of Anthropological Practice 38.2/Biosocial Inheritance 189
Anthropologists utilizing a biocultural approach can be found in every subset of an-
thropological research from medical anthropology to DOHaD (developmental origins
of health and disease). Because biocultural anthropology is still evolving, there are per-
spectives and lines of inquiry that it has yet to fully incorporate. For instance, while
there has been a call for the production of biological ethnography (Wiley 2004)orthe
incorporation of more qualitative methods into the study of human biology, this new
anthropological genre has yet to make its way into mainstream practice. As biocultural
scholars, we continue to struggle to incorporate ethnographic insight and detail into our
work. Other scholars argue that there has been a lack of attention to culture or that
we operate with an unproblematized notion of culture (Dressler 2005). Finally, while
biocultural work has long acknowledged the utility of a life-course perspective, there has
been a lack of biocultural research examining the role of intergenerational and transgen-
erational forces on contemporary human variation and health. As we continue to address
these gaps, increasingly incorporating methods and theory from cultural anthropology,
perhaps we may gain greater understanding of these important processes.
Life-course Epidemiology and the DOHaD
Life-course epidemiology originated in the early 20th century with the early work of
Kermack et al. (1934), and it expanded in the 1990s through three primary strands
of research: the study of biological programming, birth cohort research, and health
inequalities research (Blane et al. 2007; for additional reviews see also Ben-Shlomo and
Kuh 2002; Gluckman and Hanson 2006;Kuhetal.2003; Smith 2003). Key to the
origin of life-course epidemiology is research examining biological programming or “the
developmental origins of health and disease” (DOHaD, see Gluckman and Hanson
2006). Research from this field has suggested a significant role for early environments
in the programming of outcomes ranging from hypothalamic–pituitary–adrenal (HPA)
axis and immune function to behavior and temperament. While much of the work
in this area is limited to animal research, revelations from the study of humans have
yielded important insights regarding early metabolic programming and increased risk
of cardiovascular disease later in life (Barker et al. 1992,1993a, 2002). Such work has
illuminated the importance of examining the process of human development in order
to understand later-life outcomes, implying an important role for consideration of the
principles of DOHaD in the study of health disparities.
Furthermore, DOHaD has dramatically shifted our understanding of the influence of
genes, environment, and their interaction in phenotypic outcomes. Research on DOHaD
has led to increased attention to and intervention during developmental periods on an
international scale and includes nutritional and aid programs targeting gestating women
and children under the age of five. However, the effectiveness of these interventions
may be limited due to a lack of contextual depth within this research; it is rarely
situated within the cultural, social, or political-economic context of the populations
examined. Furthermore, the majority of work in the field of DOHaD relies heavily on
experimental animal models, which reinforces the notion that political economy and
other complicating factors can be ignored (for further consideration of the use of animal
models see Williams et al. 2004). Finally, there is a lack of explicit intergenerational or
190 Annals of Anthropological Practice 38.2/Biosocial Inheritance
transgenerational focus; rather the “prenatal environment” or the maternal environment
is considered as if it emerged de novo. Rather than acknowledging the maternal body
as the product of ongoing physiological, social, and political-economic processes, these
influences on maternal physiology are often placed within an analytical black box and
ignored.
Also crucial to the development of life-course epidemiology is the use of large birth-
cohort studies, particularly the National Survey of Health and Development undertaken
in Europe (Mann et al. 1992; Wadsworth 1991). In the 1990s, the members of this cohort
were reaching the stage of life when chronic diseases begin to emerge allowing for the
observation of correlations between early-life events and later-life outcomes. Such work
was supported by findings from additional studies such as the Dutch famine study and
others, which linked nutritional stress in utero with increased risk of cardiovascular
disease later in life (Lumey 1992; Lumey et al. 2009; Roseboom et al. 2001,2006).
These findings have also led to proliferation of cohort studies, which although costly and
time consuming continue to provide significant insight on the role of development and
cumulative effects in health throughout the life course.
Finally, life-course epidemiology was largely influenced by research on health inequal-
ities. A number of major studies have revealed the significant relationship between social
class and socioeconomic status and health (Blane 1995; Goldblatt 1990;Marmotetal.
1984; Rose and Marmot 1981; Wilkinson 1996), some of which have explicitly consid-
ered their effect on health across the life course. Research in this area has revealed the
social gradient in health whereby health improves incrementally, in step with increasing
social and economic status (Adler et al. 1993). Research focusing on the social gradient
has demonstrated an important role for objective and subjective, or perceived, status in
determining health outcomes (Adler et al. 2000). While a life-course perspective may
account for some intergenerational influences (Halfon and Hochstein 2002), explicit
consideration of intergenerational variables is usually limited to some measure of in
utero experience and possibly a consideration of parental anthropometrics. Furthermore,
this research often lacks an explicit discussion of the cyclical nature of social inequalities
and a framing that understands these inequalities as both trans- and intergenerational
phenomena.
Relating Early Health and Environments to Socioeconomic Capital Later in Life
While research in the field of life-course epidemiology has demonstrated the relationship
between early environments and health outcomes, research in the fields of economics,
demography, and sociology has connected these early-life inputs with health and socioe-
conomic outcomes later in life. Like biocultural anthropology, much of this research
considers the origins and perpetuation of poverty and inequality (Palloni 2006). This
extensive body of work includes examinations of prenatal environments and experience
(for review see Almond and Currie 2011 and Currie 2011) and early-childhood health and
experience (for review see Elo and Preston 1992; Currie and Almond 2011). These bodies
of work suggest that early health and experience have significant long-term effects on
health and socioeconomic outcomes such as cognitive abilities, educational attainment,
Annals of Anthropological Practice 38.2/Biosocial Inheritance 191
earnings, and employment (Almond et al. 2012). The relationship among prenatal/early
nutrition, growth, and later socioeconomic outcomes is now well established and serves
as an important example of this body of work (Behrman 1996; Martorell 1995,1999;
Martorell et al. 1995, Martorell et al. 2010a, 2010b).
Unlike research in life-course epidemiology and DOHaD that demonstrates a some-
what limited acknowledgement and engagement with intergenerational influences, schol-
arly work considering the effects of child health and early environments on social and
economic capital has paid explicit attention to the reproduction of inequalities across
generations (Currie and Moretti 2005; Palloni 2006) and the mechanisms through which
it occurs. This body of work considers the effects of parental characteristics (Behrman
and Wolfe 1987; Ermisch and Francesconi 2001), in utero exposures/prenatal health
(Almond 2006; Figlio et al. 2013; Lin and Liu 2012; Neelson and Stratman 2010;
Nelson 2010;Steinetal.1975), and the multigenerational transmission of maternal health
shocks or exposures experienced by the mother in early childhood (Almond and Chay
2006; Almond et al. 2011; Currie and Moretti 2005;Stewartetal.1980) on health and
socioeconomic outcomes. Research in this area has faced criticism for its limited ability
to assign causality or directionality as well as a difficulty in measuring or operationalizing
complicated variables such as child health status. While some scholars acknowledge these
limitations (Almond et al. 2011; Palloni 2006), much of the work in this field continues
to be riddled with hyperdeterministic and overly causal language and plagued by grainy
measures of exposure. Furthermore, it is difficult to identify mechanisms in many of the
large cohort studies utilized in this research. Still, this work makes an important con-
tribution to our understanding of social mobility and the transmission of social status
across generations. This contribution is made even stronger when synthesized with work
from other disciplines.
BIOSOCIAL INHERITANCE: THE MODEL
We present the framework of biosocial inheritance, which is designed to facilitate the in-
tegration of political-economic analysis and sociohistorical variables into considerations
of intergenerational health outcomes. We seek to unite work from the fields of biocultural
anthropology and DOHaD with research on the long-term effects of early environments
on socioeconomic attainment. It is only when these lines of inquiry meet that we gain
a holistic and clearer understanding of issues of poverty, health disparities, and social
mobility in this increasingly globalized world. Importantly, this approach highlights the
intergenerational mechanisms, both social and biological, that lead to the transmission of
socioeconomic status and health across generations. Further, these mechanisms are inti-
mately connected such that deeming them either biological or social is highly misleading,
thus we adopt the term biosocial.
Emerging research shows that these intergenerational forces have a major influence
on an individual’s health and social trajectory and, therefore cannot be ignored in con-
temporary considerations of population health and well-being. Specifically, our interest
lies in examining the parental histories and political-economic contexts that create early
192 Annals of Anthropological Practice 38.2/Biosocial Inheritance
offspring environments. These environments provide the basis for biological and social
programming that affects and is in turn affected by socioeconomic status in adulthood,
ultimately leading to disparate health outcomes and a potential reification of the cycle.
Above all else, this framework should help to ground biological work examining inter-
generational and transgenerational influences on health within the larger sociocultural,
historical, and political economic context of the population and people in question. We
begin with a brief history of its individual components and a definition of biosocial
inheritance. This is followed by a brief presentation of the framework and a discussion of
the possible mechanisms through which biosocial inheritance may operate. We conclude
a consideration of how biosocial inheritance extends previous biocultural research on
health disparities.
On the Origin of Biosocial Inheritance
Before defining biosocial inheritance as a concept it is perhaps most useful to first
consider the meaning of its component parts. In anthropology, the term biosocial has
at times been a controversial one, in large part due to its semantic association with the
field of sociobiology (Goodman and Leatherman 1998b). However, many scholars in
anthropology have used the term successfully to describe phenomena that are not easily
categorized as one or the other (Leonard and Thomas 1989; Panter-Brick 1998;Singer
2011;Thomasetal.1988). Others have used the term to explicitly reference the inextricable
nature of the social and the biological (Pike 2004; Pike and Williams 2006). In the 1990s,
with the advent of the human genome project, Rabinow (1992,1996)employedthe
term “biosociality” to describe the formation of new social relations around a biological
condition; in the case of his work, a certain genotype or the possession of a certain gene.
Within the framework of biosocial inheritance, we use the term biosocial as a way of
dissolving the false dichotomy of the biological and social, acknowledging the mutually
constitutive nature of these seemingly opposing categories, and recognizing the limits of
our language and our past conceptualization of these issues.
The term biosocial also highlights our interest in the larger social, political, and
economic structures that both facilitate and constrain contemporary human health.
While culture plays an important role, the term biosocial reminds us to examine the larger
structures of power at work in creating and maintaining health disparities, answering a call
by political-economic scholars such as Wolf (1982)andRoseberry(1998). Furthermore,
while cultural norms and traditions often play a significant role in population health,
there is a tendency to culture blame, a form of victim blaming wherein a population’s
poor or poorer health status is blamed on either specific or general “cultural practices.”
Culture blaming is ineffective and counterproductive, as we have seen in the case of female
genital cutting (Abusharaf 2006; Gruenbaum 2001; Shell-Duncan 2008). Additionally,
culture blaming often overlooks the relations of power involved in the clash of cultures,
particularly that occurs when the clash involves belief systems and actions based on
the seemingly monolithic and impenetrable practice of science and empirics and those
based in other world understandings. A similar problem is the conflation of inequality
or poverty with culture or “otherness” and the discussion of “the culture of poverty”
Annals of Anthropological Practice 38.2/Biosocial Inheritance 193
(Farmer 2003). Such conflation can lead scholars and others to fail to acknowledge and
engage with the dynamics of power that create and sustain inequality and poverty at
local and global levels (Bourgois 2001). While biosocial inheritance lies squarely within
the purview of biocultural anthropology, by employing the term biosocial rather than
biocultural we hope to keep the aforementioned issues of power at the forefront of our
analysis.
The second part of the term, inheritance, references the process through which traits,
materials, and status are transferred from one generation to the next. The term inher-
itance can refer either to biological or genetic inheritance, such as a genetic disorder, a
uterine environment, or phenotypic appearance (affected by both genetic and epigenetic
influences). It can also refer to socioeconomic or material inheritance that one receives
from parents, grandparents, or other relations such as wealth, land, or status that can
affect life outcomes. Significantly, it also captures those cultural practices and norms
that are shared across generations, as well as elements such as language, memory, and
heritage that are passed on to help to make sense of the past. The term inheritance
applied in this manner harkens back to Lamarck’s (n.d.) theory of inheritance of ac-
quired characteristics, which has received some renewed attention in light of our growing
understanding of epigenetics and the existence of environmentally induced traits that
may transfer across generational lines (Kuzawa 2005). It should be noted that inheritance
does not necessarily imply a sort of static transference; the trait, material, or practice may
shift or change somewhat between generations. Furthermore, biosocial inheritance can
be positive, negative, or both depending on the environmental context. For example, a
child who is born to a relatively poor family living in a small farming community in the
Andes Mountains of Peru where nutrition is highly variable by season may benefit from
the prenatal programming of a metabolic phenotype that allows for rapid fat storage.
Fat stores built during and following the harvest season can buffer against periods of
reduced food availability that precede the next harvest. A child whose family is relatively
well-off may avoid the development of such a metabolic phenotype through supplement-
ing the preharvest period with purchased foods, largely eliminating seasonal variability.
However, as the community becomes incorporated into larger economic systems and
begins the shift toward increasing wage labor, the nutritional highs and lows are replaced
by consistent, readily available, calorie-dense, cheap foods and reduced physical activity.
The poorer child with the “thrifty phenotype” is likely to be at greater risk for energy
imbalance, obesity, diabetes, and other health issues.
Biosocial Inheritance: A Definition and Framework
We can thus define “biosocial inheritance” as the process through which social adversity
or advantage is transmitted across generations through mechanisms both biological and
social in nature. Health and health disparities play a key role in biosocial inheritance as
outcome and mechanism. For example, poor health in adults can lead to the transmission
of poor health to offspring, but it can also lead to reduced productivity and social capital
in the following generation. Additionally, an impoverished environment inherited from
parents can lead to poor child-health outcomes, reduced social capital in the form of
194 Annals of Anthropological Practice 38.2/Biosocial Inheritance
education attainment, employment, and poor health outcomes when those children
reach adulthood. Thus intergenerational influences often have the effect of perpetuating
or amplifying biosocial traits across generations, thereby increasing the importance of an
intergenerational perspective. Biosocial inheritance leads us to consider the generations
prior to that generation with whom we are directly working. It forces us to think
about what cultural practices, social norms, ideologies, and economic opportunities or
constraints are transmitted from one generation to the next and how they interact with
human biology to harm or enhance health. Furthermore, it brings to focus the ways in
which health disparities and poverty interact to compound in a population across time,
increasingly reducing the mobility for some and increasing the opportunity for success
for others, thus contributing to the ever-widening disparities in contemporary society.
The many mechanisms of biosocial inheritance
Biosocial inheritance occurs though a number of different mechanisms. While many of
them appear to be principally biological in nature, their activation, function, and cessation
are often dependent on biological and social inputs. For example, although the pathways
through which epigenetic changes occur and take affect are considered biological, these
changes may be induced by endocrine signals stimulated by a social phenomenon such
as stress, making them biosocial in nature. Many anthropologists are already examining
the numerous mechanisms through which biosocial inheritance may occur (see Table 2).
We will briefly review several key examples that represent important ways in which the
social environment can “get under the skin” and stay there for generations.
Developmental programming represents a major mechanism of biosocial inheritance.
Programming during development can affect numerous systems including metabolism,
the HPA axis, and immune systems. Epigenetic changes, in the form of chromosomal
alterations, histone modifications, and DNA methylation are the proposed mechanisms
for much of the developmental programming that leads to adult disease (Callinan and
Feinberg 2006; Peaston and Whitelaw 2006). However, it is possible, and even likely, that
there are other undiscovered or unidentified mechanisms through which this program-
ming can occur. It is also possible that rather than programming, allostasis or the body’s
constant adjustment to internal and external changes (Sterling 2004,2012; Sterling and
Eyer 1988) may lead to a more rapid decline of body systems and the emergence of disease
states (Seeman et al. 1997), particularly when challenges to allostasis are frequent and
severe. Primarily the result of one’s immediate environment, this allostatic burden can
be understood as, at least in part, a result of intergenerational forces acting through their
influence on our current environment.
Nutrition represents a prime example of a factor in our immediate environment that is
highly mediated by intergenerational influence. For instance, while an infant may express
preferences for certain foods early on, what the infant eats is largely a result of the way
caregivers (most often parents) navigate their own environments. How long a mother
is able to breastfeed her infant may depend on her economic resources, occupation,
education, cultural beliefs, length of maternity leave, spousal and/or family support, and
access to her child or a private place to express and store milk during the workday. The
Annals of Anthropological Practice 38.2/Biosocial Inheritance 195
TAB L E 2. Mechanisms of Biosocial Inheritance
Mechanism Explanation Citations
Epigenetics Epigenetic modifications are changes
induced by environmental exposures
that can be transmitted to
subsequent generations through the
germ line. In certain contemporary
environments, this may lead to
disparate health outcomes.
Kuzawa and Sweet (2009), Anway and
Skinner (2008), O’Brien (2007),
Uddin et al. (2010)
Immune
function
Immune function is often dependent
on intergenerational inputs such as
caretaker choices/restrictions
regarding breastfeeding (antibody
and microbiome transmission) and
early microbial exposures.
Grindstaff et al. (2003), McDade (2002,
2003)
Growth and
development
Growth and development are directly
affected by caretaker status and
genetics as well as long-term social
and cultural factors that are
inherited.
Ramakrishnan et al. (1999),
Venkataramani (2011)
HPA axis
function
Early environments, shaped by
caretakers, may lead to the
development of offspring who
experience HPA activation easily,
return to normal levels more quickly,
or remain activated for long periods
of time, all of which can have
distinct implications for health.
Cottrell and Seckl (2009), Davis and
Sandman (2010), Kapoor et al.
(2008), Tegethoff et al. (2009)
Metabolic
programming
Metabolic programming can take place
during prenatal and early
development as a result of
intergenerational influences or it
may be the result of long term
transgenerational signaling.
Barker (1998), Barker et al. (1993b,
2002), Desai and Hales (1997),
Kuzawa (2005)
range and quality of supplementary foods an infant is given depend on family income,
access, education, and ecology among other things.
Additionally, political or civil conflict may cause individuals to flee conflict areas
for the safety of a nearby city or other locations. Such movement drastically changes
the nutritional environment the next generation is born into. This shift in nutritional
environment also occurs during rapid economic transition when agriculture begins to
take a back seat to the purchase of processed foods. In any of these cases, the change creates
a mismatch of transgenerational metabolic programming and contemporary nutritional
environment implicated in the rise of obesity in middle- and low-income countries (Hales
and Barker 2001; Popkin 2001; Popkin and Nielsen 2003). This same mismatch can be
196 Annals of Anthropological Practice 38.2/Biosocial Inheritance
created by larger social and economic structures, such as the food deserts that emerge
when large corporations refuse to place stores in certain locations due to perceived low
profitability. Even with a rapid shift in nutritional environment, children are often not
the primary agents in the decisions about how and what they eat. These early nutritional
inputs have significant impact on social and biological outcomes throughout the life
course of the child and are largely governed by intergenerational influences. What is
more, some scholars have proposed that these early nutritional inputs may be translated
into epigenetic changes and metabolic programming that may have effects on subsequent
generations (Kuzawa 2005). Therefore, one’s nutritional environment is largely inherited
from parents and other predecessors and also serves to inform the programming of future
generations.
Immune function represents another important mechanism through which inter-
generational social influences interact with biology to affect health. A person’s immune
system develops over the course of childhood and early exposures greatly impact im-
mune function throughout the life course. For instance, a child’s increased microbial
exposure via contact with animals and soil can help to prime the child’s immune system
to develop regulatory networks necessary for effective immune responses in adulthood
(McDade 2003,2005). These exposures can even moderate the relationship between stress
and inflammation in adulthood (McDade et al. 2012), which is in turn associated with
increased risk of cardiovascular disease (Ridker et al. 1998), type II diabetes (Pradhan
et al. 2001), late-life disability (Kuo et al. 2006), and mortality (Harris et al. 1999). These
early-life exposures, as with nutrition, are largely dependent on the socioeconomic status
and activities of the prior generation, parents, and caretakers. While microbes represent
a biological exposure influenced by social factors, inherited social status and subsequent
stress can also represent an important exposure. McDade’s work (2001,2002)inSamoa
demonstrated that status incongruity between inherited, Samoan matai titles and mate-
rial wealth, characterized by technology and Western prestige materials, led to increased
psychosocial stress and reduced immune function. Without careful attention to local,
cultural structures of status inheritance and globalized, Western ideas of prestige, this
health outcome would have remained unexplained and unaddressed.
Beyond these more apparently biological mechanisms, there are also mechanisms
appearing more social or economic that become biological as they begin to impact health.
For example, an impoverished prenatal and childhood environment can lead to impaired
adult productivity both physically through health (Barker et al. 1989,1990,1993b; Blane
et al. 2007; Lawlor et al. 2006;Valdezetal.1994) and cognitively (Richards et al. 2001),
thereby reducing socioeconomic potential. This reduced economic potential/success can
in turn compromise adult health or exacerbate pre-existing health problems. Poor adult
health, specifically poor maternal health, can lead to adverse birth outcomes such as low
birth weight and preterm birth (Dodd et al. 2011), thereby setting up negative biosocial
trajectories for the next generation and thus reinforcing the cycle. Alternatively, social
hierarchies based on race or ethnicity can create stressful environments for gestating
women who may subsequently experience the same adverse birth outcomes noted above
Annals of Anthropological Practice 38.2/Biosocial Inheritance 197
(Collins et al. 2000,2004;CollinsandDavid2009; Giurgescu et al. 2011; Mustillo et al.
2004), leaving their children the same negative trajectories.
The mechanisms of biosocial inheritance are not limited to these examples or those
listed in Table 2, and certainly are more likely to come to light with increased research
focus in this area. The mechanisms through which biosocial inheritance occurs allow
for what can be considered both positive and negative health effects; just as there is a
synergistic relationship between poor health and poverty, there is a similar relationship
between health and material and social success. When compounded across generations,
it is not difficult to see how such mechanisms allow for the concretization and deeper
entrenchment of health disparities. Those individuals with an advantage continue to
improve across generations while those at a disadvantage face significant obstacles that
impede social, economic, and health-based improvements.
CASE STUDY: BIOSOCIAL INHERITANCE OF STATURE IN THE ANDES
We now present a case study that will apply some of the concepts discussed throughout
this article (summarized in Table 3). We apply the notion of biosocial inheritance to better
identify and understand the determinants of growth and adult stature in Nu˜
noa, Peru.
Scholars and policy makers have been investigating the anthropometrics of Peruvians for
nearly a century. The ideas regarding the most powerful determinants of growth have
shifted dramatically across that time period with changing research agendas and shifting
theoretical foci, making it an excellent case study for demonstrating the importance
of incorporating biosocial inheritance in our considerations of contemporary health
disparities.
Despite our knowledge of the immense plasticity of human phenotypes, scholars
continue to seek a genetic explanation for the heritability of height (Weedon et al. 2008;
Yang et al. 2010). This would appear to be a straightforward form of inheritance as one’s
genes represent direct biological inheritance passed from one generation to the next.
However, it has become apparent that certain environmental conditions must be met
for an individual to approach their genetic potential (Silventoinen et al. 2000). This
complicates the conceptualization of genetic inheritance, making it largely dependent
on the social environment as well as the biological environment, and thus biosocial.1
Numerous explanations for reduced Andean stature have been offered through the years,
including adaptation to high altitude (Frisancho 1969,1976,1977; Frisancho and Baker
1970) and chronic malnutrition as a result of poverty and marginalization (Leonard 1989;
Leonard and Thomas 1989; Leonard et al. 1990, Leatherman et al. 1995). Indeed, this
work has demonstrated shifts in levels of chronic malnutrition in communities in the
district due to shifting economic circumstances, and irrespective of relative elevation in
the district (Leatherman 1994). Thus, it has become increasingly clear that a complex
constellation of macro and micro forces, reaching across generations, shape child growth
in Nu˜
noa and it is to these forces that we now turn.
Nu˜
noa is currently segmented into three distinct ecological and economic zones, the
lower valley, the upper valley, and the centrally located town. Though it has traditionally
been a center for alpaca and sheep wool production, which supplied most of the wealth
198 Annals of Anthropological Practice 38.2/Biosocial Inheritance
TAB L E 3. Sociocultural, Historical, and Political-economic Factors in Biosocial Inheritance
Explanation Example from the Case Study
Historical
context
Analyses should include relevant
historical context both recent and
more long term as needed.
Despite having occurred more than 20
years ago, the civil conflict between
the Shining Path and the military
continues to have significant
repercussions for the growth of the
population of Nu˜
noa.
Political-
economic
context
Analyses should consider relevant
political-economic factors such as
government policies, international
involvement, economic development
efforts, and present political conflict.
Local and federal government programs
that seek to provide economic and
nutritional support to children at
risk for stunting. Aid from NGOs
that may alter a family’s economic
and nutritional circumstances.
Mining companies providing wages,
education, and nutritional
supplementation to workers and
their families.
Relations of
power
Analyses should consider past and
present social, political, and
economic hierarchies that may affect
or be affected by health disparities.
Due to their economic or racial/ethnic
status, individuals may experience
difficulty in finding work, receiving
healthcare, and so on, because of
inherited hierarchies present in the
Andes for hundreds of years.
in the region, the last ten years have seen a major economic shift with the emergence of a
dairy industry in some lower sectors. Though there have been some government aid and
subsidies to this industry, the growth is largely driven by an increasing demand for cheese
to be sold in markets and utilized in the restaurants, particularly pizzerias, catering to
tourists in the nearby city of Cusco. This rapid economic change appears to have led to
improvements in the nutritional and economic status of those families living in the lower
valley where water is plentiful and alfalfa for dairy cattle can be grown easily. Families
living in the upper valley have not had the same level of dairying success, as they are
unable to sustain the alfalfa production and herd quality that allows for adequate milk
production year round. Most of these families have continued to rely on herding alpacas
for income, with very limited subsistence agriculture. A relatively low international price
for alpaca fiber over the past decade has severely limited earnings. Important changes
have also occurred for the population that lives in the main town of Nu˜
noa where most
families now rely on wage labor or mercantile activities and foods purchased in markets
for a large part of subsistence, rather than or in addition to agricultural fields outside
of town. These economic differences, largely formed as a result of international political
economic forces (including a recent 9percent growth rate in the Peruvian economy
largely driven by mining revenues and decentralization of income distribution by the
state), may result in significant differential effects on several mechanisms of the biosocial
Annals of Anthropological Practice 38.2/Biosocial Inheritance 199
inheritance of growth: nutrition, metabolic programming, and immune function. We
will explore each of these mechanisms in turn.
Like elsewhere in the world, nutrition in Nu˜
noa is largely dependent on access to high-
quality foods. Access implies proximity, availability, and ability to obtain these foods.
Access in Nu˜
noa is largely influenced by where someone is born and/or where their family
lives (in the upper, lower, or urban zones) and thus what economic activities, access to
land, animals, and labor are available to them. Further, within each of the economic
zones, inequalities may have emerged or perhaps been exacerbated by the activities of
international NGOs who lend money or gift animals to a limited number of families.
In addition to influencing economic activities, poor road conditions and weather during
the rainy season make traveling between the upper zone and the town difficult and costly.
This reduces the number of trips a family can make and thus the quantity of food they
can purchase and bring home. Depending on a family’s economic means, coming into
town may require a six- to eight-hour walk, one way. Even if a family has sufficient money
to purchase food, if they cannot afford transportation or to pay someone to care for their
herds while they are away, they may not be able to spend it. What is more, infrequent
trips mean that fresh produce often fails to last the full interval, leaving the family with
only dried and stored foods for a period of time. However, if a family that lives far into
the countryside and has transportation—as many wealthy landowners still do when they
are not in their second or third homes in nearby cities—or a large social network that can
be tapped into for labor, nutritional, or economic support, access to quality nutrition is
generally good. Thus, it is not simply a matter of where one’s parents live but also their
economic and social means.
Beyond differential economic activities, historical and contemporary social hierarchies
also play a role in influencing how and where people live, as do recent and more long-term
historic events. For example, in the 1980s the civil conflict between the militant Maoist
group the Sendero Luminoso or Shining Path and the Peruvian state brought a period of
upheaval, violence, and fear to Nu˜
noa. In terms of nutritional effects, there was reduced
food availability, increased hunger and malnutrition alongside trauma and high levels of
psychosocial stress (Leatherman et al. 1995).
Scholarship is showing the ways in which these traumas can echo through generations.
As we have seen in previous research, periods of food shortage, political strife, and ethnic
discrimination can have significant effects on stature and other health outcomes, such
as birth weight, in subsequent generations (Lauderdale 2006;Lumey1992; Lumey et al.
2009,2011; Roseboom et al. 2000,2001). Thus, metabolic programming that can take
place when an individual is exposed to starvation in utero or in early childhood, represents
an important biosocial mechanism through which the experience of these traumas may
have been transmitted across generations. This conflict also took a tremendous toll on
human capital. One study has shown that within Peru, the presence of the civil conflict
led to a reduction of human capital and particularly education in the areas hardest hit
(Le´
on 2012). Furthermore, this period of turbulence brought with it additional reshuffling
of property as large landholders and wealthier cooperatives saw much of their land and
animals re-appropriated. While many of Nu˜
noa’s inhabitants fled during this time, others
200 Annals of Anthropological Practice 38.2/Biosocial Inheritance
lacked the means to do so. In some cases, this may have deepened the disparities between
the wealthier population, who were able to seek shelter in the cities, and those poorer
families who had to stay behind to experience hunger, violence, and trauma, all of which
can leave intergenerational traces (O’Brien 2007; Uddin et al. 2010).
Immune function and the experience of illness are also mechanisms through which
intergenerational forces affect growth in Nu˜
noa. First and foremost there is a direct
connection between growth faltering and episodes of illness, particularly diarrheal illness
(Moore et al. 2001; Rowland et al. 1988). In infants, these episodes often occur beginning
with the introduction of complimentary foods depending on issues of caretaker hygiene,
education, and resources. Adult illness can also take a toll on infant and child growth.
Among the agro-pastoral families living in the upper and lower zones of Nu˜
noa, adult
illness often leads to the removal of older children from school to complete the role
in economic production that an ailing adult can no longer fulfill (Leatherman 2005;
see also Leatherman and Jernigan this issue). This often leads to these children falling
behind in school and eventually dropping out, significantly harming their chances of
social mobility and leaving them in a similarly vulnerable position in the future. With
incomplete educations and limited skills, they often opt to remain in the agricultural
fields of their parents rather than obtaining their own or starting new careers. Further,
when economic production falters, though there is effort on the part of mothers to
shield their children (Leonard 1991), food insecurity rises and the nutritional status of the
family may decrease leading to increased experience of stress and worry, compromised
immune function, and growth faltering in children. All of these impacts affect health
outcomes across both generations and have repercussions throughout the life course and
in generations to follow. Stature itself is a part of this picture of reduced social mobility.
Several studies have shown that physical height is linked to career success (Judge and
Cable 2004), income (Bassino 2006; Steckel 1983), socioeconomic status, and education
(Mayer and Selmer 1999).
Finally, we must acknowledge the longstanding racial and ethnic hierarchies in place
since the colonial period that continue to play an important role in social mobility
and status in Nu˜
noa. Markers of indigeneity, such as reliance on Quechua as a primary
language or traditional dress, often inherited from one’s parents, can reduce a person’s
status and potential for social mobility (de la Cadena 2000; Mannheim 1984)andthus
health. Persistent and fierce hierarchies, which also represent inherited social ideas, mean
that these markers are a social burden that children carry with them even as they adopt an
increasingly Westernized lifestyle. The ethnicity and economic choices of their parents
and family, combined with societal norms and ideas, can propel the next generation
forward or limit their mobility and health. When the cycle is established and health
disparities are exacerbated by years of inherited social hierarchies, limited social mobility,
and constrained physiology, the situation cannot be overturned with targeted, single-
point interventions. For example, Leonard (1989) demonstrated a slight improvement
in rates of stunting for the most well-off children and Leatherman et al. (1995) noted
a slight secular increase in growth when compared with the 1960s data. However, a
restudy in the late 1990sbyPawsonetal.(2001) following the political and economic
Annals of Anthropological Practice 38.2/Biosocial Inheritance 201
upheaval associated with the Shining Path showed no improvement over the 1960sand
indicated that positive growth trends may have in fact reversed during that period. Thus,
eliminating malnutrition alone will not alleviate this cycle; political-economic and social
instabilities must also be addressed. Tackling malnutrition may aid in improving certain
aspects of population health. However, the weight of intergenerational forces, such as
nutritional access, metabolic programming, illness, and ethnic or racial hierarchies, mean
a more holistic understanding and intervention is needed.
One reason for the need of holism is that rather than growth based solely on his
or her own individual conditions, the growth of this contemporary Peruvian cohort
is heavily influenced by inheritance from previous generations. First, children are often
bequeathed a similar socioeconomic status as their parents. Although, if aided by a strong
kinship network and collective resources such as a wealthy aunt and uncle in a nearby
city, children may have an opportunity to live away from home, experience a different
nutritional environment, and attend school undisturbed by family illness. Either outcome
functions as a form of biosocial inheritance. This status has a strong effect on their access
to good nutrition and thus growth. Furthermore, their parents’ occupation means the
contemporary cohort may inherit an environment where the balance between survival and
starvation, education or illiteracy, depends on one major incident of parental or family
illness (Leatherman 2005). Second, they may have inherited metabolic programming
based on their mother’s experience of trauma and malnutrition during the 1980sand
early 1990s under the threat of the Shining Path and counterinsurgency efforts. This
programming, as in the case with previously researched famines, may have significant
consequences for their metabolic health both in childhood and later in life (Lauderdale
2006;Lumey1992; Lumey et al. 2009,2011; Roseboom et al. 2000,2001). They have
also inherited a social hierarchy in place since the colonial period in which their dress,
language, and lifestyle may earn them scorn and derision or ease their passage through
society and thus their access to nutrition, health care, and social capital. Even if they adopt
new ways of dressing and earning a living, the choices of their parents and family continue
to mark them until they are able to move far enough away to hide their social inheritance.
It is likely that there are additional elements of biosocial inheritance affecting the lives
and growth of this contemporary Peruvian cohort; we have outlined a few of the most
apparent here. These examples demonstrate the way in which intergenerational forces
play an important role in contemporary health disparities, how these forces act through
mechanisms that function on both social and biological levels, and how by examining
these forces we can better incorporate political economic analysis and historical context
in our biocultural work.
To begin to actually document the effects of biosocial inheritance on growth in
Nu˜
noa, a multitiered study design incorporating population-level growth comparisons
across time and more-detailed contemporary data collection could be employed. The
long history of studying growth in Nu˜
noa may allow for the tracking of generational
shifts in growth. Additionally, it may make feasible the consideration of growth before
and after major political events such as the land reforms occurring just after the study
period in the 1960s and preceding that of the 1980s or before and after the emergence of
202 Annals of Anthropological Practice 38.2/Biosocial Inheritance
the Shining Path or the dairy industry. While datasets from each generation are limited
in that they represent only a cross section and are somewhat variable in age range, they
still lend themselves to meaningful analysis when placed in larger historical contexts.
Such analysis would be complemented by a study of early growth and infant feeding
contextualized in relatively detailed family histories. This initial study would serve as a
baseline for followup over the next 20 years and beyond with collection of anthropometric
and dietary data as well as measures of immune function, incidence of illness, and
other health-related biomarkers. Subsequent periods of data collection would include
repetition of all biological and nutritional data but could also be expanded to include
data on cognitive function, academic achievement, experience of psychosocial stress
or life stressors, and other measures that may shed light on the ways in which early
environments are connected to later-life social and health outcomes. While such studies
do exist elsewhere, by beginning this study with attention to the nuances of social,
economic, and political inequalities as they are in infancy and develop across the life
course, this study would offer the opportunity to make a meaningful contribution to the
biosocial inheritance theory and the study of health disparities.
DISCU SSION: TH E PRACTICE OF STUDYING B IOSOCIAL I NHE RITANCE
Extending Our Understanding of Health Disparities
At this juncture we might ask how exactly the notion of biosocial inheritance moves for-
ward the study of health disparities within biocultural anthropology and more broadly.
First and foremost, biosocial inheritance helps to place bodies in their cultural context,
space, and time, doing away with ideas of universality and instead highlighting their
inextricability from larger historical, political-economic forces (Lock 2013). When we
force ourselves to consider what a contemporary cohort has inherited from previous
generations, we begin to blur the lines of the individual, thereby challenging concepts of
the self and the body and questioning their status as clearly bounded entities (Lock 2013).
But most importantly, by situating contemporary health outcomes in their generational
context, we are forced to grapple with the larger historical and political-economic forces
that have acted across those generations. Biosocial inheritance serves as a natural exten-
sion of work on the spaces of vulnerability and spaces of hope begun by Leatherman
(2005) as well as the study of the biology of poverty (Goodman et al. 1988; Leatherman
and Goodman 1997;Thomas1998). Further, it compels us to reconsider the transmission
of poverty, social mobility, and the popular rhetoric of independence, self-sufficiency,
and pulling oneself up by one’s bootstraps. What is more, biosocial inheritance helps to
demonstrate how these multidimensional, intergenerational processes take place, high-
lighting the complexity of health phenomena that cannot be cured with a magic bullet.
Rather, biosocial inheritance can be used to advocate for the evaluation of programs
based on long-term results rather than immediate outcomes.
Additionally, biosocial inheritance can be understood as a form of embodiment,
whereby the social, political, and economic world around us manifests and perpetuates
itself through our physiology (Krieger 2005). In other words, as we move through life, our
Annals of Anthropological Practice 38.2/Biosocial Inheritance 203
bodies are continually internalizing and then manifesting our social, political, ecological,
and economic environments in the form of physiological phenomena, from the speed
and extent of our growth to the darkness of our freckles or the way we comport ourselves
around others. Anthropological scholars have already utilized the notion of embodiment
to describe the process by which racial inequalities have led to highly disparate health
outcomes between black and white Americans (Gravlee 2009; Kuzawa and Sweet 2009).
Although much of the anthropological work on embodiment has paid limited attention
to the biological body, the use of embodiment in the examination of health disparities
represents a prime opportunity to redress much of the black boxing of physiology that
has taken place in the social sciences and embodiment scholarship throughout the 20th
century (Lock 2013). Furthermore, there has been a call from biocultural scholars to
better engage with this important theoretical concept that has been largely limited to
cultural and medical anthropology (Leidy Sievert 2006).
Biosocial inheritance forces us to reconsider the way we think about contemporary
environments. Environments, be they social, ecological, or otherwise, must be understood
in part as products of intergenerational forces. It is an age-old adage; you cannot choose
your family. Each of us is born into an environment that we inherit from our parents,
an environment that is constantly shaped by larger historical, political-economic forces
that echo across generations. Just as early work in biocultural anthropology and political
ecology complicated our understanding of nature by demonstrating that the “natural”
is often a product of human action (Hvalkof and Escobar 1998; Neumann 2005), so
must biocultural scholars reconsider contemporary environments as a form of biosocial
inheritance. Certainly, individuals have the capacity to modify, manipulate, or seek out
new social and ecological environments, but even when all social ties are cut, biosocial
mechanisms cause us to carry our inheritance with us wherever we go. Furthermore, the
tendency for perpetuation and even exacerbation of health disparities across generations
can be strong, making their examination of the utmost importance to the study of health
disparities.
Finally, rather than being bound by the limitations, semantic and historical baggage of
notions of adaptation, pathology, and evolution, biosocial inheritance provides a greater
context for physiological and cultural characteristics that can hinder or aid individuals and
groups across a changing environment or throughout the life course. Since cultural and
biological “adaptations” have been shown to bring about new challenges, creating new
problems that must be adapted to (McDade and Nyberg 2010;Schell1997), it is perhaps
more useful to see them as a part of a larger, longer picture. When we contextualize them
in this way, questions of adaptability may become less relevant, with focus on context and
outcome rather than evaluation coming to the fore. Each of us carries with us generations
of biosocial history. While we often know the names and stories of our grandparents and
great grandparents, we rarely consider the ways in which our lives and bodies have been
directly and indirectly shaped by theirs. The consideration of biosocial inheritance helps
us to incorporate this intergenerational perspective as well as creating an opportunity for
the further integration of historic, cultural, and political economic analysis into the work
of biocultural anthropology.
204 Annals of Anthropological Practice 38.2/Biosocial Inheritance
NOTE
Acknowledgement. The authors would like to acknowledge Ruby Fried, Aaron Miller, Jared Bragg, Stephanie
Levy, and Kim McCabe for their helpful comments on early drafts and throughout the conceptualization of
this article. We would also like to thank our two anonymous reviewers for their helpful comments.
1. One step further would be the understanding that who procreates with whom, who is able to comingle
their genetic material to be passed on to the next generation, is largely influenced by both social and physical
geographies and thus, even our DNA can be considered largely biosocial.
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