The Biosocial Approach to Human Development, Behavior, and Health Across the Life Course Kathleen Mullan Harris and T homas W. M cDade

Social, cultural, economic, and biological fac- Bringing Together the tors are widely recognized as critical determi- Biological and the Social nants of well-­being across the life course. Yet The term biosocial is widely used in the social an integrative understanding of the multilevel sciences, but rarely defined. Perhaps its mean- biosocial pathways linking society, , ing is self-­evident. And though the term has health, and socioeconomic attainment remains appeared in the scientific literature for more elusive. The objective of this issue is to show- than fifty years, approaches and applications case research that integrates theory, data, and in biosocial research have shifted qualitatively methods from the social and biological sci- over the past fifteen years.1 In this section, we ences to advance our understanding of social discuss these developments and the synergies and biological processes that contribute to, or afforded by integrating perspectives from the derive from, social stratification across the life social and biological sciences. course. In this introduction, we describe the We definebiosocial as a broad concept refer- state of current research and discuss both the encing the dynamic, bidirectional interactions­ motivation for and relevant concepts underly- between biological phenomena and social rela- ing a biosocial perspective. We review the tionships and contexts, which constitute pro- themes and research contributions in this is- cesses of human development over the life sue, and chart a course forward for understand- course. It is difficult, if not impossible, to rep- ing biosocial pathways of well-­being across the resent the complexities of these biosocial dy- life course. namics in two dimensions, but we attempt to

Kathleen Mullan Harris is James E. Haar Distinguished Professor of Sociology and faculty fellow of the Caro- lina Population Center at the University of North Carolina at Chapel Hill. She is also a member of the National Academy of Sciences and director of the National Longitudinal Study of Adolescent to Adult Health (Add Health). Thomas W. McDade is Carlos Montezuma Professor of Anthropology and faculty fellow of the Institute for Policy Research at Northwestern University. He is also a senior fellow in the Child and Brain Development Pro- gram of the Canadian Institute for Advanced Research.

© 2018 Russell Sage Foundation. Harris, Kathleen Mullan, and Thomas W. McDade. 2018. “The Biosocial Ap- proach to Human Development, Behavior, and Health Across the Life Course.” RSF: The Russell Sage Foundation Journal of the Social Sciences 4(4): 2–26. DOI: 10.7758/RSF.2018.4.4.01. Direct correspondence to: Kathleen Mullan Harris at [email protected], University of North Carolina, 206 W. Franklin St., Chapel Hill, NC 27516; and Thomas W. McDade at t-­[email protected], Northwestern University, 1810 Hinman Ave., Evanston, IL 60208. Open Access Policy: RSF: The Russell Sage Foundation Journal of the Social ­Sciences is an open access journal. This article is published under a Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License.

1. In 1969, both the Journal of Biosocial Science and Social Biology began publishing. In 2008, Social Biology was renamed to Biodemography and Social Biology, the journal of the Society for Biodemography and Social Biology. the biosocial approach to human development 3

Figure 1. Conceptual Model of Biosocial Dynamics Across the Life Course

Political economy

Social structure and stratification

Neighborhoods and schools

Interpersonal relationships

Brain and Body gestation infancy childhood adolescence adulthood older adulthood

Biochemical messengers

Organs and tissues

Physiological and neurological systems

Genome and epigenome

Source: Authors’ compilation. do so in figure 1, which builds on prior efforts and multidimensional, and are illustrated by to highlight the multilevel domains and path- the relationships and interactions among indi- ways of particular importance in biosocial ap- viduals living in groups and within social con- proaches to health and social inequality (Kuh texts (families, neighborhoods, schools) who and Ben-Shlomo­ 2004; Glass and McAtee 2006). share the norms, institutions, and hierarchies The top boxes represent the set of nested and that structure them. The social realm can also interacting social contexts “outside” the body include aspects of the physical environment of that affect the developing brain and body of an relevance to biology (such as exposure to envi- individual throughout all stages of the life ronmental contaminants, public space for rec- course. Similarly, the bottom boxes represent reation) that are structured by social relations the nested and interacting levels of biological and hierarchies. organization “inside” the brain and body that A biosocial perspective, therefore, draws on respond to, and shape, social worlds. What con- models and methods from the biological, med- stitutes biological can be characterized as pro- ical, behavioral, and social sciences. It concep- cesses and structures within an individual that tualizes the biological and the social as mutu- contribute to the growth, reproduction, and ally constituting forces, and blurs boundaries maintenance of the soma from conception to between phenomena inside the body and out- death. Biology is typically organized across mul- side of the body. It implies that attempts to un- tiple levels, including the genome, molecular derstand one without the other are incomplete. interactions (such as gene expression, hormone It is a transdisciplinary approach to under- production); integrated physiological and neu- standing human development, behavior, and rological systems (such as the cardiovascular health, developed and applied by scholars that system; the sympathetic adrenal medullary often have disciplinary backgrounds in anthro- axis); organs and other tissues; and cells and pology, psychology, epidemiology, sociology, cellular processes. economics, public health, , medicine, Social phenomena are similarly complex and demography.

rsf: the russell sage foundation journal of the social sciences 4 biosocial path ways across the life course

Ongoing calls for a more integrative, multi-­ to biology has the potential to illuminate mech- method, multilevel interdisciplinary approach anisms through which socioeconomic, demo- to research on human development, health, graphic, and psychosocial factors shape human and social inequality underscore the impor- development and health within the context of tance and potential contribution of a biosocial everyday life. perspective (Halfon and Hochstein 2002; Harris The importance of context to human biol- 2010; Weinstein, Vaupel, and Wachter 2007). ogy is evident across multiple time dimensions The recent expansion of methodological op- (Lasker 1969). In the short term, homeostasis tions for collecting biological samples in non- and allostasis—processes of adaptation to clinical settings has facilitated this effort, and changes in current or anticipated environ- innovative biological measures are increasingly ments (McEwen 1998; Sterling and Ayer 1988)— being incorporated into social science research facilitate physiological or behavioral responses designs and data collection efforts. A new gen- to the shifting demands and opportunities of eration of biosocial research is poised to bridge local environments. For example, a perceived the gap between community-­ and clinic-­based danger or social threat increases the produc- approaches to understanding the dynamic in- tion of cortisol, a hormone that plays a central terplay of biology and social context across the role in mobilizing the body’s response to stress. life course. When the threat is removed, cortisol produc- tion returns to baseline (Gruenewald et al. Integrating Biology into the Social 2004). But repeat, or chronic, exposure to ad- and Behavioral Sciences verse environmental conditions can reset regu- Why should social and behavioral scientists latory set points, resulting in “wear and tear” care about biology? Although we recognize that on key physiological systems (Seeman et al. most, if not all, social and economic outcomes 2001). Lower socioeconomic status—a source have some biological component, social scien- of chronic stress—is associated with high cor- tists—with a few notable exceptions—have gen- tisol in the evening and with a flatter rhythm erally not considered biological processes with of production across the day relative to the nor- specificity or depth. This position does not al- mative pattern of declining cortisol production ways derive from theoretical or epistemological over the day to low levels in the evening (Cohen, stances, and is often due to gaps in data, con- Schwartz, et al. 2006; DeSantis et al. 2007). Lon- straints of training and motivational structures ger term effects of environments on biological that are set within disciplinary frameworks, systems emerge from critical or sensitive peri- and logistical challenges associated with col- ods of development, when exposures can have lecting biological measures in nonclinical set- disproportionate, enduring effects on biologi- tings. Many of these gaps are narrowing. cal structure and function. Continuing with the Putting the bio in biosocial has the potential example of cortisol, individuals born with a to make important contributions to the social lower birth weight have elevated cortisol in and behavioral sciences for several reasons. adulthood (Phillips et al. 2000), pointing to- First, humans are biological creatures, embed- ward a biological mechanism through which ded in families, social networks, communities, lower socioeconomic status (a strong predictor and cultures. Context matters to human biol- of lower birth weight) may affect health within ogy, and engagement with biological concepts and across generations. and measures reflects this reality. This is espe- By getting “under the skin,” biological mea- cially clear in the case of human health, where sures provide direct, objective information on the importance of social determinants is well pathophysiological processes that contribute established and widely known (Adler et al. 1994; to the emergence of disease, before clinically Glass and McAtee 2006; Link and Phelan 1995), diagnosable disease is evident. For example, and where social impacts on underlying phys- relative levels of blood pressure—a robust in- iological processes are apparent and increas- dicator of future risk of cardiovascular dis- ingly elaborated (Uchino, Cacioppo, and ease—tend to track from childhood into adult- Kiecolt-­Glaser 1996; Yang et al. 2016). Attention hood (Berenson et al. 1995; Li et al. 2004).

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Although measuring blood pressure in child- adverse effects on cognitive development and hood or young adulthood will reveal few clini- adult educational attainment (Conley and Ben- cal cases of hypertension, it will identify indi- nett 2000; Figlio et al. 2014). Education level is viduals most at risk for the future development also a partial function of inherited genotype, of cardiovascular disease and early death and common genetic factors can account for (Nguyen et al. 2011). Biological measures there- some of the well-­established association be- fore enhance our understanding of how social tween education and health (Boardman, environments influence pre-­disease pathways Domingue, and Daw 2015; Okbay et al. 2016). and provide opportunities for intervention Biological processes, therefore, influence indi- prior to the emergence of clinical disease. vidual life course trajectories, shape social and Attention to biology can also identify which educational attainments, and inform selection aspects of social and physical environments are into social and physical environments that can most detrimental to health and socioeconomic feed back onto biological processes. When well-being,­ as well as point toward resiliency scholars do not consider how biological mech- and protective factors that buffer groups of in- anisms shape developmental outcomes, or in- dividuals from the effects of adverse environ- teract with social environments to influence ments. The concept of embodiment has been social stratification across the life course, mod- invoked repeatedly in the social sciences to un- els may be incomplete or misspecified, param- derscore the social and political nature of the eter estimates of environmental effects over- human body and its responsiveness to social stated, and results biased. and cultural context (Gravlee 2009; Krieger A biosocial perspective is also important for 2005; Scheper-Hughes­ and Lock 1987; Seligman translating social science research into policy. 2014). The body tells stories—literally and figu- As noted, biological measures can reveal the ratively—and biological measures offer oppor- quality of social conditions, and in some cases tunities to access information that reflects the may motivate action to improve conditions to quality of social environments. Recent work on prevent disease rather than treat individuals “skin deep resilience” provides a case in point: already on the path toward disease. For exam- among African Americans from low socioeco- ple, lead screening in children can be used to nomic status (SES) backgrounds, measures of inform housing policy, where initiatives aimed self-­control predict better psychosocial out- at reducing lead exposure can prevent the de- comes, such as lower depression or lower like- velopment of costly cognitive and behavioral lihood of substance use, but worse physical disorders. Consistent evidence on the impor- health outcomes, as revealed by several biolog- tance of social relationships for biological pro- ical measures (Brody et al. 2013; Miller et al. cesses affecting health suggests that routine 2015). Biological measurement may therefore health screenings should include questions add an important dimension to our under- about the quantity and quality of individuals’ standing of health; that is, self-report,­ psycho- social connections, and physicians should be social, and biological measures may tell differ- encouraged to ask their patients about their ent stories. They may be particularly useful in relationships as part of their annual wellness settings where accurate self-­reports are espe- check-­ups (Yang et al. 2016). cially difficult to obtain, such as in research Biological measures can also add important with children, or across international settings dimensions to the evaluation of social policies. where linguistic or cultural factors may con- For example, the Moving to Opportunity dem- tribute to variation in perception, experience, onstration project was initiated in 1994 to in- or reporting (Hahn 1995; Kleinman 1986). vestigate the impact of residential contexts on Social factors affect biological process and educational attainments, income, and overall health outcomes, but the reverse is also true. well-being.­ Families in public housing were For example, lower birth weight—a biological randomized into an experimental condition variable reflecting the quality of the prenatal that subsidized their move into a low-­poverty environment, which is in turn shaped by ge- neighborhood; controls were not offered new netic, developmental, and social factors—has assistance. The intervention had limited effects

rsf: the russell sage foundation journal of the social sciences 6 biosocial path ways across the life course on education and income—the outcomes of of contextual factors on human development, primary interest when the study was de - physiology, and health. For example, in the signed—but large impacts on health: assign- early 1900s, the anthropologist Franz Boas ment to the low-poverty­ group resulted in a 13 showed that cranial form—at the time inter- percent to 19 percent reduction in obesity and preted as a fixed, inherited marker of racial 22 percent reduction in diabetes relative to the identity—was in fact malleable, and that it control group (Ludwig et al. 2011). A biosocial changed within a single generation of immi- approach to policy evaluation can identify the grants to the United States in response to en- biological processes and pre-­disease pathways vironmental influences (Boas 1912). For more that are affected by contextual factors like than forty years, social scientists and social neighborhood poverty, and point toward social epidemiologists have reported that interper- programs that improve health. Given the high sonal relationships affect health, and that so- costs of health care, this kind of information cial isolation is a risk factor for early death may add an important, but often overlooked, that is comparable in magnitude to estab - component to cost-­benefit analyses of social lished risk factors such as smoking, obesity, policies. and lack of physical activity (House, Landis, and Umberson 1988). More recently, social iso- The Importance of “Socializing” Biology lation has been associated with physiological The biosocial approach occupies an important dysregulation in all stages of human develop- and expanding space in the social and behav- ment, pointing toward biological mechanisms ioral sciences, where the emphasis has been through which social relationships affect on integrating biological concepts and meth- health (Yang et al. 2016). Socioeconomic sta- ods into research designed to address ques- tus—of keen interest to many social scien- tions of interest to social and behavioral sci- tists—is consistently associated with multiple entists (Harris 2010; Weinstein et al. 2007). Less measures of physiological function, morbidity, appreciated is the opportunity we have to col- and mortality (Adler et al. 1994; Yang et al. 2017; onize the biological sciences—as well as pub- Wolfe, Evans and Seeman 2012). lic discourse regarding the determinants of Human biology is a social biology and it is health—to have an impact on how we concep- probably up to social scientists to make this tualize and study human biology. point. Biosocial research, conducted in diverse, For the most part, research in the biological community-­based settings, encourages an epis- sciences privileges explanations “inside the temological shift that reframes human biology, body,” and is speeding down a reductionist development, and health as complexly deter- road that elaborates cellular and molecular pro- mined by multiple forces inside and outside the cesses while ignoring contextual influences body. It engages issues and processes of interest outside the body (Lewontin and Levins 2007). to biological scientists, but foregrounds social As just one example, the sequencing of the hu- and contextual factors as potentially important man genome, accomplished in 2003, was cel- contributors to variation in human physiologi- ebrated as providing “the first glimpse at our cal function and health (Stinson, Bogin, and own instruction book,” and “the possibility of O’Rourke 2012). This should be familiar ground achieving all we ever hoped for in medicine.”2 for developmental and social-behavioral­ scien- Clinical medicine also privileges reduction, tists who have long emphasized the complex seeking to isolate single, proximate factors as interplay of genes, biology, and society across causes of disease and as targets for treatment. the life course (Engel 1978; Glass and McAtee Pathogens cause infection. Tumors cause can- 2006; Gottleib 1991; Shanahan and Boardman cer (Ahn et al. 2006). 2009). With an increasingly sophisticated tool- In contrast, for more than a hundred years, kit for integrating biological measures into social scientists have documented the impact community-­based, social science research, the

2. “What They Said: Genome in Quotes,” BBC News Science/Nature, June 26, 2000, http://news.bbc.co.uk/2 /hi/science/nature/807126.stm (accessed October 4, 2017).

rsf: the russell sage foundation journal of the social sciences the biosocial approach to human development 7 time is right for a new generation of biosocial studies such as the National Longitudinal Study scholarship that enriches both the biological of Adolescent to Adult Health (Add Health), the and the social sciences and helps build stronger Health and Retirement Study, the National So- links between them. cial Life, Health, and Aging Project, and Moving to Opportunity. International studies, including Methodological Developments the Cebu Longitudinal Health and Nutrition Historically, community- ­and population-based­ Survey, the Mexican Family Life Survey, and the research in the social sciences has relied on Study on Global AGEing and Adult Health are vital records or self-­reported, survey-­based collecting tens of thousands more. In another measures of health and disease. Information example, Add Health developed its own kit for can be readily collected from large representa- the collection of buccal cell DNA in 1996 to test tive samples across a wide range of settings, for the zygosity of sampled twin pairs. Ten years but insight into biological processes is limited. later, commercial kits for saliva DNA collection In contrast, biomedical research employs in-­ (such as Oragene) are routinely used by multi- depth biological measures collected in con- ple studies to collect thousands of DNA samples trolled clinical or laboratory settings, but typi- both in the home setting and through the mail cally relies on smaller, select groups of via self-­collection. participants who are invited to participate The integration of objective measures of bi- based on preexisting criteria. Generalizability ological function and health has advanced the and external validity are limited, and social fac- biosocial perspective by directly contributing tors are generally not considered, beyond stan- to our understanding of how social, economic, dard measures of socioeconomic status or self-­ and community factors shape human biology reported health behaviors. and health, and vice versa. These methods also Methodological options for collecting and address the goal of socializing biology. By tak- generating biological data have expanded ing our methods into the community, where greatly over the past fifteen years, allowing us participants are living their daily lives, we to bridge this gap (Weinstein et al. 2007). Low-­ greatly expand the range of environmental vari- cost, field-friendly­ options for collecting blood, ation that can be evaluated in relation to bio- saliva, or urine in the home or local community logical phenomena. Contextual factors are allow investigators to gain access to physiolog- therefore brought into relief as potentially ical information from large numbers of par- ­important determinants of human physiologi- ticipants in naturalistic settings (Adam and Ku- cal function and health in ways not possible mari 2009; McDade, Williams, and Snodgrass with lab- ­or clinic-based­ research designs. Last, 2007). Developments in assay technology have these methods serve as a catalyst for productive facilitated the measurement of proteins, gene collaboration among social, life, and biomedi- transcripts, epigenetic marks, and DNA se- cal scientists. The growing availability of social quences with higher resolution in smaller and biological data in large, representative quantities of sample, at lower costs (Dedeur- samples, and the emphasis on interdisciplin- waerder et al. 2011; McDade et al. 2016). Por- ary scholarship, has laid fertile ground for the table devices and low-­cost monitors facilitate integration of complementary expertise to gen- assessment of sleep, physical function and ac- erate novel insights into the ways in which so- tivity, blood pressure, and body size and com- cial and biological processes interact in path- position (Lindau and McDade 2007; Marino et ways of human development. al. 2013). These methodological innovations have en- The Importance of the couraged wide-­scale integration of objective Life Course biological measures into social science surveys. Human development has social and biological For example, dried blood spots—drops of whole determinants and intergenerational linkages blood collected from a simple finger stick— beginning in utero and continuing throughout have been collected from more than thirty-five­ all stages of the human life span (Hertzman thousand participants in the United States in and Boyce 2010). Despite a consensus that early

rsf: the russell sage foundation journal of the social sciences 8 biosocial path ways across the life course life conditions and childhood experiences mat- 1), aging research has led the field in study de- ter for subsequent social and biological de­ signs incorporating inputs across social and velopment in adolescence, early adulthood, biological levels of analysis. Understandably, mid-adulthood­ and old age, most social and this line of research focuses on phenomena biomedical research does not capture the ways such as disability, illness and disease, and lon- in which developmental processes are linked gevity and mortality. The biosocial approach and interrelated across phases of human life, in aging research, however, has not been well-­ nor does it capture the dynamic interactions informed by a life course perspective. of social and biological forces that underlie de- For a long time, aging research used self-­ velopment across time and space. Part of this reported health and behavioral information research gap is due to a lack of longitudinal, and cross-sectional­ designs to study, for exam- multilevel life course data and intergenera- ple, the age distribution of the prevalence of tional study designs, and part to disciplinary illness and chronic disease (see, for example, approaches designed to identify disciplinary-­ National Center for Health Statistics 2016; specific determinants of social, behavioral, or Ward, Schiller and Goodman 2014), activities health outcomes at a point in time. of daily living and instrumental activities of A life course perspective is essential in bio- daily living designed to assess whether older social research because outcomes at any point adults can independently care for themselves reflect the product of prior interactions be- (Freedman and Spillman 2014), and family and tween social and biological forces that occur social relationships among the elderly (Waite across human development (Shanahan, Hofer, and Das 2010). Demographic studies also use and Shanahan 2003). Life phases and social cross-sectional­ data but dynamic life table roles are often intimately tied to biological methodology to document onset and years of events or trajectories (George 2009). For exam- disability and chronic illness and to estimate ple, a woman’s first birth marks her transition mortality risks and life expectancy based on into parenthood just as menopause defines the point-in-­ ­time rates of these respective events end of the reproductive phase of her life. Al- (Crimmins, Zhang, and Saito 2016). Perhaps the though a woman can biologically become a most influential contribution of aging research mother when she reaches puberty in adoles- with implications for the biosocial paradigm is cence, most young people in the United States long-­standing evidence of large and persistent delay parenthood until well after puberty to social gradients in health and mortality (Adler continue social and emotional maturation and et al. 1994; Wolfe, Evans, and Seeman 2012; Mar- invest in human capital and career develop- mot and Wilkinson 2005). Still, this research ment before becoming a parent. Thus, social remains primarily cross-sectional,­ document- and biological forces jointly shape transitions ing how SES is associated with aging-­related between roles and patterns of continuity and outcomes at a given point. discontinuity that extend across the phases of With the advent of nationally representative life. Biosocial approaches, therefore, require longitudinal aging studies in the 1990s, such the researcher to dynamically assess both bio- as the Health and Retirement Study (HRS) and logical and social features of the developing the National Social Life, Health and Aging Proj- person and their changing social context ect (NSHAP), and longitudinal community-­ through time and across generations to achieve based aging studies, such as Framingham, Ath- a full understanding of the determinants of so- erosclerosis Risk in Communities Study (ARIC) cial and physical well-­being. and Reasons for Geographic and Racial Differ- ences in Stroke (REGARDS), a life course design Biosocial Processes Across the Life Course could now be applied to understand how previ- Within social and behavioral sciences, research ous social, behavioral, and environmental con- on aging has been at the forefront of biosocial ditions were related to health and disease out- approaches. Because aging integrates forces in- comes among older adults. In addition, as new side the body and outside the body to shape survey field methods for measuring objective function and health in older adulthood (figure health outcomes were incorporated into many

rsf: the russell sage foundation journal of the social sciences the biosocial approach to human development 9 of these ongoing longitudinal aging studies, jor impact on epidemiologic research on the the ability to understand biological mecha- determinants of adult disease risk, with a par- nisms and markers of health and disease fur- ticular emphasis on cardiovascular diseases ther enhanced longitudinal life course data for and the physiological processes through which biosocial research. However, the life phase ex- they are influenced by early life nutritional en- amined in these studies is still limited to older vironments (Gluckman et al. 2008; Wadhwa et adults because observations begin when indi- al. 2009, Smith and Ryckman 2015; Kuh and viduals are forty-­five (ARIC) or fifty (HRS) or Ben-­Shlomo 2004). Numerous studies have fifty-­five (NSHAP), thus missing earlier life linked uterine, birth, and childhood exposures stages that certainly bear on aging processes to adult physical health and disease (Bengtsson that arguably begin at birth. The social gradi- and Broström 2009; Cameron and Demerath ent in health and mortality, for example, can 2002; Crimmins and Finch 2006; Smith and now be studied by examining how SES trajec- Ryckman 2015). Demographic and social re- tories beginning at age fifty influence the onset search on the “long arm of childhood” has also of disease or death. demonstrated the value of a life course perspec- From a life course perspective, this research tive when early life circumstances are both di- design has four major limitations. First, SES rectly and indirectly associated with health out- does not change much beyond age fifty because comes that emerge decades later in adulthood the components of socioeconomic status—ed- (Blackwell, Hayward, and Crimmins 2001; Case ucation, occupation, income and wealth—are and Paxson 2010; Elo and Preston 1992; Hay- typically developed earlier in the life course ward and Gorman 2004; Preston, Hill, and during adolescence, early, and mid-­adulthood Drevenstedt 1998). and vary little in old age. Second, the early and Most of this research, however, links early midlife biological precursors to disease onset life conditions with physiological processes or and death are not observed in these studies. chronic disease outcomes in later adulthood Relatedly, the lack of biological data from ear- with cross-­sectional research designs, paying lier ages precludes opportunities to consider limited attention to what happens in between— how early life course health and biological pro- during the majority of the early life course from cesses shape SES attainments in adulthood. later childhood to adulthood. From a biosocial Fourth, studies that begin observation at older perspective, this means we are missing a lot, ages miss those who have died, typically the especially the social processes and contexts more disadvantaged individuals who have been that structure, mediate, and moderate biology more exposed to earlier life trauma and ill- over the life course. nesses and have fewer resources for health Adolescence and the transition to adult- care, thus biasing the SES-health­ relationship. hood, for example, are life stages when young One solution to this lack of prior life course people first begin to choose their environ- information that bears directly on older age so- ments, health behaviors, habits, and future life- cial and physical well-being­ is to collect retro- styles (Harris 2010). These life course choices spective information about status earlier in life, shape or alter social and biological pathways­ enabling a modified life course perspective for originating in childhood and moving into biosocial research. Here aging studies have fo- adulthood. Investments in human capital be- cused on retrospective reports of SES and gin in early childhood, but intensify and be- health conditions at birth and during early come more self-­directed during adolescence childhood. and the transition to adulthood. Profound and Along these lines, research on the develop- protracted physical, biological, and neurologi- mental origins of health and disease (DOHaD) cal changes linked to puberty occur throughout has exploded, following early biomedical re- adolescence and early adulthood. Hormonal search by Barker documenting significant links changes prompt a literal remodeling of cortical between birth weight and later cardiovascular and limbic circuits in the brain that were previ- disease risk within cohorts (Barker 1997, 1998, ously organized in the perinatal period and 2006). The life course approach has had a ma- that, in combination with adolescent social ex-

rsf: the russell sage foundation journal of the social sciences 10 biosocial path ways across the life course periences and contexts, affect general cogni- cial well-­being in that life stage and beyond. tion, decision making, and behavior into adult- Biosocial research cannot examine social and hood (Sisk and Zehr 2005). Behavioral changes biological forces in all life stages in one project and exploration in diet, exercise, sleep pat- or with one dataset, but should contribute terns, substance use, sexual activity, and knowledge about how social and biological ­aggression during adolescence and young phenomena operate in distinct life stages and adulthood further shape social and biological are linked to health and social inequities in pathways into adulthood (Hubert et al. 1987). subsequent stages across the life course. Al- DOHaD research, and other life course perspec- though we have made some progress docu- tives, ideally should include the contributions menting the association between early life of these critical developmental stages to illu- ­conditions and late life health and disease out- minate health and well-­being pathways into comes (such as lower birth weight and in- adulthood. creased cardiovascular disease risk), we need Although neurological development and to move beyond cross-sectional­ designs to un- change slows down as young people settle into cover the underlying life course processes that adulthood, these demographically dense years explain these associations. Both the intergen- bring new stresses to daily life as young adults erational precursors that lead to these condi- juggle the multiple interrelated life domains tions and the subsequent intragenerational life of relationships, schooling, work, and family. course pathways such conditions initiate are Stress processes are perhaps the most com- yet to be explored. However, a small but grow- monly considered biological mechanisms ing literature links these kinds of early life through which the social environment gets un- health factors to later social attainments, but der the skin to affect health and development more research on health and biology as under- in biosocial models, as described earlier in the lying factors in social stratification processes case of the stress hormone cortisol (Gruene­ is needed. wald et al. 2004; McEwen and Lasley 2002). Mid- dle adulthood may usher in greater life course Biosocial Study Designs of Health and Social stability and security in socioeconomic status, Inequality Across the Life Course work, and family, but only for certain sub- Here we describe two general life course orien- groups of the population. Middle adulthood tations for understanding how biological phe- has become more dynamic and demanding in nomena are related to social and economic sta- contemporary U.S. society—high divorce and tus and opportunities in direct, indirect, and repartnering rates, greater dependency from reciprocal ways. One orientation examines how both the child and parent generations, and un- social stratification processes across the life certain work schedules, low wages, and a lack course are related to subsequent health out- of employment benefits for those with low ed- comes in different life stages. This orientation ucation or few job skills. Relative to other life stems primarily from the large literature on the stages, middle-aged­ adults are highly embed- social gradients of health discussed (or more ded in social relationships with aging parents, commonly, the social determinants of health), children, the parents of children’s friends, but with a life course perspective. Social strat- neighbors, work colleagues and within com- ification is both an inter- ­and intragenerational munity institutions. These relationships are process. At birth, we enter a social hierarchy important social mechanisms that can buffer tied to parental SES that determines access to (through social support) or exacerbate (through material and social resources for both physical strain and conflict) the daily stresses of middle and social development. The developing indi- adulthood (Yang, Schorpp, and Harris 2014; vidual then faces constraints and opportunities Yang et al. 2016). in each life stage that determine social and eco- All life course stages have unique social and nomic status across time. Inter- ­and intragen- biological forces that determine life-­long hu- erational social stratification processes have man development and that operate indepen- both direct and indirect effects on health across dently and jointly to influence physical and so- the life course. Understanding the social and

rsf: the russell sage foundation journal of the social sciences the biosocial approach to human development 11 biological mechanisms for how social stratifi- ular developmental periods may operate to in- cation processes get under the skin to influence crease health risk in subsequent life stages. health is at the heart of this conceptual orien- The top model illustrates sensitive period tation of biosocial research. timing effects in which exposures during sensi- Ideally, social stratification is measured lon- tive periods of development have stronger ef- gitudinally, as a life course process, conceived fects on health outcomes than they would at as social exposures that can be positive (sup- other life stages (Hayward and Gorman 2004; portive parenting behaviors, college education) Gluckman et al. 2008; Cohen, Janicki-­Deverts or negative (childhood poverty, neighborhood et al. 2010). Sensitive period effects operate disadvantage). Exposure to both beneficial and through a biological embedding mechanism adverse experiences over the life course will whereby social exposures during sensitive win- vary for each individual and constitute a unique dows of development have the potential to in- social stratification trajectory. Biological out- duce structural and functional changes to the comes are conceived of as the consequence of developing individual through biological pro- exposure in social stratification trajectories and gramming that cannot be reversed regardless can be measured at a point in time in a par- of intervening experience. Thus, the dark shad- ticular life stage or over time as biological and owed line represents a direct effect of exposure developmental change. Importantly, social in the earlier stage of development with no in- stratification trajectories represent social pro- direct effects and no direct effects of subse- cesses that enable the biosocial researcher to quent social disadvantage on later life health. explore fundamental life course mechanisms This life course model posits that the effect of involving the timing, duration, and intensity of the sensitive period exposure is typically latent beneficial or adverse social exposures that oc- in that its impact on health outcomes may cur in different and across phases of life and not appear until later life stages, often decades that affect health and development in subse- later. quent life stages. The ability to measure the Duration effects of social stratification pro- timing, duration, and intensity of social expo- cesses can be explored through the accumula- sures across the life course allows for testing tion life course model (middle model of figure life course models for how social experiences 2), which emphasizes the role of persistent that occur outside the body are linked to bio- ­advantage or disadvantage over time—both logical mechanisms inside the body that affect in specific life stages and over life stages—on health and well-being.­ health and development. The effects of multi- The stress response framework is the most ple exposures over the life course are both ad- prominent biosocial paradigm to explicate how ditive and interactive and combine in synergis- trajectories of social structural inequalities are tic ways to influence biological mechanisms associated with greater exposure to stress and and, in turn, health and development out- its biological and health-related­ manifestations comes. Cumulative effects can either be mul- (Pearlin 1989; Aneshensel 1992; McEwen 1998; tiple exposures to a recurrent stressor (such as McEwen and Lasley 2002). When social expo- chronic poverty) or a series of exposures to dif- sures are intense, or the magnitude of struc- ferent social environments or life experiences. tural disadvantage is high (depth of poverty, For example, poverty experienced only during multiple disadvantages of poor neighbor- childhood is not as detrimental as poverty dur- hoods), stress response is chronic and biologi- ing childhood, adolescence, and the transition cal dysregulation is greater, resulting in poor to adulthood on subsequent adult health. health and developmental outcomes. The life A third life course model that might explain stage timing of social exposures, however, may how social stratification processes are related differ for both the biological mechanisms and to health outcomes is the pathway model, subsequent health outcomes associated with which tracks how social exposures in one life stress exposure. Figure 2 provides an illustra- stage influence the probability of related social tion of various life course models that describe exposures in subsequent stages. Also known as how exposure to social disadvantage in partic- the chains of risk model, it emphasizes pathway

rsf: the russell sage foundation journal of the social sciences 12 biosocial path ways across the life course

Figure 2. Life Course Models of Social Disadvantage Trajectories and Health

Sensitive Period

Social Social Health risk disadvantage disadvantage

Age

Accumulation

Social Social Health risk disadvantage disadvantage

Age

Pathway

Social Social Health risk disadvantage disadvantage

Age Source: Authors’ compilation. effects whereby early experiences set in motion explained by the SES pathway where early life a chain of events that put individuals on paths SES determines adult SES, which in turn, is a differentiated by types and levels of stress ex- more proximate and important predictor of posures to social and biological factors (Mar- adult health and disease (Yang et al. 2017). mot et al. 2001; Pudrovska and Anikputa 2014). A second orientation for understanding bio- This model elaborates on the ways in which social pathways in well-­being across the life inter-­ and intragenerational social stratification course is consideration of the role of biology pathways are linked across the life course. For or health in social stratification processes (Pal- example, the connection between early life con- loni 2006). In this orientation, biological mech- ditions and adult health and disease may be anisms and health trajectories are important

rsf: the russell sage foundation journal of the social sciences the biosocial approach to human development 13

Figure 3. Role of Health in Social Stratification Processes

Adolescence Young adulthood Adulthood

Obese Obese Social and economic status Not obese Not obese

Mechanisms

Source: Authors’ compilation. contributors to subsequent socioeconomic out- illuminate the often missing, underlying role comes and attainment. For example, economic of biology in social stratification processes. Un- research has focused on how childhood health derstanding whether, when, and how biological influences human capital and labor force out- processes matter for social and economic out- comes in adulthood, including educational at- comes across the life course will help identify tainment, labor force participation, income, when biomedical interventions might be most and occupation (Case, Lubotsky, and Paxson effective for reducing social inequality. These 2002; Currie and Stabile 2003; Currie and models are not mutually exclusive and in real- Moretti 2007; Case, Fertig, and Paxson 2005; ity coexist (see Hallqvist et al. 2004). Most im- Almond and Currie 2011). Figure 3 illustrates portant, they provide a framework for biosocial the case of how early life course health can in- research made possible by longitudinal data fluence later socioeconomic status. In particu- and study designs that enable researchers to lar, life course trajectories of obesity during identify the social and biological processes that adolescence and into young adulthood contrib- operate in pathways of well-­being across the ute to social stratification outcomes in adult- life span. The life course perspective articulates hood. Research findings indicate that those the longitudinal and multidimensional of so- who become obese in early adolescence and cial and biological forces that operate in all life remain obese for a longer period of time have stages and underlie human development lower levels of education, marriage, wages, across time, emphasizing the need to concep- household income, employment, assets, and tualize social conditions and biological mech- subjective social status (Cawley 2004; Han, Nor- anisms as dynamic constructs that unfold ton, and Stearns 2009; Harris and Lee 2011; across time, beginning in early life and continu- Glass, Haas, and Reither 2010). These life ing into young adulthood, midlife, and old age. course effects of obesity operate through such mechanisms as low self-­esteem, social isola- Social Genomics tion, societal views of attractiveness and lost Consensus is now widespread that social, be- work days, illustrating the biosocial connec- havioral, and health outcomes are a function tions. Similar effects are found for chronic of both nature and nurture, and are best un- health conditions and diabetes during adoles- derstood in a life course context. Even in the cence and young adulthood, which truncate age of genome mapping, research on the im- educational trajectories and reduce the stabil- pact of genetic variance alone has limited ex- ity of work (Fletcher and Richards 2012). planatory power, and is often of less interest to In this orientation, life course models will social and behavioral scientists given that in-

rsf: the russell sage foundation journal of the social sciences 14 biosocial path ways across the life course dividual DNA sequence is fixed and not subject spective, GxE research has come under signifi- to intervention. Instead, social and behavioral cant criticism for a poor record of replication, scientists have been drawn to understanding lack of statistical power for GxE associations, gene-­environment interplay, or how environ- and the endogenous nature of most measures mental and genetic factors interact over time of E (Boardman, Daw, and Freese 2013; Conley to affect social, behavioral, and health out- 2016; Freese and Shostak 2009; Charney and comes, along the lines of figure 1. Because such English 2012; North and Martin 2008). Still, outcomes represent the cumulative history of promising GxE research is on the horizon us- a person’s social experiences as they combine ing natural or quasi-­experimental designs and with genetic makeup, gene-­environment inter- larger samples afforded through genetic con- play reflects life course processes. Although ap- sortia (see, for example, Boardman et al. 2012; preciation is widespread that the links between Okbay et al. 2016; Schmitz and Conley 2016, genes and behavioral outcomes, for example, 2017; Rietveld et al. 2013). are conditioned by the social environment, Research exploring gene-­environment cor- consideration of the dynamic features, of social relations is especially valuable to social and be- environments and life experiences as processes havioral sciences because it confronts the wor- occurring across the life course is not. To date, risome endogeneity problem of estimated most studies of gene-­environment interplay are environmental effects being due to unobserved cross-­sectional or use longitudinal data with- heterogeneity (genetic factors) associated with out explicit modeling of life course features, the selection of one’s environment. Gene-­ including pathways, transitions, trajectories, environment correlation (rGE) refers to pro- durations, or timing (Conley 2016; Shanahan cesses by which genetic factors are associated and Boardman 2009). with features of the environments in which in- Nevertheless, substantial social science re- dividuals live their lives (friendships, peer search examines gene-­environment interplay groups, romantic relationships, schools, neigh- focusing on two general approaches: gene-­ borhoods, work environments, and so on). environment interactions (GxE) and gene-­ Sorting out and controlling for genetic variance environment correlations (rGE). GxE research in selection of these environments enables so- has captured social science attention by elabo- cial and behavioral scientists to isolate the rating on processes by which the effect of ge- causal impacts of social environments on so- netic factors on a social or biological outcome cial, behavioral, and health outcomes. In sum, is conditioned by environmental factors and perhaps the main impact of the evidence on vice versa (Boardman, Daw, and Freese 2013; gene-­environment interplay has been to dispel Hutter et al. 2013). Provocative findings have notions of the nature-nurture­ dichotomy and been published, for example, showing that ge- build consensus on the need for integrative netic effects on children’s cognition are damp- models of genetic and social factors to better ened in low SES environments, genetic propen- understand human development and health. sities for adolescent substance use are But two general weaknesses remain: the G (gen- enhanced or suppressed according to the prev- otype) is still fixed and therefore unresponsive alence of substance use in the adolescents’ to social change; and the dynamics of life schools, stressful life events increase the risk course changes and their biological interac- of depression depending on one’s genetic pro- tions with changing social environments and file for processing neurotransmitters, and ado- experiences over time have not been exploited lescents are more genetically similar to their in either GxE or rGE research. friends in more highly structured and segre- Human social genomics, on the other hand, gated environments (Boardman, Domingue, is an emerging field of research that examines and Fletcher 2012; Caspi et al. 2003; Rowe, Ja- why and how external social conditions affect cobson, and Van den Oord 1999; Daw et al. the activity of the genome (Slavich and Cole 2013). Although such findings highlight the im- 2013; Boyce and Kobor 2015). Social genomics portant role of the social environment in includes the study of gene expression (tran- ­genetic processes, especially from a policy per- scriptome) and (epigenome). It

rsf: the russell sage foundation journal of the social sciences the biosocial approach to human development 15 emerges from the scientific understanding that sion—a key reason why epigenetics has cap- while the gene sequences we inherit from our tured the attention of many social scientists. parents are fixed, the expression of these genes Methylation of DNA has been the major focus is shaped by forces “outside” the body. The of human research, and involves the binding focus of social genomics is inherently bioso- of methyl groups to cytosine residues in CpG cial as it seeks to uncover how social experi- dinucleotides (Bird 2002). At a point in an in- ences can alter gene expression and thereby dividual’s life course, analysis of DNA methyla- affect physiological function and social and tion may reveal how and which genes have been behavioral outcomes (Cole 2014; Hertzman modified in response to the cumulative life 2012). Indeed, human social genomics re - course environmental, behavioral, and biolog- search is demonstrating that certain genes ical trajectories of that individual. Thus, epi- can be “turned on” or “turned off” by different genetics specifies a life course biosocial process social-­environmental conditions, and in some that entails the dynamic interactions and feed- cases these social exposures can affect the ac- back loops of social and genetic phenomena tivity of hundreds of genes in a coordinated both inter- ­and intragenerationally over the life manner. course (Boyce and Kobor 2015). For example, This line of research provides new opportu- epigenetic patterns have been shown to be al- nities for understanding how social and genetic tered by a range of environmental conditions factors interact to shape complex biological such as diet, tobacco smoking, exercise, and and social pathways of well-­being. Indeed, it exposure to chemicals (Christensen et al. 2009; has the potential to reframe our understanding Grönniger et al. 2010; Langevin et al. 2011). of the genome as a dynamic substrate that in- More broadly, measures of socioeconomic and corporates information from the environment psychosocial adversity in childhood have been over developmental time, rather than the pre- linked to patterns of DNA methylation later in vailing view of the genome as static sequences life (Essex et al. 2013; Needham et al. 2015). Epi- of DNA that are fixed at conception. Current genetic patterns have also been shown to affect social genomics research has examined, for ex- physical traits and appearance, behavior, and ample, how social processes, such as social sta- health outcomes (IHEC 2013). Thus, environ- tus, social supports or isolation, social capital, mental variation may routinely change epigen- early life adversity, exposure to toxicants and etic patterns, and those epigenetic patterns microbes, and health behaviors, alter the ex- may in turn influence developmental outcomes pression of hundreds of human genes (such as over time. suppression of antiviral and antibody-­related One of the more provocative—but also con- genes and stimulation of pro-­inflammatory troversial—findings for social scientists is that genes) to affect human development and health environmentally triggered behavior or biologi- over many years (see, for example, Cole 2013, cal change might be transmitted across gen- 2014; Murphy et al. 2012; Miller et al. 2009; Fry erations through epigenetic mechanisms and et al. 2012; McDade et al. 2017). without the involvement of DNA sequence The term epigenetics—first used by Wadding- (Jablonka and Lamb 2015; Thayer and Kuzawa ton in the 1940s (Jablonka and Lamb 2002)— 2011). Fascinating early evidence on the inher- literally means above or on top of genetics, and itance of epigenetic marks comes from re- refers to chemical modifications to DNA and search done on mice. Human evidence is much its packaging that change the accessibility of more difficult to establish. Studies of humans gene regions to transcription factors, and whose ancestors survived through periods of thereby affect the level of transcription. These starvation in Sweden and the Netherlands sug- modifications alter the physical structure of gest that the effects of famine on epigenetics DNA in ways that are relatively stable and con- and development can pass through at least served with cell replication. Therefore, epi- three generations (Heijmans et al. 2008; Tobi genetics represents a biological mechanism et al. 2009). Nutrient deprivation in a recent through which the body “remembers” prior en- ancestor seems to prime the body for diabetes vironmental exposures to shape gene expres- and cardiovascular problems, a biological re-

rsf: the russell sage foundation journal of the social sciences 16 biosocial path ways across the life course sponse that may have evolved to mitigate the Disadvantage, Discrimination, and Health effects of future famines. The findings on in- The impact of social adversity on human wel- tergenerational epigenetic inheritance could fare is of long-­standing concern to social sci- have far-reaching­ significance. Much social sci- entists. A biosocial perspective addresses ques- ence research documents how parental char- tions regarding the health impacts of adversity, acteristics, such as lifestyle, behaviors, and liv- and the biological mechanisms through which ing habits, influence children’s well-­being. social environments “get under the skin” to Epigenetic processes may provide biological impact human development and health. This mechanisms through which lifestyles and be- aspect of biosocial research has been greatly haviors are stored and transmitted to children advanced by recent methodological develop- and their children’s children, who do not have ments that have facilitated the collection of any direct environmental exposure to these life- objective biological data in nonclinical, com- styles or behaviors. munity-­ and population-­based settings. All Social genomics is an exciting area for future three articles in this section showcase the value biosocial research that emphasizes the instru- of these kinds of measures for advancing our mental role of the social environment in alter- understanding of how social adversity affects ing how genes are expressed to affect behavior, health. biology, and social and health outcomes. The Douglas Massey and his colleagues build on more we understand how the social environ- a long tradition of scholarship on neighbor- ment regulates genes that affect health and so- hood effects, with a particular emphasis on cial stratification processes, the more potential residential segregation and concentrated pov- we have for intervening on those environmen- erty. The majority of this work has been socio- tal exposures to reduce health and social in- logical, including some links to health but pay- equalities. The molecular models of social ge- ing little attention to biology. Massey and his nomics do require new methodological skills colleagues introduce a biosocial framework for and technical capacities for working with these linking spatially concentrated disadvantage at data, over and above the application of stan- the geographic level with an individual-­level dard social science methods used in GxE and biological measure of cellular aging (telomere rGE analysis. Interdisciplinary training, how- length) to reveal mechanisms through which ever, is already coming online to equip social social structure contributes to race-­based dif- scientists with these skills through summer ferences in morbidity and mortality in the boot camps and graduate training programs United States. Aside from contributing to the around the country. The potential for under- literature on neighborhood effects, the article standing these social and biological phenom- demonstrates how collaborative, interdisciplin- ena has captured the attention of the scholarly ary teams can leverage novel insights from and public worlds alike. The ability of social ­molecular biology to cast new light on long-­ genomics to fill diverse gaps in our understand- standing social science questions. ing of human development and health and to The article by Bridget Goosby and her col- provide scientific explanations of the mecha- leagues also investigates the health impact of nisms underlying our lived experiences makes social disadvantage, but at the individual, it a compelling avenue for future biosocial re- micro-­social level of analysis. Their focus on search. perceived discrimination draws on a well-­ established line of biosocial research that at- Issue Themes and Chapters tends to the appraisal of stress as a key part of Contributors to the volume represent a wide the causal pathway linking social adversity with range of disciplines, and their work advances physical health. Sleep quality and quantity are the biosocial perspective on human develop- the key outcomes in their study, based on re- ment, behavior, and health across the life cent clinical and epidemiological research course. The issue is loosely organized around demonstrating the importance of sleep for a three themes. wide range of physical and mental health out-

rsf: the russell sage foundation journal of the social sciences the biosocial approach to human development 17 comes. Like Massey and colleagues, the article actions between social and biological forces showcases the value of borrowing from biolog- that occur across human development. ical and health sciences to illuminate issues of Yang Qu and his colleagues use a biopsycho- interest to social scientists. This point is un- social approach to understand adolescent de- derscored by the counterintuitive nature of velopment among a growing U.S. ethnic minor- their results. Global ratings of discrimination ity group—Mexican American youth. They are negatively associated with sleep quantity identify adolescence as a dynamic life stage in and quality, as one might expect, but partici- which neural changes in both brain function pants slept better the night following a day and brain structure are likely associated with when they reported a discriminatory encoun- individual differences in academic and psycho- ter. This finding reveals the potential of bio- logical adjustment. They also argue that the logical or health-­related measures to provide environment becomes especially salient during novel insights into psychosocial dynamics that adolescence for Mexican American youth when might otherwise be obscured. It also highlights ethnic parents attempt to socialize children the value of measuring these dynamics at mul- about their cultural values and heritage, and tiple levels of analysis. at the same time, adolescents yearn to spend The article by Elizabeth McClure and her more time with peers and fit into adolescent colleagues provides an excellent example of social life and activities. The longitudinal re- how a biosocial approach can be used to “so- search in this article examines the independent cialize” biology. The ratio of signal-joint­ to beta and interactive effects of adolescents’ brain de- T-­cell receptor excision circles (sj/beta-­TREC velopment and their family and peer environ- ratio) is used by immunologists to measure the ment in determining educational achievement function of the thymus, an organ that produces and substance use. By integrating imaging data cells (T lymphocytes) that are essential for im- with rich social variables on cultural socializa- munity. In their study in Detroit following the tion and peer deviance, Qu and his colleagues Great Recession, they show that neighborhood-­ find important independent contributions of level measures of home foreclosure and aban- biological and psychosocial factors in youth’s donment predict lower levels of thymic func- achievement and adjustment. tion among residents. Furthermore, reduced The Margot Jackson and Susan Short article social cohesion—a product of home foreclo- uses a life course intragenerational design to sure and abandonment—predicts lower thymic examine both gender differences in physical function. With a clearly articulated conceptual health in young adulthood and the ways in model that informs their study design and an- which adolescent development and social en- alytic strategy, McClure and her colleagues vironments might explain those differences. move from the macro- ­(Great Recession) to the Documenting gender differences in objective mezzo-­ (neighborhood characteristics) to the biological markers of health (inflammation and micro- ­(thymic function) levels of analysis to immunosuppression) during young adulthood demonstrate how human biology is a social bi- is a contribution to the health disparities lit- ology. erature given most research on sex differences focuses on older aged populations. Jackson and Developmental and Intergenerational Short report strong differentials in inflamma- Processes tion and immune function that disadvantage The two articles in this section use a biosocial women in these biological systems. They ex- approach to assess the biological and social fea- plore a wealth of childhood, adolescent, and tures of the developing person and their chang- early adulthood circumstances—including de- ing social contexts through time and space to mographic, family socioeconomic, health be- provide new insights into the determinants of havior, and young adult family formation and social and physical well-being.­ The articles ex- socioeconomic attainment—as potential ex- amine outcomes in specific life stages as a func- planatory factors underlying the sex differ- tion of earlier life course exposures and inter- ences. Identifying gender disparities early in

rsf: the russell sage foundation journal of the social sciences 18 biosocial path ways across the life course the adult life course is critical to curbing their molecular data, Mills and her colleagues find growth throughout adulthood by designing in- relatively low levels of predictive power for the terventions to improve female health and re- PGSs based on the entire genome, revealing duce the disparities before chronic disease and the more predominant role of social- ­ long-­term physical damage occurs. environmental and behavioral factors in deter- mining age at first birth and number of chil- Genes and Environments over the dren ever born. Rather than speculating on how Life Course much environmental and behavioral estimates Interest in gene-­environment interplay has cap- are overstated by their confounding with ge- tured the imagination of biosocial researchers, netic effects, social scientists use these models who isolate the role of genes in relation to en- to control for this genetic confounding while vironmental influence and focus on the ways estimating the importance of social and behav- in which genes and environments operate to- ioral factors of fertility behavior. Moreover, age gether in social stratification processes across at first birth and number of children ever born the life course. The advent of new sources of are well-­established markers of social stratifi- molecular genetic data, especially genome-­ cation, illustrated by the voluminous literature wide data, and statistical tools for analyzing on teenage childbearing and family size (re- massive amounts of individual-­level genome-­ views in Furstenberg 2003; Powell et al. 2016; wide data linked to survey and biomarker in- Sweeney and Raley 2014; Wilcox and Lerman formation in large studies has opened exciting 2014). Indeed, Mills and her coauthors show new research opportunities for understanding that the PGSs are also correlated with other fer- gene-­environment interplay in biosocial mod- tility traits, such as childlessness, and are in- els of attainment and behavior. Two articles in dependent of the effects of education. this issue take advantage of an analytic ap- Benjamin Domingue and his colleagues fo- proach that combines the genetic associations cus on the role of genes in environmental se- with specific phenotypes (that is, behavioral, lection processes, or gene-­environment corre- attainment, and health outcomes) across the lation. They examine geographic clustering of entire genome using polygenic scores (PGS) PGSs for multiple phenotypes related to an- based on genome-­wide association studies thropometry, education, and physical and men- (GWAS). A PGS is a linear combination of the tal health by state of residence at different effects of genetic variants present in the entire points in the life course to explore the extent genome specific to a phenotype that can be in- to which state-­level genetic composition ex- terpreted as a single quantitative measure of plains state-level­ clustering of various pheno- genetic predisposition for that phenotype. types and how these relationships change over The article by Melinda Mills and her col- age. Domingue and his coauthors expertly dis- leagues builds on their recently published cuss the important motivation behind examin- meta-­GWAS study on human reproductive be- ing gene-­environment correlations, the mech- havior (Barban et al. 2016) by examining the anisms through which gene-­environment predictive power of two PGS discovered in that correlations may operate, and test for the pen- study for age at first birth and number of chil- etrance of PGSs (association of genotype and dren ever born in four independent extant data phenotype) at both the individual and ecolog- sources. Despite extensive research on the role ical (that is, state) levels. For most of the phe- of genetics in such outcomes as obesity, sub- notypes they examine, the authors find that the stance use, and education, little attention has ecological correlations are much larger than been directed to fertility behavior, perhaps be- the individual correlations, suggesting the en- cause age at first birth and number of children vironmental context of the state may moderate ever born are complex outcomes related to bi- the genotype-­phenotype associations. In par- ological fecundity, behavioral choice, and ticular, they identify two phenotypes, depres- socio-­environmental factors. Consistent with sion and educational attainment, for which the most of the research on genetic influence using genetic context of a state is especially salient.

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Conclusions Identifies 12 Loci Influencing Human Reproduc- The articles in this issue advance our under- tive Behavior.” Nature Genetics 48(12): 1462–72. standing of the biosocial pathways of well-­ DOI: 10.1038/ng.3698. being across the life course, and their complex Barker, David J. P. 1997. “Maternal Nutrition, Fetal associations with social stratification. They Nutrition, and Disease in Later Life.” Nutrition build on a solid foundation of biosocial re- 13(9): 807–13. DOI: 10.1016/s0899– search in the social sciences, and they show- 9007(97)00193–7. case the value of blurring the boundaries be- ———. 1998. Mothers, Babies and Health in Later Life, tween phenomena outside the body and inside 2nd ed. Edinburgh: Churchill Livingstone. the body. In some cases, they use novel meth- ———. 2006. “Adult Consequences of Fetal Growth ods to cast new light on old questions. In oth- Restriction.” Clinical Obstetrics and Gynecology ers, novel methods reframe the questions and 49(2): 270–83. DOI: 10.1097/00003081– open new lines of inquiry. In all cases, the in- 200606000–00009. tegration of biological information with mea- Bengtsson, Tommy, and Göran Broström. 2009. “Do sures of social environments and behavior Conditions in Early Life Affect Old-­Age Mortality across the life course is generating unique in- Directly and Indirectly? Evidence from 19th-­ sights and unprecedented opportunities for Century Rural Sweden.” Social Science and Med- discovery. In many ways, this issue can be seen icine 68(9): 1583–90. DOI: 10.1016/j.socs- as marking the “coming of age” of a new gen- cimed.2009.02.020. eration of biosocial scholarship, and the future Berenson, Gerald S., Wendy A. Wattigney, Weihang looks bright for those of us who are invested Bao, Sathanur R. Srinivasan, and Bhandaru Rad- in illuminating the complex pathways linking hakrishnamurthy. 1995. Rationale to Study the society, biology, and health across the life Early Natural History of Heart Disease: The Bog- course. alusa Heart Study. The American Journal of the Medical Sciences 310(S1): S22–28. References Bird, Adrian. 2002. “DNA Methylation Patterns and Adam, Emma K., and Meena Kumari. 2009. “Assess- Epigenetic Memory.” Genes and Development ing Salivary Cortisol in Large-­Scale, Epidemio- 16(1): 6–21. DOI: 10.1101/gad.947102. logical Research.” Psychoneuroendocrinology Blackwell, Debra L., Mark D. Hayward, and Eileen M. 34(10): 1423–36. DOI: 10.1016/j. Crimmins. 2001. “Does Childhood Health Affect psyneuen.2009.06.011. Chronic Morbidity in Later Life?” Social Science Adler, Nancy E., Thomas Boyce, Margaret A. and Medicine 52(8): 1269–84. DOI: 10.1016/ Chesney, Sheldon Cohen, Susan Folkman, Robert s0277–9536(00)00230–6. L. Kahn, and S. Leonard Syme. 1994. “Socioeco- Boardman, Jason D., Jonathan Daw, and Jeremy nomic Status and Health: The Challenge of the Freese. 2013. “Defining the Environment in Gradient.” American Psychologist 49(1): 15–24. Gene–Environment Research: Lessons from So- Ahn, Andrew C., Muneesh Tewari, Chi-­Sang Poon, cial Epidemiology.” American Journal of Public and Russell S. Phillips. 2006. “The Limits of Re- Health 103(S1): S64–72. DOI: 10.2105/ ductionism in Medicine: Could Systems Biology ajph.2013.301355. Offer an Alternative?” PLoS Medicine 3(6): e208. Boardman, Jason D., Benjamin W. Domingue, and DOI: 10.1371/journal.pmed.0030208. Jonathan Daw. 2015. “What Can Genes Tell Us Almond, Douglas, and Janet Currie. 2011. “Killing Me About the Relationship Between Education and Softly: The Fetal Origins Hypothesis.” Journal of Health?” Social Science and Medicine 127 (Febru- Economic Perspectives 25(3): 153–72. DOI: ary): 171–80. DOI: 10.1016/j.socs- 10.1257/jep.25.3.153. cimed.2014.08.001. Aneshensel, Carol S. 1992. “Social Stress: Theory Boardman, Jason D., Benjamin W. Domingue, and and Research.” Annual Review of Sociology 18(1): Jason M. Fletcher. 2012. “How Social and Ge- 15–38. DOI: 10.1146/annurev.soc.18.1.15. netic Factors Predict Friendship Networks.” Pro- Barban, Nicola, Rick Jensen, Ronald de Vlaming, Ah- ceedings of the National Academy of Sciences mad Vaez, et al. 2016. “Genome-­Wide Analysis 109(43): 17377–81. DOI: 10.1073/pnas.1208975109.

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