Joumal of Family Psychology 2014, Vol, 28, No. 3, 357-367

© 2014 American Psychological Association 0893-3200/14/$! 2.00 http://dx.d0i.0rg/J 0.1037/a0036818

For Better and for Worse: Genes and Parenting Interact to Predict Future Behavior in Romantic Relationships April S. Masarik and Rand D. Conger

M. Brent Donnellan

University of California, Davis

Michigan State University

Michael C. Stallings

Monica J. Martin

University of Colorado, Boulder

University of California, Davis

Thomas J. Schofield and Tricia K. Neppl

Laura V. Scaramella

Iowa State University

University of New Orleans

Andrew Smolen

Keith F. Widaman

University of Colorado, Boulder

University of Califomia, Davis

We tested the differential susceptibility hypothesis with respect to connections between interactions in the family of origin and subsequent behaviors with romantic partners. Focal or target participants (G2) in an ongoing longitudinal study (A' = 352) were observed interacting with their parents (Gl) during adolescence and again with their romantic partners in adulthood. Independent observers rated positive engagement and hostility by Gl and G2 during structured interaction tasks. We created an index for hypothesized genetic plasticity by summing G2's allelic variation for polymorphisms in 5 genes (serotonin transporter gene [linked polymorphism], 5-HTT; ankyrin repeat and kinase domain containing 1 gene/dopamine receptor D2 gene, ANKK1/DRD2; dopamine receptor D4 gene, DRD4\ dopamine active transporter gene, DAT; and catechol-0-methyltransferase gene, COMT). Consistent with the differential susceptibility hypothesis, G2s exposed to more hostile and positively engaged parenting behaviors during adolescence were more hostile or positively engaged toward a romantic partner if they had higher scores on the genetic plasticity index. In short, genetic factors moderated the connection between earlier experiences in the family of origin and future romantic relationship behaviors, for better and for worse. Keywords: differential susceptibility, gene by environment interaction (GxE), parenting, romantic relationships, behavior Supplemental materials: http://dx.doi.org/10.1037/a0036818.supp

Earher research suggests that both genetic and environmental factors have a significant influence on the quality of interactions in

romantic relationships (Jerskey et al., 2010; Simons et al., 2013; Spotts et al, 2005, 2004). Because past research indicates that supportive romantic relationships may promote and hostile relationships may impair psychological and physical well-being (e.g., Braithwaite, Delevi, & Eincham, 2010; Reis, ColUns, & Berscheid, 2000), improved understanding of these genetic and environmental influences has both theoretical and practical value. The present study addresses these issues by examining the degree to which certain genetic characteristics interact with parenting history during adolescence to predict the quality of behaviors toward a romantic partner in adulthood. In the present study, we draw upon the differential susceptibility perspective to develop a set of testable hypotheses regarding the interplay between genetic and environmental factors (Belsky, Bakermans-Kranenburg, SL van IJzendoom, 2007; Belsky & Pluess, 2009; Ellis & Boyce, 2011). According to this theoretical framework, certain individuals have a genetic propensity toward susceptibility to environmental influences, whether positive or negative, whereas other individuals are relatively impervious to such influences. We applied this idea to connections between

This article was published Online First May 12, 2014. April S. Masarik and Rand D. Conger, Department of Human Ecology, University of Califomia, Davis; M. Brent Donnellan, Department of Psychology, Michigan State University; Michael C. Stallings, Department of Psychology and Instimte for Behavioral Genetics, University of Colorado, Boulder; Monica J. Martin, Department of Human Ecology, University of Califomia. Davis; Thomas J. Schofield and Trieia K. Neppl, Department of Human Development and Family Studies, Iowa State University; Laura V. Scaramella, Department of Psychology, University of New Orleans; Andrew Smolen, Institute for Behavioral Genetics, University of Colorado, Boulder; Keith F. Widaman, Department of Psychology, University of California, Davis. Support for this work was provided by grants from the Eunice Kennedy Shriver National Institute of Child Health and Human Development (HD047573, HD051746, and HD064687). Correspondence concerning this article should be addressed to April S. Masarik, Department of Human Ecology, University of Califomia, Davis, One Shields Avenue, Davis, CA 95616. E-mail: [email protected] 357

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experiences in the family of origin and subsequent behaviors in romantic relationships in adulthood. Specifically, we tested whether an index hypothesized to represent genetic plasticity interacted with parental positive and negative behaviors during adolescence to predict future positive and negative behaviors with romantic partners in adulthood. We used observational ratings of parent (Gl) and adolescent (G2) behaviors derived from a longterm community study of G2s followed from early adolescence to the young adult years (the Family Transitions Project or FTP; Conger & Conger, 2002).

Parenting and Genetic Influences on Romantic Relationship Development Theory and research suggest that children's experiences with parents are internalized and carried over into their adult relationships (e.g.. Reis et al., 2000; Sroufe, Egeland, Carlson, & Collins, 2005). For example, prospective longitudinal research has documented an association between qualities in the parent-infant relationship (e.g., infant's attachment, parental sensitivity) and social competence in elementary school; attachment to friends in adolescence; and to beliefs, experiences, and behaviors in adult romantic relationships (e.g.. Orina et al., 2011; Roisman, Collins, Sroufe, & Egeland, 2005; Simpson, Collins, Tran, & Haydon, 2007). Although these studies primarily focused on qualities of the early parent-child relationship as predictors of adult romantic relationship outcomes, other work suggests that later parenting also plays an important role in this process. Steinberg (2001) notes that parenting during adolescence involves a unique and influential developmental context because it represents autonomy-related changes and important negotiations in the parent-child relationship. Through experiences with parents, adolescents may learn specific interpersonal skills that promote or hinder future competence in romantic relationships. For example, a study by Whitton et al. (2008) showed that positive engagement and hostility expressed by parents toward their adolescent child during family conflict resolution tasks predicted positive engagement and hostility expressed by offspring toward their spouses in a marital conflict task 17 years later. Similarly, earlier findings from the longitudinal study used for the present analyses showed that nurturant-involved parenting practices during adolescence predicted greater warmth and support and lower hostility toward a romantic partner in adulthood (Conger, Cui, Bryant, & Elder, 2000; Masarik et al, 2013). Moreover, these associations remained signiflcant after controlling for personality traits of the target adolescent (Donnellan, Larsen-Rife, & Conger, 2005). Thus, it appears that both earlier and later parenting may play an important role in romantic relationship development during the adult years. In addition to these parenting influences, findings from twin studies suggest that genetic variability also plays a role in the development and course of romantic relationships. For example, Jerskey et al. (2010) found that genetic effects accounted for 58% of the variability in the likelihood of ever marrying and 32% of the variability in the likelihood of ever divorcing. In a sample of adult twin women, Spotts et al. (2005, 2004) reported that genetic effects accounted for 22% to 36% of the variance in romantic relationship quality. Thus, a complete understanding of the origins of behavioral variation in romantic relationships during the early adult years requires consideration of both genetic and environmental

factors. Indeed, contemporary research and theory about human development has increasingly emphasized the importance of gene by environment interplay, or GxE (e.g., Rutter, Moffitt, & Caspi, 2006).

GxE Research: The Differential Susceptibility Hypothesis Although the impetus for GxE research using candidate polymorphisms came from the diathesis-stress perspective, which suggests that certain genetic characteristics create psychological vulnerability to negative or stressful environments, more recent work has begun to address the differential susceptibility hypothesis. This approach proposes that genetic variation serves to increase responsiveness ("plasticity") to both positive and negative environmental conditions (Belsky et al., 2007; Belsky & Pluess, 2009; Ellis & Boyce, 2011). According to this perspective, individuals labeled genetically "at risk" for poor developmental outcomes in harsh environments may be the same individuals with the best developmental outcomes if exposed to positive environments. A limitation to date for much of the research on differential susceptibility, however, has been a failure to include measures of both positive and negative environments and positive and negative developmental outcomes in any given study (Belsky & Pluess, 2009), an issue we address in the present investigation. In the following review, we first consider the only study of which we are aware that has examined GxE interactions in predicting from parenting during adolescence to behavior in early adult romantic relationships, and then we turn to the empirical and theoretical underpinnings for the genotypes used in the present analyses (serotonin transporter gene (linked polymorphism), 5-HTT; ankyrin repeat and kinase domain containing 1 gene/dopamine receptor D2 gene, ANKK1/DRD2\ dopamine receptor D4 gene, DRD4\ dopamine active transporter gene, DAT; and catechol-O-methyltransferase gene, COMT).

Differential Susceptibility to Parenting in Romantic Relationships To our knowledge, only one published study has reported GxE effects linking qualities of parenting and later qualities in adult romantic relationships (Simons et al., 2013). Consistent with the differential susceptibility hypothesis, Simons and colleagues found that African American children who had relatively common variants on the GABRA2 gene reported more aggressive behavior (e.g., shouting, hitting) toward their romantic partner in their early twenties if they had been subjected to harsh parenting during late childhood and early adolescence (observed during a parent-child interaction task). Yet, they exhibited less aggression toward their partner in adulthood if their parents avoided harsh parenting practices. Important strengths of this study included its prospective, longitudinal research design and the use of observed parenting in the family of origin. In addition, the parenting scale ranged from high support and low hostility to low support and high hostility, thus encompassing both positive and negative family environments in the contextual variable. There were significant limitations, however, in the measure of hostility (i.e., aggression) in young adult romantic relationships. First, this measure did not include a positive dimension; rather, low hostility was interpreted as evidence of positive behaviors in the romantic relationship. As

GENES, PARENTING, AND ROMANTIC RELATIONSHIPS Belsky and Pluess (2009) have noted, this measurement limitation provides only an incomplete evaluation of the differential susceptibility hypothesis. Second, hostility toward a romantic partner was based on self-reports rather than direct observations of couple interactions, a measurement strategy which may introduce unknown biases involving the characteristics of the reporter (Lorenz, Conger, Simons, Whitbeck, & Elder, 1991). Because we did not have a set of polymorphisms on the GABRA2 gene, we were not able to directly replicate this important study. Nonetheless, the fact that these findings indicate that variations in genotypes related to the GABRA2 gene may create greater plasticity to environmental influences suggests that other gene systems may have a similar effect. We do not know from this study, however, whether greater parental supportiveness toward a child or adolescent would interact with genetic plasticity in the GABRA2 gene (or other hypothesized plasticity genes) to influence positive romantic relationship outcomes, such as supportive behaviors toward a romantic partner. We build on the Simons et al. study by investigating a different set of polymorphisms hypothesized to indicate plasticity to both negative and positive dimensions of parenting expected to predict future negative and positive behaviors in adult romantic relationships.

Genotype by Parenting Interactions Predicting to Social and Emotional Development In the present study, we focused on five candidate polymorphisms that have been shown to interact with variations in parenting to predict the social and emotional development of children and adolescents. The current investigation provides an important extension of this earlier research by evaluating the degree to which similar GxE effects can be observed during early adulthood. These polymorphisms were selected both because they are linked to biological processes expected to affect plasticity and also because there is preliminary empirical support for this expectation. Each of the selected polymorphisms has been linked to functioning in either the dopamine system {ANKK1/DRD2, DRD4, DAT, COMT) or the serotonin system {5-HTT). In general, the dopamine system has been associated with reward sensitivity and sensation seeking (e.g., Dreher, Kohn, Kolachana, Weinberger, & Berman, 2009; Stice, Yokum, Burger, Epstein, & Smolen, 2012), and the serotonin system has been associated with experiences of threat or displeasure (e.g., Caspi, Hariri, Holmes, Uher, & Moffitt, 2010). Thus, rewarding or threatening environmental experiences (e.g., positive or negative parenting histories) may exert influence most powerfully for individuals who have lower genetic thresholds for pleasure and displeasure, respectively (Belsky & Pluess, 2009). Furthermore, the low-activity or minor alíeles that are hypothesized to act as plasticity alíeles for these polymorphisms are linked to increased activity in the brain's limbic system (e.g., the amygdala), thereby increasing emotional responsiveness to environmental stimuli. Biologically, then, individuals with more (compared with those with less) genetic plasticity should have more sensitive central nervous systems that are more responsive to environmental influences. Eariier research on parenting behaviors provides preliminary support for this hypothesis for the five polymorphisms used in the present investigation. First, in three independent studies, Hankin and colleagues (2011) found that children aged 9-15 years old carrying the short

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(s) alíele of the 5-HTTLPR (serotonin transporter linked polymorphism) had the highest and lowest levels of positive affectivity under conditions of high and low supportive parenting, respectively—consistent with differential susceptibility. However, this study did not examine a negative pole forthe developmental outcome (e.g., negative affectivity). Second, Mills-Koonce and colleagues (2007) found that 3-year-old children carrying at least one Al alíele for the Taql A polymorphism of the ANKKl (ankyrin repeat and kinase domain containing 1) gene were less likely to have affective problems if matemal sensitivity was high, whereas no effect of matemal sensitivity was documented for children not carrying the Al alíele, afindingalso consistent with differential susceptibihty. In this case, however, the study did not include a measure reflecting psychological health (e.g., positive añectivity). It is also important to note that the Taq 1A polymorphism has previously been studied in association with DRD2; however, it has been suggested thai ANKKl, which exists downstream fkim DRD2, may be responsible for at least some of the effects earher attributed to DRD2 (Neville, Johtistone, & Walton, 2004). To account for this observation, we refer to this genotype as ANKK1/DRD2. Third, preschool children carrying one 7R alíele for the exon-3 Variable Number Tandem Repeat (VNTR) of the DRD4 gene exhibited a greater number of externalizing problems when exposed to less maternal sensitivity at 10 months of age, but fewer externalizing problems when exposed to greater matemal sensitivity (Bakermans-Kranenburg & van IJzendoom, 2006). Also consistent with differential susceptibility, children (ages 1-3) carrying the 7R alíele demonstrated the greatest decrease in externalizing behaviors after an intervention in which parents were trained to be more sensitive and use more positive disciplining strategies (Bakermans-Kranenburg, van IJzendoorn, Pijlman, Mesman, & Juffer, 2008). The major limitation in these studies was the absence of a positive developmental outcome (e.g., prosocial behavior). Fourth, Laucht and colleagues (2007) found that adolescents homozygous for the lOR alíele of the DAT (dopamine active transporter) 5' VNTR polymorphism whose parents reported greater exposure to family adversity displayed more inattention and hyperactivity-impulsiveness than others. Although this study did not explicitly test the differential susceptibility hypothesis, their results are consistent with differential susceptibility because adolescents homozygous for the lOR alíele had lower levels of inattention and hyperactivity-impulsiveness under conditions of low family adversity. Finally, Laucht and colleagues (2012) found support for differential susceptibility because adolescents homozygous for the Met alíele of the Vall58Met polymorphism in the COMT gene exhibited greater drinking activity when their parents were less involved and provided less supervision; however, drinking activity decreased with more parental involvement and supervision. Yet, no explicitly positive dimension of development was included in the analyses. Collectively, these findings suggest that genetic plasticity for each one of the five genotypes examined in the present study increased children's sensitivity to the quality of parenting behavior. As noted, however, none of these studies simultaneously considered both positive and negative dimensions of parenting and also positive and negative children's social development. For that reason, none of these investigations provides a complete test of the differential susceptibility hypothesis. Moreover, none of these studies examined whether the tested parenting by genotype interacfions predicted beyond childhood or adolescence and into the adult years.

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360 The Present Study

In the present study, we hypothesized that our measure of genetic plasticity would interact with both negative and positive parenting behaviors experienced in adolescence to predict future negative and positive behaviors toward a romantic partner in adulthood. This strategy directly tests the central tenets of differential susceptibility; that G2 individuals with greater genetic plasticity will fare worse in their adult romantic relationships if exposed to Gl negative parenting environments (hostility) during adolescence, but will fare better if exposed to positive parenting environments (positive engagement). Because the positive and negative behaviors of parents and adolescents tend to be significantly correlated (Conger & Ge, 1999), one problem in making inferences about the influence of earlier parenting on G2 behaviors in adult relationships is that this association may simply reflect continuity in G2 behavior over time. That is, Gl parenting may have no influence on later G2 behavior once earlier G2 behavior to parents is taken into account. For that reason, we controlled for adolescents' behavior toward their parents in all of our statistical models. That is, we hypothesized that the interaction between genetic plasticity and parenting would predict future romantic relationship behavior over and above the continuity in G2 behavior over time. An important feature of the present study is that we used trained observer ratings to assess Gl parents' behavior toward their G2 adolescent, the target adolescent's behavior toward their parents, and the target's behavior toward a romantic partner in adulthood. Indeed, many of the reviewed GxE studies relied on self-reports of a particular behavior (e.g., parent report of their parenting, youth report of affective problems); therefore, we hoped to complement and expand this line of work by using measures of observable behavior. Moreover, the earlier studies using the same genotypes employed in the present investigation limited their investigation of GxE effects to childhood or adolescence. The present analyses extend that earlier work by examining the interaction between Gl parenting and G2 genetic variability as a predictor of G2 behavior toward a romantic partner in adulthood, 16 years later. In addition, although most GxE work has examined one genetic polymorphism at a time, recent studies of differential susceptibility have used composite measures of hypothesized genetic plasticity by summing allelic variation across a set of polymorphisms (see Beaver, Sak, Vaske, & Nilsson, 2010; Belsky & Beaver, 2011). This approach operates in much the same way as indices of contextual risk in which multiple environmental risk factors are combined to create a single score of overall risk (e.g., Evans, Li, & Sepanski Whipple, 2013). That is, individuals with the greatest number of plasticity alíeles are hypothesized to be the most susceptible to environmental influences. We adhere to this strategy given evidence that multiple genes, particularly those just reviewed (i.e., 5-HTF, ANKK1/DRD2, DRD4, DAT, COMT), may operate in a cumulative fashion as plasticity genes (Belsky & Pluess, 2009). Moreover, this approach is consistent with the notion that each individual candidate gene will likely have only a small effect on behavior and that larger effects are more likely to be observed when a number of these small effects are combined in a cumulative fashion. To summarize, we proposed that parenting experienced in adolescence would interact with an adolescent's cumulative genetic

plasticity to predict his or her future behaviors toward a romantic partner in adulthood. We tested the "strong differential susceptibility" hypothesis following analytic procedures recently proposed by Belsky, Pluess, and Widaman (2013). This hypothesis proposes that environmental factors such as the quality of parenting will have no statistical effects on individual behavior in the absence of genetic plasticity. Support for differential susceptibility will be observed if both (a) the genetic index amplifies the association between parental hostility and later hostility by the adult child toward a romantic partner and (b) the genetic index amplifies the association between parental positive engagement and later positive engagement with a romantic partner.

Method Participants Data come from a prospective, longitudinal study of over 500 families living in the Midwest and initiated in 1989 (FTP; Conger & Conger, 2002). Focal or "target" adolescents (G2), a close-aged sibling, and their biological parents (Gl) were visited in their homes by trained interviewers. Starting in 1991, a supplementary sample of 108 single-parent families (mother-headed) was added to the study. At this time, 82% of the adolescents were from two-parent families and 18% were from single mother families. Because the present study focused on associations between parenting experienced during adolescence and romantic relationship functioning in adulthood, for these analyses we selected targets (N = 352; 195 females) who participated with a romantic partner in 2007 (M age = 31.64, SD = .46). The 2007 assessment was the most recent wave of available observational data involving romantic partners. We used parenting data from the 1991 assessment, when targets were on average 15.09 years old {SD = .41). The 1991 wave of data collection represents the earliest assessment involving both two-parent and single-parent families; thus, we chose this time period for parent-adolescent behaviors so that we could maximize the available sample size. These selected participants came from both two-parent families {n = 294) and singlemother families {n = 58). Of the 352 targets who participated with a romantic partner, 285 were married, 54 were cohabiting fulltime, and 13 were in steady dating relationships (including parttime cohabiting). Two same-sex cohabiting couples were included in the study. The ethnic/racial background is predominately European American reflecting the demographics of the region at study initiation.

Procedure During home visits, families were videotaped during structured interaction tasks designed to elicit information about social skills and emotional responses. During adolescence (1991), trained interviewers began videotaped sessions by asking each family member to independently complete a short questionnaire to identify issues of concern that led to disagreements within the family (e.g., responsibility for chores, use of money, etc.). Family members then gathered around a table and were given a set of cards and instructed to read questions aloud and discuss their answers to the questions (Task 1). These cards contained general questions about family life such as important family events, approaches to parent-

GENES, PARENTING, AND ROMANTIC RELATIONSHIPS ing, and household chores. This first task lasted about 30 min and was designed to provide family members with the opportunity to express a range of emotions including positive sentiment. Task 2 (15 min) also involved all family members. For this task, interviewers selected three issues based on the quesdonnaires completed at the beginning of the assessment and family members were asked to discuss and resolve these issues that they had identified as leading to confiict in their family. Task 2 also was designed to elicit a range of emotion, especially negative affect, and is comparable to confiict tasks typically used in studies of family interaction (see Melby & Conger, 2001 for more details). Starting in 1995 (1 year post high school), the romantic partner of each target participated if one was available and the study's focus shifted from the family of origin to the target's adult relationships. Targets and their romantic partners were videotaped biennially thereafter using similar procedures as the family of origin assessments; however, only one romande relationship task was completed during the target's adult years. In brief, targets and their romantic partner were asked to discuss the history and current status of their relationship, enjoyable events and activides they had engaged in during the past year, areas of agreement and disagreement, and plans for the future. This task was designed to elicit both positive and negative affect and was an attempt to generate the same range of emotions elicited by Tasks 1 and 2 during adolescence. Approximately 20% of all videotaped interaction tasks (for both the family of origin assessments and adult romantic reladonship assessments) were randomly assigned for rating by a second, independent observer. The primary and secondary radngs were then used to generate estimates of interobserver reliability.

Measures Parenting behaviors toward target adolescent. In 1991, parenting was assessed via observer ratings of two videotaped interaction tasks (Tasks 1 and 2), Observers rated verbal and nonverbal behavior by the mother and father (when available) to the target adolescent using the Iowa Family Interaction Rating Scales (IFIRS; Melby & Conger, 2001), The IFIRS has been udlized in a variety of cross-sectional and longitudinal studies examining diverse topics such as economic stress, parendng, adolescent development, and romantic relationships, and has acceptable reliability and validity (Melby & Conger, 2001), The parental hostility construct consisted of three ratings (Hostility, Angry Coercion, and Antisocial Behavior) of parent behavior toward the target adolescent in Tasks 1 and 2, More information about the parental hosdlity scales, as well as the other scales discussed below, can be found in Melby and Conger (2001), Scores for the three scales were averaged separately for mothers and fathers in each task. Mother and father behaviors were significantly correlated in Task 1 (r = ,44, p < ,001) and Task 2 (r = ,37, p < ,001) and were averaged together to refiect parental hosdlity in each task. Parental hosdlity in Task 1 was significandy correlated with parental hosdlity in Task 2{r= ,63, p < ,001) and radngs from each task were averaged together to refiect overall parental hostility. Across tasks, the average interobserver reliabilities and internal consistencies (separated by a slash) for parental hostility were ,93/,87 (tnother and father) and ,90/,88 (single mothers). The averaged scores ranged from 1,17-9,00 (M = 3,63, SD = 1,48),

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The parental positive engagement construct consisted of four ratings (Listener Responsiveness, Positive Assertiveness, Positive Communication, and Prosocial Behavior) of parent behavior toward the target adolescent in Tasks 1 and 2, Scores for the four scales were averaged separately for mothers and fathers in each task. Mother and father behaviors were significandy correlated in Task I (/• = .46, p< ,001) and Task 2 (r = ,43, p < ,001) and were averaged together to reflect parental positive engagement. Parental posidve engagement in Task 1 was significantly correlated with parental posidve engagement in Task 2 (r = ,41, p < ,001) and ratings from each task were averaged together to refiect overall parental positive engagement. Across tasks, the average interobserver reliabilities and internal consistencies (separated by a slash) for parental positive engagement were ,87/,84 (mother and father) and .92/,88 (single mothers). The averaged scores ranged from 1,50-8,50 (M = 4,49, SD = 1,02). Target adolescent behaviors toward parents. In 1991, adolescent behaviors toward parents were assessed in Tasks 1 and 2, To be consistent, we used the same observadonal ratings for hostility and positive engagement from Tasks 1 and 2 as used for the parenting constructs. Target adolescent hostility toward their mother in Task 1 was significantly correlated with target's hostility toward their father in Task 1 (r = ,80, p < ,001) and likewise in Task 2 (r = ,77, p < ,001), Hosdlity enacted in each task was then averaged together to refiect target hostility to parents. Target hostility to parents in Task 1 was significantly correlated with target hostility to parents in Task 2 (r = ,66, p < ,001) and radngs from each task were then averaged together to reflect overall target hostility to parents. Across tasks, the average interobserver reliabilities and internal consistencies (separated by a slash) for target hostility were ,94/,89 (mother and father) and ,92/,90 (single mothers). The averaged scores ranged from 1-9 (M = 3,99, SD = 1,90), Target adolescent positive engagement toward their mother in Task 1 was significantly correlated with positive engagement toward their father in Task 1 (r = ,90, p < ,001) and likewise in Task 2 (r = ,88, p < ,001), Positive engagement enacted in each task was then averaged together. Target posidve engagement to parents in Task 1 was significantly correlated with target positive engagement to parents in Task 2 (r = ,40, p < ,001) and ratings from each task were then averaged together to reflect overall target positive engagement to parents. Across tasks, the average interobserver reliabilides and internal consistencies (separated by a slash) for target posidve engagement were ,84/,79 (mother and father) and ,88/,84 (single mothers). The averaged scores ranged from 1-7,25 (M = 3,23, SD = ,92), Target behaviors toward romantic partner in adultbood. In 2007, couples participated in one discussion task. Again, we used the same ratings of target's behaviors toward their romande partner as we used for the parent-to-adolescent and adolescent-toparent measures. Scores ranged from 1 to 9 (M = 3,40, SD = 1,61) for romantic relationship hostility (reliabilities = .92/, 87) and from 1 to 9 (M = 5,79, SD = 1,56) for romantic relationship positive engagement (reliabilities = ,95/,92), Genotyping. Saliva samples were obtained from target participants with Oragene (DNA Genotek, Ontario, Canada) collection kits. Genotyping was conducted at the University of Colorado's Institute for Behavioral Genetics. Genomic DNA was isolated with Agencourt DNAdvance DNA Isoladon Kits (Beckman

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Coulter, Brea, CA) using a Beckman-Coulter Biomek EX workstation according to company protocols. Methods for genotyping the DAT and DRD4 VNTRs are detailed in Anchordoquy, McGeary, Liu, Krauter, and Smolen (2003), and the method for 5-HTTLPR is in Whisman, Richardson, and Smolen (2011). Genotyping of the Taql A polymorphism and Vall58Met polymorphism in COMT are outlined in Haberstick and Smolen (2004). Based on past research (e.g., Belsky & Pluess, 2009), we consider the following alíeles to confer plasticity: short (s) alíele of 5-HTTLPR in 5-HJT (accounting for SNP rs25531); Al alíele of the Taql A polymorphism in ANKK1/DRD2\ 7R alíele of exon-3 VNTR in DRD4\ lOR alíele of the 5' VNTR in DAT, and the Met alíele of the Vall58Met polymorphism in COMT. Polymorphisms received a score of "0" if none of these alíeles were observed, a score of " 1 " if one of these alíeles was observed, and "2" if two of these alíeles were observed. Einally, these scores were summed to create a cumulative genetic index of hypothesized plasticity. Although index scores could theoretically range from 0-10, the observed range was 1-8 indicating that all participants had at least one of the hypothesized plasticity alíeles (M = 4.40, SD = 1.38). Based on examination of duplicate controls and Mendelian inconsistencies among family members, genotype error rates were less than 1% for all five polymorphisms; and alíele and expected genotype distributions were in Hardy-Weinberg equilibrium. Moreover, the alíele frequencies were in line with other Caucasian populations (see http://alfred.med.yale.edu; Rajeevan, Soundararajan, Kidd, Pakstis, & Kidd, 2012). Table 1 provides the intercorrelations and descriptive statistics for all genotypes used in the present study. These correlations ranged from —.14 to -I-.IO, indicating that scores on these polymorphisms were primarily unrelated to one another.

Results We initially inspected gene-environment correlations (KJE) between targets' genetic index and parenting (see Table 2 for correlations among all study constructs). The possibility of an evocative genetic effect on parenting was mied out because the index was not significantly related to either parental hostility ir = -.02, p = .11) or positive engagement (r = .04, p = .52). In other words, there was no evidence that adolescents received particular parenting behaviors that were influenced by the adolescents' genotype (see Rutter et al, 2006). In fact, the genetic index was not significantly correlated with any of the behavioral measures (Table 2). All of the behavioral measures were significantly interrelated in the expected directions (e.g., r = .25, p < .001, between parent hostility and target hostility toward a romantic partner). Next, we conducted regression analyses (Table 3). Model 1 is labeled the "hostility model" (parental hostility -^ romantic relationship hostility) and Model 2 is the "positive engagement model" (parental positive engagement -^ romantic relationship positive engagement). To ensure that the hostility and positive engagement measures should be examined as separate constructs, confirmatory factor analyses (CEAs) were conducted with parenting and romantic relationship behaviors. Eirst, all individual rating scales comprising hostility and positive engagement behaviors were assigned to a single constmct. Model fit was poor (Comparative Eit Index [CEI] = .622/.765; Tucker-Lewis Index [TLI] = .433/.647; Root Mean Square Error of Approximation [RMSEA] = .347/.308 for

parenting/romantic relationship behaviors, respectively) and many of the factor loadings were poor (e.g., .26/.47 for parental/romantic relationship rating of Angry Coercion). Model fit was better when indicators of hostility and positive engagement were separated to load on their respective constructs (CEI = .996/.987; TLI = .990/.976; RMSEA = .046/.080 for parenting/romantic relationship behaviors, respectively). Furthermore, all factor loadings in these models were of acceptable magnitude (loadings ranged from .64-.98 across parenting and romantic relationship models). These analyses suggested that it was appropriate to separate hostility and positive engagement constructs in the following regression models.' Out of 352 targets, 303 (86%) had complete genetic data. Regression analyses were conducted in Mplus Version 6 (Muthén & Muthén, 1998-2011) using full information maximum likelihood estimation (EIML). Analyses also were conducted using list-wise deletion. Results did not differ in statistical significance or magnitude compared with findings using EIML. Target gender was added as a control variable in the analyses. Gender did not significantly interact with any of the predictors (e.g., genetic index, parenting, genetic index X parenting). As noted, we controlled for target behavior toward parents during adolescence in all of our models to assure that parenting effects were not spurious and a result of continuity in target behavior. To test for strong differential susceptibility as proposed by Belsky et al. (2013), we set the parenting effect to zero in both the hostility (Model 1) and positive engagement models (Model 2). Constraining this parameter to zero implies no environmental (i.e., parenting) effect for individuals with predicted scores of zero for the genetic index. The standardized coefficients in Table 3 indicate that the GxE interaction effects were statistically significant for both the hostility (ß = .184, p < .05) and positive engagement models (ß = .201, p < .05). These coefficients suggest that, even after controlling for target adolescent's behavior toward parents, (a) higher scores on the genetic index magnified associations between parental hostility and hostility toward a romantic partner, and (b) higher scores on the genetic index magnified associations between parental positive engagement and positive engagement toward a romantic partner. To further evaluate the fit of the strong differential susceptibility model (Belsky et al, 2013), we allowed the main effects of parenting to be freely estimated in follow-up analyses. That is, we tested whether the weak differential susceptibility hypothesis— which posits that individuals with no genetic plasticity would be ' Although results from CFAs suggested it was more appropriate to separate measures of hostility from positive engagement rather than combine them into a single construct, the zero-order association between hostility and positive engagement is quite strong for both parents' behavior toward target [r = -.52, p < .001) and target's behavior toward parents (r = -.62,p < .001; Table 2). Although the correlation is not perfect (i.e., 1), it may suggest that hostility and positive engagement reflect opposite dimensions of the same continuum, in exploratory analyses (not shown), we averaged ratings of hostility and positive engagement to create a single continuum of behavior ranging from high positive engagement/low hostility to low positive engagement/high hostility. None of the major findings (i.e., GxE effects) reported in the following sections differed when we used this approach. However, we chose to separately assess the negative and positive dimensions of behavior in the present report consistent with the results of the CFA and to provide a more detailed analysis of the differential susceptibility hypothesis.

GENES, PARENTING, AND ROMANTIC RELATIONSHIPS

363

Table 1

Intercorrelations and Descriptives for Genotypes Percentage Genotype

1

2

1. 5-HTT 2. ANKK1/DRD2 3. DRD4 4. DAT 5. COMT

.04 -.06 .10+ -.06

— -.14* .03 .03

3

— -.01 .00

4

M

SD

0

1

2

Total N

— .00

0.99 0.39 0.36 1.50 1.15

.69 .57 .52 .63 .71

24% 65% 66% 7% 19%

54% 31% 32% 36% 48%

23% 4% 2% 57% 34%

313 317 311 314 314

Note. 5-HTT = serotonin transporter gene (linked polymorphism); ANKK1/DRD2 = ankyrin repeat and kinase domain containing 1 gene/dopamine receptor D2 gene (TaqlA polymorphism); DRD4 = dopamine receptor D4 gene (exon-3 VNTR); DAT = dopamine active transporter gene (5' VNTR polymorphism); COMT = catechol-O-methyltransferase gene (Vall58Met polymorphism). Scores of 0, 1,2 represent 0, 1, and 2 plasticity alíeles for each genotype, respectively. Percentages may not equal 100 percent because of rounding.

V < .05.

affected by parenting, but to a lesser degree than those carrying more genetic plasticity—would fit the data better (see Belsky et al., 2013). In these models (not shown), parenting main effects failed to reach statistical significance for both Model 1 and Model 2. Because these models were less parsimonious and did not lead to significant improvements in model fit over the strong version of differential susceptibility (Model 1: Ax^ = 2.188, Mf = 1, not statistically significant [ns\; Model 2: Ax^ = .937, Ad/ = 1, ns), we concluded that the strong differential susceptibility models best fit the data. To examine the GxE interactions in more detail, we plotted separate lines for predicted scores of behaviors toward a romantic partner as a function of earlier parenting behaviors across the range of predicted values (0-8) for the cumulative genetic index (Figure 1A and B). We used an online tool for estimating the magnitude of the simple slopes between parenting behaviors and adult children's behavior toward their romantic partner for each predicted score on the cumulative genetic index (Preacher, Curran, & Bauer, 2006). These estimates, which are reported in Figure 1, demonstrate that as the score on the genetic index increases, the magnitude of the simple slope increases linearly. Although we did not observe a genetic index score of zero (i.e., all participants had at least one plasticity alíele), the estimate of predicted values included the zero point. This procedure allowed us to directly test the strong differential susceptibility hypothesis that individuals with a predicted

value of zero for the genetic index should be unaffected by parenting. As shown in Figure lA, parental hostility in adolescence predicted romantic relationship hostility in adulthood except for targets with a predicted score of "0" on the genetic index. As the number of plasficity alíeles increased, the magnitude of the slope increased. Figure IB demonstrates the same findings for parental positive engagement experienced in adolescence predicting romantic relationship positive engagement in adulthood (except for targets with a predicted score of "0" on the genetic index). To increase understanding of these genetic effects, we conducted additional exploratory analyses. First, to assure that no single genotype was accounting for the observed effects of the genetic index, we reran the analyses for Models 1 and 2 five different times using just four polymorphisms in each analysis. For example, one analysis was run with the DAT genotype omitted, then with COMT omitted, and so forth. Each of these regressions produced essentially the same results as reported in Table 3 indicating that none of the individual genotypes fully accounts for the effects of the genetic index. Second, we evaluated each individual genotype to determine if it interacted with parenting during adolescence to predict romantic relationship behaviors in the same way that the cumulative genetic index did. In these analyses, we used the same procedures as used for the cumulative genetic index models. Results are presented in supplemental material available online. In brief, the interaction

Table 2 Intercorrelations Among Study Constructs (N = 352; Full Information Maximum Likelihood) Hostility

Positive engagement

Construct Hostility 1. Parent —> target 2. Target —» parent 3. Target -^ romantic partner Positive engagement 4. Parent -^ target 5. Target —> parent 6. Target -^ romantic partner 7. Target genetic index

5 — .67 .25 -.52 -.40 -.21 -.02

.32*

36" 52" 22" 04

-.23" -.23" -.62" -.03

— .51" .21" .04

.26" .01

-.04

Note. Parent -* target = parent behavior toward target (1991); Target -^ parent = target behavior toward parent (1991); Target -^ romantic partner = target behavior toward romantic partner (2007). 'p < .01. " p < .001.

MASARIK ET AL.

364 Table 3

Regression Coefficients for Predicting Hostility (Model 1) and Positive Engagement (Model 2) Toward a Romantic Partner as a Function of Gender, Parent-Target Adolescent Behavior, Target Genetic Index, and the Interaction Between Parent Behavior and Target Genetic Index Model 1; Hostility (Target --* romantic partner) Predictors Intercept

Female Target —» parent Parent —> target Target genetic index Parent —> target X target genetic index R^ {SE)

B 2.481 .729"' .163" .000 -.156+ .035*

SE B

ß

.354 .163 .227 .057 .194 .000 .000 .083 -.136 .184 .016 .148 (.034)*"

Model 2:: Positive engagement (Target —> romanticpartner) ; B

SE B

.444 4.983 -.171 .162 .350** .102 .000 .000 -.231* .109 .040* .020 .086 (.030)**

ß -.054 .207 .000 -.205 .201

Note. Female = 1 (0 = male). Target —> parent = target adolescent behavior toward parent; Parent —> target = parent behavior toward target adolescent; B = unstandardized coefficient; SE = standard error; ß = standardized coefficient. Models were estimated using full information maximum likelihood (FIML; total N = 352). Parental hostility and positive engagement was constrained to "0" to model strong differential susceptibility. V
hypothesized to be involved in plasticity. However, we advance this earlier research by considering negative and positive parenting environments and negative and positive romantic relationship outcomes in separate models. In doing so, we add substantial new evidence in support of the differenrial suscep-

tibility hypothesis (Belsky et al., 2007; Belsky & Pluess, 2009; Ellis & Boyce, 2011). Also important, the Simons et al. study did not control for continuity in G2 behavior from adolescence to adulthood. That added control in the present study increases confidence in the findings.

366

MASARIK ET AL,

We also acknowledge limitations in the current study. First, participants were ethnically and geographically homogeneous, Sitnilar GxE research is needed with other ethnic/racial groups to increase confidence in generalizability of findings. The Simons et al, study of African American youth helps to meet that need. Second, romantic relationship functioning is undoubtedly affected by other environmental infiuences (e,g,, interparental relationships), individual differences (e,g,, personality), and other biological factors (e,g,, physiological reacdvity) not invesdgated in this report. Future research should build upon our findings to provide a more comprehensive and interdisciplinary understanding of romande reladonship development. Finally, because we did not have prospecdve information about parenting received as a child or adolescent for the romande partners in the study, we were unable to consider these data in ottr analyses. Future research with similar background data for both partners would further etmch this type of invesdgadon. These limitadons are balanced by a consideration of the strengths of the present report. First, trained observers assessed the parenting and romantic reladonship constructs during an interacdon task. This approach is an important complement to self-report measures because it offers a relatively objective perspective on how families and couples actually behave. Second, data come from a prospective, longitudinal study that has followed the same cohort of participants and their families from adolescence to adulthood, whereas cross-sectional or short-term longitudinal research designs may not fully capture developmental-contextual processes. Third, we considered both negative and positive environmental predictors and outcomes thereby providing comprehensive empirical support for the basic tenets of the differential susceptibility perspective (Belsky & Pluess, 2009), Perhaps most important, these findings contribute to an improved understanding of the factors that influence behaviors in romantic relationships inasmuch as romantic relationships are an integral component of psychological and physical well-being (e,g,, Braithwaite et al,, 2010; Reis et al,, 2000), The fact that the most genetically sensitive individuals benefitted more and suffered more in their romande relationships as a function of earlier exposure to positive and negative parenting behaviors, respectively, also has possible clinical implicadons. Specifically, these findings and other recent studies (e,g,, Simons et al, 2013) suggest that genetic variations should make some individuals more and others less responsive to therapeutic efforts to improve the quality of romantic and parent-child relationships. Indeed, recent empirical evidence from differential suscepdbility experiments demonstrated that the seemingly "most vulnerable" children (indicated by variation in temperament, physiological reacdvity, or candidate genes) profited the most from family based intervendon programs (see van IJzendoom & Bakermans-Kranenburg, 2012), These observations suggest a more encouraging view of the potential for behavioral change through environmental enrichment (e,g,, prevendon and/or intervention). Especially important, genetically sensitive individuals may be the ones who reap the most benefits from couple-based or parent-child education or treatment programs and genetic informadon may become an important element in tailoring specific interventions to specific individuals. In conclusion, the current article provides novel evidence to suggest that certain genetic factors confer increased suscepdbility to earlier family environments and subsequent behaviors in adult romantic relationships—for better and for worse. This work points

to important roles for both nature and nurture, and we hope that future studies continue to unravel the complex ways that these two forces interact to shape human development and improve the quality of romantic relationships.

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Received September 17, 2013 Revision received March 14, 2014 Accepted March 25, 2014 •

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For better and for worse: genes and parenting interact to predict future behavior in romantic relationships.

We tested the differential susceptibility hypothesis with respect to connections between interactions in the family of origin and subsequent behaviors...
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