International Journal of Child Care and Education Policy Copyright 2013 by Korea Institute of Child Care and Education

2013, Vol. 7, No. 2, 15-31

Differential Susceptibility to Environmental Influences

Jay Belsky
University of California, Davis & King Abdulaziz University
USA

Evidence that adverse rearing environments exert negative effects particularly on children and adults
presumed ―vulnerable‖ for temperamental or genetic reasons may actually reflect something else:
heightened susceptibility to the negative effects of risky environments and to the beneficial effects of
supportive environments. Building on Belsky’s (1997, 2005; Belsky & Pluess, 2009) evolutionary-inspired
differential susceptibility hypothesis stipulating that some individuals, including children, are more
affected—both for better and for worse—by their environmental exposures and developmental
experiences, recent research consistent with this claim is reviewed. It reveals that in many cases, including
both observational field studies and experimental intervention ones, putatively vulnerable children and
adults are especially susceptible to both positive and negative environmental effects. In addition to
reviewing relevant evidence, unknowns in the differential-susceptibility equation are highlighted.

Key words: early childhood education and care, preschool, policy, ethnic minority children, special needs
children

Introduction most work still focuses on effects of

environmental exposures and developmental
Most students of child development experiences that apply equally to all
probably do not presume that all children—so-called main effects of
children are equally susceptible to parenti7ng or poverty or being reared
rearing (or other environmental) effects; by a depressed mother—thus failing to
a long history of research on consider interaction effects, which
interactions between parenting and reflect the fact that whether, how, and
temperament, or parenting-by-temperament how much these contextual conditions
interactions, clearly suggests otherwise. influence the child may depend on the
Nevertheless, it remains the case that child’s temperament or some other
characteristic of individuality.
Correspondence regarding this article should Research on parenting-by-temperament
be addressed to Jay Belsky, professor, University
interactions is based on the premise
of California, Davis, USA and King Abdulaziz
University, Saudi Arabia. Electronic mail may be that what proves effective for some
sent to jbelsky@ucdavis.edu.

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Jay Belsky

individuals in fostering the development of Diathesis-Stress, Dual-Risk and

some valued outcome—or preventing Vulnerability
some problematic one—may simply not
do so for others. Commonly tested are The view that infants and toddlers
diathesis-stress hypotheses derived manifesting high levels of negative
from multiple-risk/transactional frameworks emotion are at special risk of
in which individual characteristics that problematic development when they
make children ―vulnerable‖ to adverse experience poor-quality rearing is
experiences—placing them ―at risk‖ of widespread. Evidence consistent with
developing poorly—are mainly influential this view can be found in the work of
when there is at the same time some Morrell and Murray (2003), who
contributing risk from the environmental showed that it was only highly
context (Zuckerman, 1999). Diathesis distressed and irritable 4-month-old
refers to the latent weakness or boys who experienced coercive and
vulnerability that that a child or adult rejecting mothering at this age who
may carry (e.g., difficult temperament, continued to show evidence, 5 months
particular gene), but which does not later, of emotional and behavioural
manifest itself, thereby undermining dysregulation. Relatedly, Belsky, Hsieh,
well-being, unless the individual is and Crnic (1998) observed that infants
exposed to conditions of risk or stress. who scored high in negative
After highlighting some research emotionality at 12 months of age and
consistent with a diathesis-stress or who experienced the least supportive
dual-risk perspective, I raise questions mothering and fathering across their
—on the basis of other findings—about second and third years of life scored
how the first set of data has been highest on externalizing problems at 36
interpreted, advancing the evolutionary- months of age. And Deater-Deckard
inspired proposition that some children, and Dodge (1997) reported that
for temperamental or genetic reasons, children rated highest on externalizing-
are actually more susceptible to both (a) behavior problems by teachers across
the adverse effects of unsupportive the primary-school years were those
parenting and (b) the beneficial effects who experienced the most harsh
of supportive rearing (Belsky, 1997, discipline prior to kindergarten entry
2005; Belsky & Pluess, 2009, 2013a). and who were characterized by
Finally, I draw conclusions and mothers at age 5 as being negatively
highlight some ―unknowns in the reactive infants.
differential-susceptibility equation.‖ The adverse consequences of the co-
occurrence of a child risk factor (i.e., a
diathesis; e.g., negative emotionality)

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 Differential Susceptibility to Environmental Influences

and problematic parenting also is reproductive fitness, that is, the

evident in Caspi and Moffitt’s (2006) dispersion of genes in future
ground-breaking research on gene-by- generations, the ultimate goal of
environment (GXE) interaction. Young Darwinian evolution. To protect against
men followed from early childhood all children being steered, inadvertently,
were most likely to manifest high levels in a parental direction that proved
of antisocial behavior when they had disastrous at some later point in time,
both (a) a history of child maltreatment developmental processes were selected
and (b) a particular variant of the MAO- to vary children’s susceptibility to
A gene, a gene previously linked to rearing (and other environmental
aggressive behaviour. Such results led influences). In what follows, I review
Rutter (2006), like others, to speak of evidence consistent with this claim
―vulnerable individuals,‖ a concept that which highlights early negative
also applies to children putatively at emotionality and particular candidate
risk for compromised development due genes as ―plasticity factors‖ making
to their behavioral attributes. But is individuals more susceptible to both
―vulnerability‖ the best way to supportive and unsupportive
conceptualize the kind of person-X- environments, that is, ―for better and for
environment interactions under worse‖ (Belsky, Bakerman-Kranenburg, &
consideration? van IJzendoorn, 2007).

Negative Emotionality as Plasticity

Beyond Diathesis-Stress, Dual- Factor
Risk and Vulnerability The first evidence which Belsky (1997,
2005; Belsky & Pluess, 2009) could point
Working from an evolutionary to consistent with his differential-
perspective, Belsky (1997, 2005; Belsky susceptibility hypothesis concerned
& Pluess, 2009) theorized that children, early negative emotionality. Children
especially within a family, should vary scoring high on this supposed ―risk
in their susceptibility to both adverse factor‖, particularly in the early years,
and beneficial effects of rearing appeared to benefit disproportionately
influence (see also Ellis, Boyce, Belsky, from supportive rearing environments
Bakermans- Kranenburg & van (Boyce & Ellis, 2005; Ellis, Boyce, Belsky,
IJzendoorn, 2011). Because the future is Bakermans-Kranenburg & van IJzendoorn,
uncertain, in ancestral times, just like 2011). Feldman, Greenbaum, and
today, parents could not know for Yirmiya (1999) found, for example, that
certain (consciously or unconsciously) 9-month-olds scoring high on negativity
what rearing strategies would maximise who experienced low levels of

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Jay Belsky

synchrony in mother–infant interaction Experimental studies designed to test

manifested more noncompliance during Belsky’s (1997, 2005) differential-
clean-up at age two than other children susceptibility hypothesis are even more
did. When such infants experienced suggestive than the longitudinal-
mutually synchronous mother–infant correlational evidence just cited. Blair
interaction, however, they displayed (2002) discovered that it was highly-
greater self-control than did children negative infants who benefited most—
manifesting much less negativity as in terms of both reduced levels of
infants. Subsequently, Kochanska, Aksan, externalizing behavior problems and
and Joy (2007) observed that highly enhanced cognitive functioning—from
fearful 15-month-olds experiencing a multi-faceted infant-toddler intervention
high levels of power-assertive paternal program whose data he reanalyzed.
discipline were most likely to cheat in a Thereafter, Klein Velderman, Bakermans-
game at 38 months, yet when cared for Kranenburg, Juffer, and van IJzendoorn
in a supportive manner such negatively (2006) found that experimentally
emotional, fearful toddlers manifested induced changes in maternal sensitivity
the most rule-compatible conduct. exerted greater impact on the
In the time since Belsky and Pluess attachment security of highly negatively
(2009) reviewed evidence like that just reactive infants than it did on other
cited, highlighting the role of negative infants. In both experiments, environmental
emotionality as a ―plasticity factor‖, influences on ―vulnerable‖ children
even more evidence to this effect has were for better instead of for worse.
emerged in the case of children. As it turns out, there is ever-growing
Consider in this regard work linking (1) experimental evidence that early
maternal empathy (Pitzer, Jennen- negative emotionality is a plasticity
Steinmetz, Esser, Schmidt, & Laucht, factor. Consider findings showing that
2011) and anger (Poehlmann et al., 2012) it is infants who score relatively low on
with externalizing problems; (2) mutual irritability as newborns who fail to
responsiveness observed in the mother- benefit from an otherwise security-
child dyad with effortful control (Kim promoting intervention (Cassidy,
& Kochanska, 2012); (3) intrusive Woodhouse, Sherman, Stupica, &
maternal behavior (Conway & Stifter, Lejuez, 2011) and infants who show few,
2012) and poverty (Raver, Blair, & if any, mild perinatal adversities—
Willoughby, 2013) with executive known to be related to limited negative
functioning; and (4) sensitive parenting emotionality—who fail to benefit from
with social, emotional and cognitive- computer-based instruction otherwise
academic development (Roisman et al., found to promote preschoolers’ phonemic
2012). awareness and early literacy (Van der

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 Differential Susceptibility to Environmental Influences

Kooy-Hofland, van der Kooy, Bus, van and reward mechanisms and one
Ijzendoorn, & Bonsel, 2012). In other variant of this polymorphism, the 7-
words, only the putatively ―vulnerable‖ repeat allele, has been linked to lower
—those manifesting or likely to dopamine reception efficiency, van
manifest high levels of negativity— IJzendoorn and Bakermans-Kranenburg
experienced developmental enhancement (2006) predicted this allele would
as a function of the interventions cited. moderate the association between
Similar results emerge among older maternal unresolved loss or trauma and
children, as Scott and O’Connor’s (2012) infant attachment disorganization.
parenting intervention resulted in the Having the 7-repeat DRD4 allele
most positive change in conduct among substantially increased risk for
emotionally dysregulated children (i.e., disorganization in children exposed to
loses temper, angry, touchy). maternal unresolved loss/trauma, as
expected, consistent with the diathesis-
Genes as Plasticity Factors stress framework; yet when children
Perhaps nowhere has the diathesis- with this supposed ―vulnerability gene‖
stress framework informed person-X- were raised by mothers who had no
environment interaction research more unresolved loss, they displayed significantly
than in the study of GXE interaction. less disorganization than agemates
Recent studies involving measured without the allele, regardless of
genes and measured environments also mothers’ unresolved-loss status
document both for better and for worse (Bakermans-Kranenburg & van IJzendoorn,
environmental effects--in the case of 2011).
susceptible individuals—as it turns out. Similar results emerged when the
Here I consider evidence pertaining to interplay between DRD4 and observed
two specific candidate genes before parental insensitivity in predicting
turning attention to research examining externalizing problems was studied in a
multiple genes at the same time. group of 47 twins (Bakermans-
Kranenburg & van IJzendoorn, 2007).
DRD4 Children carrying the 7-repeat DRD4
One of the most widely studied allele raised by insensitive mothers
genetic polymorphisms in research displayed more externalizing behaviors
involving measured genes and than children without the DRD4 7-
measured environments pertains to a repeat (irrespective of maternal
particular allele (or variant) of the sensitivity), whereas children with the
dopamine receptor gene, DRD4. 7-repeat allele raised by sensitive
Because the dopaminergic system is mothers showed the lowest levels of
engaged in attentional, motivational, externalizing problem behavior

19
Jay Belsky

(Bakermans-Kranenburg & van IJzendoorn, (Fortuna et al., 2011). Especially

2007). Such results suggest that noteworthy, perhaps are the results of a
conceptualizing the 7-repeat DRD4 meta-analysis of GXE research involving
allele exclusively in risk-factor terms is dopamine-related genes showing that
misguided, as this variant of the gene children eight and younger respond to
seems to heighten susceptibility to a positive and negative developmental
wide variety of environments, with experiences and environmental exposures
supportive and risky contexts promoting, in a manner consistent with differential
respectively, positive and negative susceptibility (Bakermans-Kranenburg
functioning. & van IJzendoorn, 2011).
In the time since I last reviewed such As in the case of negative emotionality,
differential-susceptibility-related evidence intervention research also underscores
(Belsky & Pluess, 2009), ever more GXE the susceptibility to 7-repeat carriers of
findings pertaining to DRD4 (and other the DRD4 gene to benefit disproportionately
polymorphisms, see below) have from supportive environments. Kegel,
appeared consistent with the notion Bus and van IJzendoorn (2011) tested
that there are individual differences in and found support for the hypothesis
developmental plasticity. Consider in that it would be DRD4 7R carriers who
this regard recent differential- would benefit from specially designed
susceptibility-related evidence showing computer games promoting phonemic
heightened—or exclusive—susceptibility awareness and, thereby, early literacy
of individuals carrying the 7-repeat in their randomized control trial (RCT).
allele when the environmental Other such RCT results point in the
predictor and developmental outcome same direction with regard to DRD4-7R,
were, respectively, (a) maternal including research on African-
positivity and pro-social behavior American teenagers in which substance
(Knafo, Israel, & Ebstein, 2011); (b) use was the outcome examined (Beach,
early nonfamilial childcare and social Brody, Lei, & Philibert, 2010; Brody,
competence (Belsky & Pluess, 2013b); (c) Chen, & Beach, 2013; Brody, Chen,
contextual stress and support and Beach, et al., 2013).
adolescent negative arousal (Beach et
al., 2012); (d) childhood adversity and 5-HTTLPR
young-adult persistent alcohol dependence Perhaps the most studied polymorphism
(Park, Sher, Todorov, & Heath, 2011); in research on GXE interactions is the
and (e) newborn risk status (i.e., serotonin-transporter gene, 5-HTTLPR.
gestational age, birth weight for Most research distinguishes those who
gestational age, length of stay in NICU) carry one or two short alleles (s/s, s/l)
and observed maternal sensitivity and those homozygous for the long

20
 Differential Susceptibility to Environmental Influences

allele (l/l). The short allele has but best—showing least problems—
generally been associated with reduced when encountering few or none (e.g.,
expression of the serotonin transporter Wilhelm et al., 2006). Calling explicit
molecule––which is involved in the attention to such a pattern of results,
reuptake of serotonin from the synaptic Taylor and associates (2006) reported
cleft—and thus considered to be related that young adults homozygous for
to depression, either directly or in the short alleles (s/s) manifested greater
face of adversity. Indeed, the short depressive symptomatology than
allele has often been conceptualized as individuals with other allelic variants
a ―depression gene‖. when exposed to early adversity (i.e.,
Caspi and associates (2003) were the problematic childrearing history), as
first to show that the 5-HTTLPR well as many recent negative life events,
moderates effects of stressful life events yet the fewest symptoms when they
during early adulthood on depressive experienced a supportive early
symptoms, as well as on probability of environment or recent positive
suicide ideation/attempts and of major experiences. The same for-better-and-
depression episode at age 26 years. for-worse pattern of results concerning
Individuals with two s alleles proved depression are evident in Eley et al.’s
most adversely affected whereas effects (2004) research on adolescent girls who
on l/l genotypes were weaker or were and were not exposed to risky
entirely absent. Of special significance, family environments.
however, is that carriers of the s/s allele The effect of 5-HTTLPR in moderating
scored best on the outcomes just environmental influences in a manner
mentioned when stressful life events consistent with differential susceptibility is
were absent, though not by very much. not restricted to depression and its
Multiple research groups have symptoms. It also emerges in studies of
attempted to replicate Caspi et al.’s anxiety (Stein, Schork & Gelernter, 2008)
(2003) findings of increased vulnerability and ADHD, particularly ADHD which
to depression in response to stressful persists into adulthood (Retz et al.,
life events for individuals with one or 2008). In all these cases, emotional
more copies of the s allele, with many abuse in childhood (Stein et al., 2008) or
succeeding (see below), but certainly a generally adverse childrearing
not all (e.g., Surtees et al., 2006; Risch et environment (Retz et al., 2008), it
al., 2009). The data presented in quite a proved to be those individuals carrying
number of studies indicates, however, short alleles who responded to
that individuals carrying short alleles developmental or concurrent experiences
(s/s, s/l) did not just function most in a for-better-and-for-worse manner,
poorly when exposed to many stressors, depending on the nature of the

21
Jay Belsky

experience in question. age showing that short-allele carriers

Since last reviewing such 5-HTTLPR- are more susceptible to the effects of
related GXE research consistent with both positive and negative developmental
differential susceptibility (Belsky & experiences and environmental exposures,
Pluess, 2009), ever more evidence in at least in the case of Caucasians (Van
line with the just-cited work has IJzendoorn, Belsky & Bakermans-
emerged. Consider in this regard Kranenburg, 2013).
evidence showing for-better-and-for- As was the case with DRD4, there is
worse results in the case of those also evidence from intervention studies
carrying one or more short alleles of 5- documenting differential susceptibility.
HTTLPR when the rearing predictor Consider in this regard Drury and
and child outcome were, respectively, associates (2012) data showing that it
(a) maternal responsiveness and child was only children growing up in
moral internalization (Kochanska et al., Romanian orphanages who carried 5-
2011), (b) child maltreatment and HTTLPR short alleles who benefited
children’s antisocial behavior (Cicchetti, from being randomly assigned to high-
Rogosch, & Thibodeau, 2012), and (c) quality foster care—in terms of
supportive parenting and children’s reductions in the display of indiscriminant
positive affect (Hankin et al., 2011). friendliness. Eley and associates (2012)
Differential-susceptibility-related findings also documented intervention benefits
also emerged (among male African- restricted to short allele carriers in their
American adolescents) when (d) study of cognitive behavior therapy for
perceived racial discrimination was children suffering from severe anxiety,
used to predict conduct problems but their design included only treated
(Brody et al., 2011); (e) when life events children (i.e., did not involve a
were used to predict neuroticism randomly assigned control group).
(Pluess, Belsky, Way, & Taylor, 2010)
and (f) life satisfaction of young adults Polygenetic Plasticity
(Kuepper et al., 2012); and (g) when Most GXE research, like that just
retrospectively-reported childhood considered, has focused on one or
adversity was used to explain aspects another polymorphism—like DRD4 or
of impulsivity among college students 5-HTTLPR. In recent years, however,
(e.g., pervasive influence of feelings, work has emerged focusing on multiple
feelings trigger action) (Carver, Johnson, polymorphisms and thus reflecting the
Joormann, Kim, & Nam, 2011). operation of epistatic (i.e., GXG)
Especially noteworthy are the results of interactions (e.g., Beaver, Sak, Vaske, &
a recent meta-analysis of GXE findings Nilsson, 2010; Conner, Hellemann,
pertaining to children under 18 years of Ritchie, & Noble, 2010), as well as

22
 Differential Susceptibility to Environmental Influences

GXGXE ones. One can distinguish anxiety and somatic symptoms

polygenetic GXE research in terms of (Cicchetti, Rogosch, & Sturge-Apple,
the basis used for creating multi-gene 2007), (b) between perceived racial
composites. One strategy involves discrimination and risk-related cognitions
identifying genes which show main reflecting a fast vs. slow life-history
effects and then compositing only these strategy (Gibbons et al., 2012), (c)
to then test an interaction with some between contextual stress/support and
environmental parameter (e.g., Docherty, aggression in young adulthood (Simons
Kovas, & Plomin, 2011). Another et al., 2011) and (d) between social class
approach is to composite genes for a and post-partum depression (Mitchell
secondary, follow-up analysis that have et al., 2011). Of note, too is evidence
been found in a first round of inquiry to that a 3-gene composite moderates the
generate significant GXE interactions relation between a hostile-demoralizing
(e.g., Brody, Chen, & Beach, 2013; community and family environment
Sonuga-Barke et al., 2009). When and aggression in early adulthood
Cicchetti and Rogosch (2012) applied (Simons et al., 2011) and that a 5-gene
this approach using four different composite moderates the relation
polymorphisms, they found that as the between parenting and adolescent self-
number of sensitivity-to-the- control (Belsky & Beaver, 2011).
environment alleles increased (i.e., S/S Given research already reviewed, it is
allele of 5HTTLPR, zero copies of CRH1 probably not surprising that there is
TAT haplotype, the TT genotype of also work examining genetically
DRD4-521C/T, and A carrier of OXTR), moderated intervention effects focusing
so did the degree to which maltreated on multi-gene composites rather than
and non-maltreated low-income singular candidate genes. Consider in
children differed on a composite this regard the Drury et al.’s (2012)
measure of resilient functioning—in a findings showing that even though the
for-better-and-for-worse manner. genetic polymorphism brain derived
A third approach which has now neurotrophic factor, BDNF, did not—all
been used successfully a number of by itself—operate as a plasticity factor
times to chronicle differential susceptibility when it came to distinguishing those
involves compositing a set of genes who did and did not benefit from the
selected on an a-priori basis before aforementioned foster-care intervention
evaluating GXE (e.g., Brody, Chen, & implemented with institutionalized
Beach, 2013). Consider in this regard children in Romania, the already-noted
evidence indicating that 2-gene moderating effect of 5-HTTLPR was
composites moderate links (a) between amplified if a child carried Met rather
sexual abuse and adolescent depression/ than Val alleles of BDNF along with

23
Jay Belsky

short 5-HTTLPR alleles. In other words, Although research summarized here

the more plasticity alleles children suggests that the concept has utility,
carried, the more their indiscriminate there are many ―unknowns,‖ several of
friendliness declined over time when which are highlighted in this
assigned to foster care and the more it concluding section.
increased if they remained institutionalized.
Consider next Brody, Chen and Beach’s Domain General or Domain Specific?
(2013) confirmed prediction that the Is it the case that some children,
more GABAergic and Dopaminergic perhaps those who begin life as highly
genes African-American teens carried, negatively emotional, are more
the more protected they were from susceptible both to a wide variety of
increasing their alcohol use over time rearing influences and with respect to a
when enrolled in a whole-family wide variety of developmental
prevention program. Such results once outcomes—as is presumed in the use of
again call attention to the benefits of concepts like ―fixed‖ and ―plastic‖
moving beyond single polymorphisms strategists (Belsky, 2005), with the latter
when it comes to operationalizing the being highly malleable and the former
plasticity phenotype. They also indicate hardly at all? Boyce and Ellis (2005)
that even if a single gene may not by contend that a general psychobiological
itself moderate an intervention (or reactivity makes some children
other environmental) effect, it could especially vulnerable to stress and thus
still play a role in determining the to general health problems. Or is it the
degree to which an individual benefits. case, as Belsky (2005) wonders and
These are insights future investigators— Kochanska, Aksan, and Joy argue
and interventionists—should keep in (2007), that different children are
mind when seeking to illuminate ―what susceptible to different environmental
works for whom?‖ influences (e.g., nurturance, hostility)
and with respect to different outcomes?
Pertinent to this idea are findings of
Unknowns in the Differential- Caspi and Moffitt (2006) indicating that
Susceptibility Equation different genes differentially moderated
the effect of child maltreatment on
The notion of differential susceptibility, antisocial behavior (MAO-A) and on
derived as it is from evolutionary depression (5HTT).
theorizing, has gained great attention in
recent years, including a special section Continuous Versus Discrete Plasticity?
in the journal Development and The central argument that children
Psychopathology (see Ellis et al., 2011). vary in their susceptibility to rearing

24
 Differential Susceptibility to Environmental Influences

influences raises the question of how to IJzendoorn’s (2007) aforementioned

conceptualize differential susceptibility: findings pertaining to DRD4, given the
categorically (some children highly link between the dopamine system and
plastic and others not so at all) or attention. Kochanska et al., (2007)
continuously (some children simply contend that the ease with which
more malleable than others)? It may anxiety is induced in fearful children
even be that plasticity is discrete for makes them highly responsive to
some environment–outcome relations, parental demands. And Belsky (2005)
with some individuals affected and speculates that negativity actually
others not at all (e.g., gender-specific reflects a highly sensitive nervous
effects), but that plasticity is more system on which experience registers
continuous for other susceptibility powerfully—negatively when not
factors (e.g., in the case of the regulated by the caregiver but
increasing vulnerability to stress of positively when coregulation occurs—a
parents with decreasing dopaminergic point of view somewhat related to
efficiency; van IJzendoorn, Bakermans- Boyce and Ellis’ (2005) proposal that
Kranenburg, & Mesman, 2008. Certainly susceptibility may reflect prenatally
the work which composites multiple programmed hyper-reactivity to stress.
genotypes implies that there is a
―plasticity gradient‖, with some
children higher and some lower in References
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environment and developmental
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outcome, but they do not elucidate the
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