to the DZ correlation of .60. Adoption data, including adopted-apart parents and offspring and adopted-apart siblings as well asMZ twins adopted apart, also point to substantial genetic influence.Model-fitting analyses that simultaneously analyze all of the fam-ily, adoption, and twin data summarized in Figure 1 yield herita-bility estimates of about 50% (Chipuer, Rovine, & Plomin, 1990;Loehlin, 1989). In other words, about half of the total variance(which includes error of measurement) can be attributed to DNAdifferences between individuals. Heritability is of course higher if corrections are made for error of measurement as in analyses of latent variables free of measurement error. Even an attempt toexplain as much of the variance of
as possible in terms of prenatal effects nonetheless yielded a heritability estimate of 48%(Devlin, Daniels, & Roeder, 1997; McGue, 1997). Although mostof this research was conducted in the United States and WesternEuropean countries, similar estimates of heritability have beenfound in countries such as Moscow, the former East Germany,Japan, and rural and urban India (Plomin, DeFries, et al., 2001).The convergence of evidence on the conclusion that individualdifferences in intelligence are substantially heritable led to a de-cline in the 1990s of genetic research on intelligence that merelyaimed to investigate the heritability of intelligence. Instead, ge-netic designs were used to go beyond estimating heritability inorder to ask questions about environmental influences, develop-mental change and continuity, and multivariate issues. Beforediscussing these three topics, it should be noted that assortativemating for intelligence is substantial. Correlations betweenspouses are only about .10 for other personality traits and about .20for height and weight, but assortative mating for intelligence isabout .40 (Plomin, DeFries, et al., 2001). The importance of assortative mating is that it increases genetic variance generationafter generation and may thus contribute to the high heritability of intelligence. Twin studies that do not take assortative mating intoaccount underestimate heritability because the genetic effects of assortative mating inflate the DZ correlation but not the MZcorrelation (Plomin, DeFries, et al., 2001).
Concerning the environment, genetic research provides the bestavailable evidence for the importance of environmental influenceson intelligence: If heritability is 50%, that means that environmen-tal factors account for the rest of the reliable variance. Two of themost important findings from genetic research are about nurturerather than nature. First, nearly all personality traits show that,contrary to theories of socialization from Freud onwards, environ-mental influences operate to make siblings growing up in the samefamily as different from one another as children growing up indifferent families (Harris, 1998; Plomin & Daniels, 1987). How-ever, intelligence is the exception to this rule of nonshared envi-ronmental influence (Plomin, 1988). Direct estimates of the im-portance of shared environmental influence come fromcorrelations of .19 for adoptive parents and their adopted childrenand .32 for adoptive siblings (see Figure 1). Because adoptivesiblings are unrelated genetically, what makes them similar isshared rearing, suggesting that about a third of the total variancecan be explained by shared environmental influences. The factorsresponsible for this shared environmental influence have not beenpinned down, although general family background variables suchas socioeconomic status are likely to contribute. There has beenone largely unsuccessful attempt to identify specific aspects of thehome environment that are responsible for the shared environmen-tal influence on intelligence in childhood (Chipuer & Plomin,1992). However, as explained in the following section on devel-opment, although shared environment is important for intelligencein childhood, its importance declines to negligible levels afteradolescence. In other words, shared environmental factors relevantto intelligence would be expected to show associations in child-hood but not later in development. Moreover, this finding suggeststhat, even for intelligence, the salient environmental factors arenonshared after childhood.The second finding has been called the
nature of nurture
(Plo-min & Bergeman, 1991). When used as outcome measures ingenetic research, environmental measures consistently point tosome genetic influence, suggesting that genetic factors influencethe way we react and interact with the environment, a type of genotype-environment correlation (Kendler & Eaves, 1986). Forexample, a widely used measure of the home environment inresearch on cognitive development is the Home Observation forMeasurement of the Environment (HOME; Bradley, Convyn,Burchinal, McAdoo, & Coll, 2001). In an adoption study compar-ing nonadoptive and adoptive siblings, genetic influences wereestimated to account for about 40% of the variance of HOMEscores (Braungart, Fulker, & Plomin, 1992). Moreover, multivar-iate genetic analysis (described below) indicated that about half of the phenotypic correlation between the HOME and children
sintelligence is mediated genetically. This research suggests that wecreate our experiences in part for genetic reasons and supports acurrent shift from thinking about passive models of how theenvironment affects individuals toward models that recognize theactive role we play in selecting, modifying, and creating our ownenvironments (Plomin, 1994). In quantitative genetics, this topic isreferred to as active genotype-environment correlation (Plomin,DeFries, et al., 2001).
Two types of developmental questions have been addressed ingenetic research. The first question is: Does heritability changeduring development? Because it is so reasonable to assume thatgenetic differences become less important as experiences accumu-late during the course of life, one of the most interesting findingsabout intelligence is that the opposite is closer to the truth. Re-search during the past decade has shown that the heritability of
increases during development. Figure 2 summarizes twin resultsby age (McGue, Bouchard, Iacono, & Lykken, 1993), showing thatthe difference between MZ and DZ twin correlations increasesslightly from early to middle childhood and then increases dra-matically in adulthood. Because relatively few twin studies of intelligence have included adults, summaries of intelligence data(see Figure 1) showing heritability estimates of about 50% restprimarily on data from childhood. Heritability in adulthood ishigher, perhaps as high as 80%, although there is some evidencethat heritability late in life might be lower (Finkel, Pedersen,McGue, & McClearn, 1995). A finer grained analysis of twinresults indicates that heritability is lower in infancy (about 20%)than in middle childhood (about 40%; Plomin, 1986). The modestheritability of intelligence in early childhood was confirmed in a
PLOMIN AND SPINATH