Plemented using the Flash browser plugin. The Bricks items were developed with “Building Bricks”, a web application developed for the purpose, and administered using the “psy.js” JavaScript library; both of these tools are open-source and freely available (see the Supplementary Methods online).MethodsMeasures.Twin data. DZ twins share 50 of their segregating genes on average, while MZ twins share 100 , but environments are shared to approximately the same extent for both MZ and DZ twins. Genetic influence on a trait is therefore indicated by the degree to which the intrapair MZ correlation exceeds the DZ correlation, and cross-twin cross-trait correlations (i.e., the correlation between twin 1 on the first trait and twin 2 on the second) allow the genetic influences common to multiple traits to be estimated. MZs and same-sex DZs are perfectly correlated for sex, and all twins are for age; it is therefore common practice to regress twin data on sex and age, to avoid the artificially inflated estimates of shared environmental influences which would otherwise result31. In addition, for each measure in the present study, outliers beyond 3 SD from the mean were removed, along with any data for those participants suspected to have suffered technical errors or to have responded randomly or PD173074 custom synthesis carelessly (see the Supplementary Methods online). Participants with severe physical or psychological disabilities, or whose mothers had experienced serious perinatal complications, were also excluded from analysis. All variables were standardised, and since the Bricks variables were slightly skewed, a van der Waerden rank transformation32 was performed to ensure that all data were normally distributed, as required for the model-fitting procedures. The study was approved by the appropriate King’s College London ethics committee, and was conducted in accordance with the approved guidelines. Participants provided informed consent. Model-fitting. The data were subjected to full-information maximum-likelihood (FIML) model-fitting procedures, accounting for missing data and combining both same- and opposite-sex DZ twins to maximise power. Univariate ACE models33 were fitted to the data, which use the expected genetic and environmental correlations between the twins (additive genetic influences correlating 1.0 for MZs and 0.5 for DZs; shared environment 1.0 for both; non-shared environment 0 for both) to apportion the variance into components attributable to: i) additive genetic influences (A); ii) shared (or “common”) environmental influences making people raised in the same family more similar to each other (C); and iii) non-shared (unique) environmental influences making them less similar (E, which also includes any measurement error). Individual components may be dropped in nested sub-models, but the full ACE models were used here despite C being non-significant for the Bricks measures, both because this tends to produce the most conservative heritability estimates, and for consistency with the other cognitive measures used (as C is significant for Raven’s Progressive Matrices; see Supplementary Table S15). All model-fitting was conducted using OpenMx34, an R package for structural equations. Multivariate ACE model-fitting uses cross-twin cross-trait correlations22 to estimate the genetic and environmental GW 4064 web sources of covariance, revealing the architecture underpinning two or more traits35. This calculates the genetic correlations (rA) between each pair of var.Plemented using the Flash browser plugin. The Bricks items were developed with “Building Bricks”, a web application developed for the purpose, and administered using the “psy.js” JavaScript library; both of these tools are open-source and freely available (see the Supplementary Methods online).MethodsMeasures.Twin data. DZ twins share 50 of their segregating genes on average, while MZ twins share 100 , but environments are shared to approximately the same extent for both MZ and DZ twins. Genetic influence on a trait is therefore indicated by the degree to which the intrapair MZ correlation exceeds the DZ correlation, and cross-twin cross-trait correlations (i.e., the correlation between twin 1 on the first trait and twin 2 on the second) allow the genetic influences common to multiple traits to be estimated. MZs and same-sex DZs are perfectly correlated for sex, and all twins are for age; it is therefore common practice to regress twin data on sex and age, to avoid the artificially inflated estimates of shared environmental influences which would otherwise result31. In addition, for each measure in the present study, outliers beyond 3 SD from the mean were removed, along with any data for those participants suspected to have suffered technical errors or to have responded randomly or carelessly (see the Supplementary Methods online). Participants with severe physical or psychological disabilities, or whose mothers had experienced serious perinatal complications, were also excluded from analysis. All variables were standardised, and since the Bricks variables were slightly skewed, a van der Waerden rank transformation32 was performed to ensure that all data were normally distributed, as required for the model-fitting procedures. The study was approved by the appropriate King’s College London ethics committee, and was conducted in accordance with the approved guidelines. Participants provided informed consent. Model-fitting. The data were subjected to full-information maximum-likelihood (FIML) model-fitting procedures, accounting for missing data and combining both same- and opposite-sex DZ twins to maximise power. Univariate ACE models33 were fitted to the data, which use the expected genetic and environmental correlations between the twins (additive genetic influences correlating 1.0 for MZs and 0.5 for DZs; shared environment 1.0 for both; non-shared environment 0 for both) to apportion the variance into components attributable to: i) additive genetic influences (A); ii) shared (or “common”) environmental influences making people raised in the same family more similar to each other (C); and iii) non-shared (unique) environmental influences making them less similar (E, which also includes any measurement error). Individual components may be dropped in nested sub-models, but the full ACE models were used here despite C being non-significant for the Bricks measures, both because this tends to produce the most conservative heritability estimates, and for consistency with the other cognitive measures used (as C is significant for Raven’s Progressive Matrices; see Supplementary Table S15). All model-fitting was conducted using OpenMx34, an R package for structural equations. Multivariate ACE model-fitting uses cross-twin cross-trait correlations22 to estimate the genetic and environmental sources of covariance, revealing the architecture underpinning two or more traits35. This calculates the genetic correlations (rA) between each pair of var.
