Association tests were performed in R (R Development Core Team 2011) with the lmekin function of the kinship package (Atkinson and Therneau 2008). This function provides a linear mixed-effects model whereby the genetic relatedness among individuals (based on the kinship coefficient) is incorporated into the covariance structure of the random effects. This adjusts the model fit and compensates for the fact that the siblings are related and, therefore, so are their genotypes and phenotypes, which would otherwise violate the assumption of independent observations in a linear regression model.