For each miRNA with conserved predicted mRNA targets (defined by TargetScan [37]), a one-sided GSE test was carried out to test if the miRNA had a significant anti-correlated association with its set of predicted mRNA targets. The GSE test was based on statistics from association tests between a particular miRNA and all of the mRNAs, one at a time, using a linear fixed-effects model adjusted for relevant covariates, with the miRNA as the predictor and one mRNA as the response (see Materials and Methods). In the GSE analysis, we test whether there was an enrichment of highly ranked test statistics within the set of predicted mRNA targets of the miRNA being tested, with the null hypothesis that the predicted target set was randomly chosen (see Materials and Methods). The analysis included 248 profiled miRNAs with conserved predicted mRNA targets, since these are thought to be more likely to have a gene-regulatory function [38]. Thus, we performed 248 GSE analyses. In abdominal adipose tissue 55 out of 248 (22%) miRNAs were found to have a significant (FDR adjusted p-value<0.05) association with
