We downloaded curated sets of known TF-targets and tissue-expressed genes from the Enrichr64,65 website (http://amp.pharm.mssm.edu/Enrichr/). TF-target gene sets were assayed by ChIP-X experiments from the ChEA66 and ENCODE67,68 projects. Tissue-expressed genes were based on the ARCHS4 database69. Gene sets were processed, mapped to entrez IDs with R package ClusterProfiler v3.10.170 (http://bioconductor.org/packages/release/bioc/html/clusterProfiler.html) and tested for over-representation by one-sided Fisher’s exact test as implemented in the R package GeneOverlap v1.18.0 (https://www.bioconductor.org/packages/release/bioc/html/GeneOverlap.html), by using 19,942 genes tested in trans-eQTL analysis as background. Multiple testing correction was conducted using Benjamini-Hochberg method17.