We identified several key regulatory transcription factors in each cell type using the expression and accessibility of all target genes within the trans-regulatory network constructed by LINGER40 (Supplementary Fig. 7d, e). For example, in D2 MSNs, the transcription factor predicted to have the most significant difference between those with and without AUD, based on the chromatin openness of its target genes, was FOXO1 (Forkhead transcriptional factor O1, Supplementary Fig. 7d), which has been previously shown to regulate energy homeostasis in neurons41 and was linked to depression in recent studies42,43.