Prediction of motif activities for each cell was performed using the RunChromVAR command in Signac. Briefly, chromVAR identifies motifs associated with variability in chromatin accessibility between cells107). Differential testing on the chromVAR z-score was performed using the FindMarkers function, setting mean.fxn=rowMeans and fc.name="avg_diff", so that the fold-change represents the average difference in z-score between the groups.