PHACTR1 and migraine disorder: By integrating sc-ST with multi-omics profiling (TWAS, SMR, PoPS), we uncovered microenvironment-specific co-localization of cross-subtype core genes (e.g., PHACTR1-STAT6 clusters in meningeal vascular niches) and developmental origins of subtype divergence, such as prenatal enrichment of migraine-associated loci in neural crest-derived tissues (e.g., jaw primordium).