Nucleus preparation from hippocampus tissue was based on previously described methods (PMID 38092918), the final nuclei is in the 1× nucleus buffer (nucleus buffer (10x Genomics). Nuclei (1uL) was stained with Trypan Blue (Invitrogen, T10282) and  manually counted. In total, 16,000–20,000 nuclei were used for the tagmentation reaction and 10x Chromium controller loading. The DNA and RNA libraries were generated according to the manufacturer’s recommended protocol (https://www.10xgenomics. com/support/single-cell-multiome-atac-plus-gene-expression). 10x multiome ATAC–seq and RNA-sequencing (RNA-seq) libraries were paired-end sequenced on the Illumina NovaSeq X systems to a depth of around 50,000 reads per cell for each modality.