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              <jats:title>METHODS<\/jats:title>\n                    <jats:p>We leveraged functional genomics data from Religious Orders Study or the Rush Memory and Aging Project (ROS\/MAP) and Mount Sinai Brain Bank (MSBB) brain biobanks and single\u2010cell RNA\u2010sequencing data from HHV\u2010infected forebrain organoids to investigate HHV\u2010infection\u2010associated transposable element (TE) dysregulation underlying AD etiologies.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>RESULTS<\/jats:title>\n                    <jats:p>\n                      We identified widespread TE dysregulation in HHV\u2010positive human AD brains, including an astrocyte\u2010specific upregulation of LINE1 subfamily TEs in HHV\u2010positive human AD brains. We further pinpointed astrocyte\u2010specific LINE1 upregulation that could potentially regulate target gene\n                      <jats:italic>NEAT1<\/jats:italic>\n                      expression via long\u2010range enhancer\u2010promoter chromatin interactions. This LINE1 dysregulation can be partially reversed by the usage of anti\u2010HHV drugs (valacyclovir and acyclovir) in a virus\u2010infected human brain organoid model. Finally, we demonstrated that valacyclovir rescued tau\u2010associated neuropathology and alleviated LINE1 activation in an experimental tau aggregation model.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>DISCUSSION<\/jats:title>\n                    <jats:p>Our analysis provides associations linking molecular, clinical, and neuropathological AD features with HHV infection, which warrants future clinical validation.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Highlights<\/jats:title>\n                    <jats:p>\n                      <jats:list list-type=\"bullet\">\n                        <jats:list-item>\n                          <jats:p>\n                            Via analysis of bulk RNA\u2010seq data in two large\u2010scale human brain biobanks, ROS\/MAP (\n                            <jats:italic>n<\/jats:italic>\n                            \u00a0=\u00a0109 pathologically confirmed AD and\n                            <jats:italic>n<\/jats:italic>\n                            \u00a0=\u00a044 cognitively healthy controls) and MSBB (\n                            <jats:italic>n<\/jats:italic>\n                            \u00a0=\u00a0284 AD and\n                            <jats:italic>n<\/jats:italic>\n                            \u00a0=\u00a0150 cognitively healthy controls), we identified widespread TE activation in HHV\u2010positive human AD brains and significantly positive associations of HHV RNA abundance with\n                            <jats:italic>APOE4<\/jats:italic>\n                            genotype, Braak staging score, and CERAD score.\n                          <\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>\n                            We identified cell type\u2010specific LINE1 upregulation in both microglia and astrocytes of human AD brains via long\u2010range enhancer\u2013promoter chromatin interactions on lncRNA nuclear enriched abundant transcript 1 (\n                            <jats:italic>NEAT1<\/jats:italic>\n                            ).\n                          <\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>We determined that usage of valacyclovir and acyclovir was significantly associated with reduced incidence of AD in a large real\u2010world patient database.<\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>Using the HEK293 tau P301S model and U2OS mt\u2010Keima cell model, we determined that valacyclovir treatment rescued tau\u2010associated neuropathology and alleviated activation of LINE1 with increased cellular autophagy\u2010level mechanistically supported clinical benefits of valacyclovir in real\u2010world patient data.<\/jats:p>\n                        <\/jats:list-item>\n                      <\/jats:list>\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1002\/alz.14595","type":"journal-article","created":{"date-parts":[[2025,2,22]],"date-time":"2025-02-22T09:54:41Z","timestamp":1740218081000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":16,"title":["Human herpesvirus\u2010associated transposable element activation in human aging brains with Alzheimer's disease"],"prefix":"10.1002","volume":"21","author":[{"given":"Yayan","family":"Feng","sequence":"first","affiliation":[{"name":"Cleveland Clinic Genome Center, Lerner Research Institute Cleveland Clinic  Cleveland Ohio USA"},{"name":"Genomic Medicine Institute, Lerner Research Institute Cleveland Clinic  Cleveland Ohio 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