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Genet."},{"key":"10.1016\/j.cell.2026.04.014_bib71","doi-asserted-by":"crossref","first-page":"535","DOI":"10.1016\/j.cell.2018.12.015","article-title":"Predicting Splicing from Primary Sequence with Deep Learning","volume":"176","author":"Jaganathan","year":"2019","journal-title":"Cell"},{"key":"10.1016\/j.cell.2026.04.014_bib72","doi-asserted-by":"crossref","first-page":"2456","DOI":"10.1016\/j.cell.2023.03.035","article-title":"Massively parallel base editing to map variant effects in human hematopoiesis","volume":"186","author":"Martin-Rufino","year":"2023","journal-title":"Cell"},{"key":"10.1016\/j.cell.2026.04.014_bib73","doi-asserted-by":"crossref","first-page":"52","DOI":"10.1126\/science.ads7951","article-title":"Transcription factor networks disproportionately enrich for heritability of blood cell phenotypes","volume":"388","author":"Martin-Rufino","year":"2025","journal-title":"Science"},{"key":"10.1016\/j.cell.2026.04.014_bib74","doi-asserted-by":"crossref","first-page":"463","DOI":"10.1016\/j.cell.2020.05.037","article-title":"Determinants of Base Editing Outcomes from Target Library Analysis and Machine Learning","volume":"182","author":"Arbab","year":"2020","journal-title":"Cell"},{"key":"10.1016\/j.cell.2026.04.014_bib75","doi-asserted-by":"crossref","first-page":"1632","DOI":"10.1002\/dvdy.24183","article-title":"Poly peak parser: Method and software for identification of unknown indels using sanger sequencing of polymerase chain reaction products","volume":"243","author":"Hill","year":"2014","journal-title":"Dev. Dyn."},{"key":"10.1016\/j.cell.2026.04.014_bib76","doi-asserted-by":"crossref","first-page":"e168","DOI":"10.1093\/nar\/gku936","article-title":"Easy quantitative assessment of genome editing by sequence trace decomposition","volume":"42","author":"Brinkman","year":"2014","journal-title":"Nucleic Acids Res."},{"key":"10.1016\/j.cell.2026.04.014_bib77","doi-asserted-by":"crossref","first-page":"1694","DOI":"10.1038\/emboj.2008.94","article-title":"Alu element-mediated gene silencing","volume":"27","author":"Chen","year":"2008","journal-title":"EMBO J."},{"key":"10.1016\/j.cell.2026.04.014_bib78","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1007\/978-1-4939-1369-5_11","article-title":"RNA fluorescence in situ hybridization in cultured mammalian cells","volume":"1206","author":"Tripathi","year":"2015","journal-title":"Methods Mol. Biol."},{"key":"10.1016\/j.cell.2026.04.014_bib79","doi-asserted-by":"crossref","first-page":"621","DOI":"10.1016\/j.cell.2020.03.006","article-title":"Distinct Processing of lncRNAs Contributes to Non-conserved Functions in Stem Cells","volume":"181","author":"Guo","year":"2020","journal-title":"Cell"},{"key":"10.1016\/j.cell.2026.04.014_bib80","doi-asserted-by":"crossref","DOI":"10.1126\/science.aaa6090","article-title":"RNA imaging. Spatially resolved, highly multiplexed RNA profiling in single cells","volume":"348","author":"Chen","year":"2015","journal-title":"Science"},{"key":"10.1016\/j.cell.2026.04.014_bib81","doi-asserted-by":"crossref","DOI":"10.1126\/science.adx3800","article-title":"High-resolution spatial mapping of cell state and lineage dynamics in vivo with PEtracer","volume":"390","author":"Koblan","year":"2025","journal-title":"Science"},{"key":"10.1016\/j.cell.2026.04.014_bib82","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1007\/978-1-61779-424-7_25","article-title":"Silencing of gene expression by gymnotic delivery of antisense oligonucleotides","volume":"815","author":"Soifer","year":"2012","journal-title":"Methods Mol. Biol."},{"key":"10.1016\/j.cell.2026.04.014_bib83","doi-asserted-by":"crossref","first-page":"e3","DOI":"10.1093\/nar\/gkp841","article-title":"Efficient gene silencing by delivery of locked nucleic acid antisense oligonucleotides, unassisted by transfection reagents","volume":"38","author":"Stein","year":"2010","journal-title":"Nucleic Acids Res."},{"key":"10.1016\/j.cell.2026.04.014_bib84","doi-asserted-by":"crossref","first-page":"2114","DOI":"10.1093\/bioinformatics\/btu170","article-title":"Trimmomatic: a flexible trimmer for Illumina sequence data","volume":"30","author":"Bolger","year":"2014","journal-title":"Bioinformatics"},{"key":"10.1016\/j.cell.2026.04.014_bib85","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1093\/bioinformatics\/bts635","article-title":"STAR: ultrafast universal RNA-seq aligner","volume":"29","author":"Dobin","year":"2013","journal-title":"Bioinformatics"},{"key":"10.1016\/j.cell.2026.04.014_bib86","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1009730","article-title":"Improved transcriptome assembly using a hybrid of long and short reads with StringTie","volume":"18","author":"Shumate","year":"2022","journal-title":"PLoS Comput. Biol."},{"key":"10.1016\/j.cell.2026.04.014_bib87","doi-asserted-by":"crossref","first-page":"1840","DOI":"10.1093\/bioinformatics\/btw076","article-title":"SplAdder: identification, quantification and testing of alternative splicing events from RNA-Seq data","volume":"32","author":"Kahles","year":"2016","journal-title":"Bioinformatics"},{"key":"10.1016\/j.cell.2026.04.014_bib88","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1007\/978-1-0716-2293-3_11","article-title":"Identification, Quantification, and Testing of Alternative Splicing Events from RNA-Seq Data Using SplAdder","volume":"2493","author":"Markolin","year":"2022","journal-title":"Methods Mol. Biol."},{"key":"10.1016\/j.cell.2026.04.014_bib89","doi-asserted-by":"crossref","first-page":"923","DOI":"10.1093\/bioinformatics\/btt656","article-title":"featureCounts: an efficient general purpose program for assigning sequence reads to genomic features","volume":"30","author":"Liao","year":"2014","journal-title":"Bioinformatics"},{"key":"10.1016\/j.cell.2026.04.014_bib90","doi-asserted-by":"crossref","DOI":"10.1186\/s13059-014-0550-8","article-title":"Moderated estimation of fold change and dispersion for RNA-seq data with DESeq2","volume":"15","author":"Love","year":"2014","journal-title":"Genome Biol."},{"key":"10.1016\/j.cell.2026.04.014_bib91","doi-asserted-by":"crossref","DOI":"10.1093\/bioinformatics\/btac830","article-title":"igv.js: an embeddable JavaScript implementation of the Integrative Genomics Viewer (IGV)","volume":"39","author":"Robinson","year":"2023","journal-title":"Bioinformatics"},{"key":"10.1016\/j.cell.2026.04.014_bib92","doi-asserted-by":"crossref","DOI":"10.3791\/3912-v","article-title":"Chromatin isolation by RNA purification (ChIRP)","author":"Chu","year":"2012","journal-title":"J. Vis. Exp."},{"key":"10.1016\/j.cell.2026.04.014_bib93","doi-asserted-by":"crossref","first-page":"841","DOI":"10.1093\/bioinformatics\/btq033","article-title":"BEDTools: a flexible suite of utilities for comparing genomic features","volume":"26","author":"Quinlan","year":"2010","journal-title":"Bioinformatics"},{"key":"10.1016\/j.cell.2026.04.014_bib94","article-title":"Discovery and Quantification of Long-Range RNA Base Pairs in Coronavirus Genomes with SEARCH-MaP and SEISMIC-RNA","author":"Allan","year":"2024","journal-title":"bioRxiv"},{"key":"10.1016\/j.cell.2026.04.014_bib95","doi-asserted-by":"crossref","first-page":"1974","DOI":"10.1093\/bioinformatics\/btp250","article-title":"VARNA: Interactive drawing and editing of the RNA secondary structure","volume":"25","author":"Darty","year":"2009","journal-title":"Bioinformatics"},{"key":"10.1016\/j.cell.2026.04.014_bib96","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1007\/978-1-4939-6380-5_7","article-title":"RIP: RNA Immunoprecipitation","volume":"1480","author":"Gagliardi","year":"2016","journal-title":"Methods Mol. 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Med."},{"key":"10.1016\/j.cell.2026.04.014_bib98","doi-asserted-by":"crossref","DOI":"10.1186\/s13059-022-02664-4","article-title":"Predicting RNA splicing from DNA sequence using Pangolin","volume":"23","author":"Zeng","year":"2022","journal-title":"Genome Biol."},{"key":"10.1016\/j.cell.2026.04.014_bib99","doi-asserted-by":"crossref","first-page":"2348","DOI":"10.1126\/science.1067179","article-title":"Resolution of the early placental mammal radiation using Bayesian phylogenetics","volume":"294","author":"Murphy","year":"2001","journal-title":"Science"},{"key":"10.1016\/j.cell.2026.04.014_bib100","doi-asserted-by":"crossref","first-page":"D186","DOI":"10.1093\/nar\/gkac999","article-title":"LncBook 2.0: integrating human long non-coding RNAs with multi-omics annotations","volume":"51","author":"Li","year":"2023","journal-title":"Nucleic Acids Res."},{"key":"10.1016\/j.cell.2026.04.014_bib101","doi-asserted-by":"crossref","first-page":"699","DOI":"10.1038\/s41586-020-2493-4","article-title":"Expanded encyclopaedias of DNA elements in the human and mouse genomes","volume":"583","author":"Moore","year":"2020","journal-title":"Nature"},{"key":"10.1016\/j.cell.2026.04.014_bib102","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1089\/crispr.2018.0014","article-title":"EditR: A Method to Quantify Base Editing from Sanger Sequencing","volume":"1","author":"Kluesner","year":"2018","journal-title":"CRISPR J."},{"key":"10.1016\/j.cell.2026.04.014_bib103","article-title":"Metascape provides a biologist-oriented resource for the analysis of systems-level datasets","volume":"10","author":"Zhou","year":"2019","journal-title":"Nat. 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