TONSL Gene

HGNC Family	Ankyrin repeat domain containing (ANKRD)
Name	tonsoku-like, DNA repair protein
Description	The protein encoded by this gene is thought to be a negative regulator of NF-kappa-B mediated transcription. The encoded protein may bind NF-kappa-B complexes and trap them in the cytoplasm, preventing them from entering the nucleus and interacting with the DNA. Phosphorylation of this protein targets it for degradation by the ubiquitination pathway, which frees the NF-kappa-B complexes to enter the nucleus. [provided by RefSeq, Jul 2008]
Summary	{"type": "root", "children": [{"type": "p", "children": [{"type": "t", "text": "\nTONSL plays a crucial role in safeguarding genome integrity during DNA replication by acting as a reader of newly incorporated histones. Specifically, the TONSL–MMS22L complex recognizes H4 tails unmethylated at lysine 20 (H4K20me0), a hallmark of post‐replicative chromatin, thereby marking regions where DNA repair via homologous recombination is needed. This recognition promotes the localization of TONSL to challenged replication forks and sites of DNA damage, facilitating the proper loading of RAD51 and ensuring efficient repair of double‐strand breaks that occur following replication stress. The loss or mutation of TONSL’s histone recognition domain is associated with increased replication fork stalling, genomic instability, and impaired cell viability."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "1", "end_ref": "5"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nBeyond its role as a histone reader, TONSL functions in concert with MMS22L to coordinate several key aspects of DNA replication and repair. The TONSL–MMS22L heterodimer not only localizes to replication forks under both unperturbed and stress conditions but also contributes to the establishment of sister chromatid cohesion and the appropriate modulation of the chromatin environment. In this context, the complex assists in the recruitment of repair factors and facilitates RAD51-mediated strand invasion required for homologous recombination. Furthermore, genetic interactions involving associated replication factors underscore the importance of the TONSL–MMS22L complex in maintaining chromosomal stability during replication, with its loss resulting in synthetic lethality when paired with deficiencies in other replication-related pathways."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "6", "end_ref": "8"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nAberrant regulation of TONSL is linked to a spectrum of human diseases. Biallelic hypomorphic variants in TONSL are causative for SPONASTRIME dysplasia, a skeletal dysplasia characterized by short stature, spinal abnormalities, and other developmental defects. In contrast, altered TONSL expression levels have been implicated in cancer: overexpression may promote epithelial cell immortalization, enhance homologous recombination capacity—in part by increasing chromatin accessibility to oncogenic transcription factors—and support the survival of cancer stem cells, thereby contributing to tumor progression and poor patient prognosis in malignancies such as breast, gastric, and hepatocellular carcinoma. In addition, regulatory elements such as the long non-coding RNA TONSL-AS1, which modulate the expression of its neighboring TONSL gene, further highlight the multifaceted roles of TONSL in cellular homeostasis and underscore its potential as a therapeutic target. Emerging evidence also suggests that TONSL (historically annotated as NFKBIL2 in some contexts) may modulate immune responses, adding another layer to its diverse cellular functions."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "9", "end_ref": "16"}]}, {"type": "t", "text": "\n"}]}, {"type": "rg", "children": [{"type": "r", "ref": 1, "children": [{"type": "t", "text": "Giulia Saredi, Hongda Huang, Colin M Hammond, et al. "}, {"type": "b", "children": [{"type": "t", "text": "H4K20me0 marks post-replicative chromatin and recruits the TONSL–MMS22L DNA repair complex."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Nature (2016)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1038/nature18312"}], "href": "https://doi.org/10.1038/nature18312"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "27338793"}], "href": "https://pubmed.ncbi.nlm.nih.gov/27338793"}]}, {"type": "r", "ref": 2, "children": [{"type": "t", "text": "Lara O'Donnell, Stephanie Panier, Jan Wildenhain, et al. "}, {"type": "b", "children": [{"type": "t", "text": "The MMS22L-TONSL complex mediates recovery from replication stress and homologous recombination."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Mol Cell (2010)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1016/j.molcel.2010.10.024"}], "href": "https://doi.org/10.1016/j.molcel.2010.10.024"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "21055983"}], "href": "https://pubmed.ncbi.nlm.nih.gov/21055983"}]}, {"type": "r", "ref": 3, "children": [{"type": "t", "text": "Brenda C O'Connell, Britt Adamson, John R Lydeard, et al. "}, {"type": "b", "children": [{"type": "t", "text": "A genome-wide camptothecin sensitivity screen identifies a mammalian MMS22L-NFKBIL2 complex required for genomic stability."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Mol Cell (2010)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1016/j.molcel.2010.10.022"}], "href": "https://doi.org/10.1016/j.molcel.2010.10.022"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "21055985"}], "href": "https://pubmed.ncbi.nlm.nih.gov/21055985"}]}, {"type": "r", "ref": 4, "children": [{"type": "t", "text": "Eris Duro, Cecilia Lundin, Katrine Ask, et al. "}, {"type": "b", "children": [{"type": "t", "text": "Identification of the MMS22L-TONSL complex that promotes homologous recombination."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Mol Cell (2010)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1016/j.molcel.2010.10.023"}], "href": "https://doi.org/10.1016/j.molcel.2010.10.023"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "21055984"}], "href": "https://pubmed.ncbi.nlm.nih.gov/21055984"}]}, {"type": "r", "ref": 5, "children": [{"type": "t", "text": "Ting-Hsiang Huang, Faith Fowler, Chin-Chuan Chen, et al. "}, {"type": "b", "children": [{"type": "t", "text": "The Histone Chaperones ASF1 and CAF-1 Promote MMS22L-TONSL-Mediated Rad51 Loading onto ssDNA during Homologous Recombination in Human Cells."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Mol Cell (2018)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1016/j.molcel.2018.01.031"}], "href": "https://doi.org/10.1016/j.molcel.2018.01.031"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "29478807"}], "href": "https://pubmed.ncbi.nlm.nih.gov/29478807"}]}, {"type": "r", "ref": 6, "children": [{"type": "t", "text": "Wojciech Piwko, Michael H Olma, Michael Held, et al. "}, {"type": "b", "children": [{"type": "t", "text": "RNAi-based screening identifies the Mms22L-Nfkbil2 complex as a novel regulator of DNA replication in human cells."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "EMBO J (2010)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1038/emboj.2010.304"}], "href": "https://doi.org/10.1038/emboj.2010.304"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "21113133"}], "href": "https://pubmed.ncbi.nlm.nih.gov/21113133"}]}, {"type": "r", "ref": 7, "children": [{"type": "t", "text": "Wojciech Piwko, Lucie J Mlejnkova, Karun Mutreja, et al. "}, {"type": "b", "children": [{"type": "t", "text": "The MMS22L-TONSL heterodimer directly promotes RAD51-dependent recombination upon replication stress."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "EMBO J (2016)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.15252/embj.201593132"}], "href": "https://doi.org/10.15252/embj.201593132"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "27797818"}], "href": "https://pubmed.ncbi.nlm.nih.gov/27797818"}]}, {"type": "r", "ref": 8, "children": [{"type": "t", "text": "Janne Jm van Schie, Klaas de Lint, Govind M Pai, et al. "}, {"type": "b", "children": [{"type": "t", "text": "MMS22L-TONSL functions in sister chromatid cohesion in a pathway parallel to DSCC1-RFC."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Life Sci Alliance (2022)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.26508/lsa.202201596"}], "href": "https://doi.org/10.26508/lsa.202201596"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "36622344"}], "href": "https://pubmed.ncbi.nlm.nih.gov/36622344"}]}, {"type": "r", "ref": 9, "children": [{"type": "t", "text": "Lindsay C Burrage, John J Reynolds, Nissan Vida Baratang, et al. "}, {"type": "b", "children": [{"type": "t", "text": "Bi-allelic Variants in TONSL Cause SPONASTRIME Dysplasia and a Spectrum of Skeletal Dysplasia Phenotypes."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Am J Hum Genet (2019)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1016/j.ajhg.2019.01.007"}], "href": "https://doi.org/10.1016/j.ajhg.2019.01.007"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "30773277"}], "href": "https://pubmed.ncbi.nlm.nih.gov/30773277"}]}, {"type": "r", "ref": 10, "children": [{"type": "t", "text": "Stephen J Chapman, Chiea C Khor, Fredrik O Vannberg, et al. "}, {"type": "b", "children": [{"type": "t", "text": "Common NFKBIL2 polymorphisms and susceptibility to pneumococcal disease: a genetic association study."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Crit Care (2010)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1186/cc9377"}], "href": "https://doi.org/10.1186/cc9377"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "21171993"}], "href": "https://pubmed.ncbi.nlm.nih.gov/21171993"}]}, {"type": "r", "ref": 11, "children": [{"type": "t", "text": "Bin Yu, Youming Ding, Xiaofeng Liao, et al. "}, {"type": "b", "children": [{"type": "t", "text": "Overexpression of TONSL might be an independent unfavorable prognostic indicator in hepatocellular carcinoma."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Pathol Res Pract (2019)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1016/j.prp.2019.01.044"}], "href": "https://doi.org/10.1016/j.prp.2019.01.044"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "30723051"}], "href": "https://pubmed.ncbi.nlm.nih.gov/30723051"}]}, {"type": "r", "ref": 12, "children": [{"type": "t", "text": "Hae Ryung Chang, Sung Yoon Cho, Jae Hoon Lee, et al. "}, {"type": "b", "children": [{"type": "t", "text": "Hypomorphic Mutations in TONSL Cause SPONASTRIME Dysplasia."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Am J Hum Genet (2019)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1016/j.ajhg.2019.01.009"}], "href": "https://doi.org/10.1016/j.ajhg.2019.01.009"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "30773278"}], "href": "https://pubmed.ncbi.nlm.nih.gov/30773278"}]}, {"type": "r", "ref": 13, "children": [{"type": "t", "text": "Aditi S Khatpe, Rebecca Dirks, Poornima Bhat-Nakshatri, et al. 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"}, {"type": "b", "children": [{"type": "t", "text": "Novel TONSL variants cause SPONASTRIME dysplasia and associate with spontaneous chromosome breaks, defective cell proliferation and apoptosis."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Hum Mol Genet (2020)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1093/hmg/ddaa195"}], "href": "https://doi.org/10.1093/hmg/ddaa195"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "32959051"}], "href": "https://pubmed.ncbi.nlm.nih.gov/32959051"}]}]}]}
Synonyms	NFKBIL2, IKBR
Proteins	TONSL_HUMAN
NCBI Gene ID	4796
API
Download Associations
Predicted Functions
Co-expressed Genes
Expression in Tissues and Cell Lines

Functional Associations

TONSL has 5,445 functional associations with biological entities spanning 9 categories (molecular profile, organism, disease, phenotype or trait, functional term, phrase or reference, chemical, structural feature, cell line, cell type or tissue, gene, protein or microRNA, sequence feature) extracted from 106 datasets.

Click the + buttons to view associations for TONSL from the datasets below.

If available, associations are ranked by standardized value

Harmonizome 3.0

TONSL Gene

Functional Associations

Please acknowledge the Harmonizome in your publications by citing the following reference(s):

	Dataset	Summary
	Achilles Cell Line Gene Essentiality Profiles	cell lines with fitness changed by TONSL gene knockdown relative to other cell lines from the Achilles Cell Line Gene Essentiality Profiles dataset.
	Allen Brain Atlas Adult Human Brain Tissue Gene Expression Profiles	tissues with high or low expression of TONSL gene relative to other tissues from the Allen Brain Atlas Adult Human Brain Tissue Gene Expression Profiles dataset.
	Allen Brain Atlas Adult Mouse Brain Tissue Gene Expression Profiles	tissues with high or low expression of TONSL gene relative to other tissues from the Allen Brain Atlas Adult Mouse Brain Tissue Gene Expression Profiles dataset.
	Allen Brain Atlas Aging Dementia and Traumatic Brain Injury Tissue Sample Gene Expression Profiles	tissue samples with high or low expression of TONSL gene relative to other tissue samples from the Allen Brain Atlas Aging Dementia and Traumatic Brain Injury Tissue Sample Gene Expression Profiles dataset.
	Allen Brain Atlas Developing Human Brain Tissue Gene Expression Profiles by Microarray	tissue samples with high or low expression of TONSL gene relative to other tissue samples from the Allen Brain Atlas Developing Human Brain Tissue Gene Expression Profiles by Microarray dataset.
	Allen Brain Atlas Developing Human Brain Tissue Gene Expression Profiles by RNA-seq	tissue samples with high or low expression of TONSL gene relative to other tissue samples from the Allen Brain Atlas Developing Human Brain Tissue Gene Expression Profiles by RNA-seq dataset.
	Allen Brain Atlas Prenatal Human Brain Tissue Gene Expression Profiles	tissues with high or low expression of TONSL gene relative to other tissues from the Allen Brain Atlas Prenatal Human Brain Tissue Gene Expression Profiles dataset.
	BioGPS Cell Line Gene Expression Profiles	cell lines with high or low expression of TONSL gene relative to other cell lines from the BioGPS Cell Line Gene Expression Profiles dataset.
	BioGPS Human Cell Type and Tissue Gene Expression Profiles	cell types and tissues with high or low expression of TONSL gene relative to other cell types and tissues from the BioGPS Human Cell Type and Tissue Gene Expression Profiles dataset.
	BioGPS Mouse Cell Type and Tissue Gene Expression Profiles	cell types and tissues with high or low expression of TONSL gene relative to other cell types and tissues from the BioGPS Mouse Cell Type and Tissue Gene Expression Profiles dataset.
	CCLE Cell Line Gene CNV Profiles	cell lines with high or low copy number of TONSL gene relative to other cell lines from the CCLE Cell Line Gene CNV Profiles dataset.
	CCLE Cell Line Gene Expression Profiles	cell lines with high or low expression of TONSL gene relative to other cell lines from the CCLE Cell Line Gene Expression Profiles dataset.
	CCLE Cell Line Proteomics	Cell lines associated with TONSL protein from the CCLE Cell Line Proteomics dataset.
	CellMarker Gene-Cell Type Associations	cell types associated with TONSL gene from the CellMarker Gene-Cell Type Associations dataset.
	ChEA Transcription Factor Binding Site Profiles	transcription factor binding site profiles with transcription factor binding evidence at the promoter of TONSL gene from the CHEA Transcription Factor Binding Site Profiles dataset.
	ChEA Transcription Factor Targets	transcription factors binding the promoter of TONSL gene in low- or high-throughput transcription factor functional studies from the CHEA Transcription Factor Targets dataset.
	ChEA Transcription Factor Targets 2022	transcription factors binding the promoter of TONSL gene in low- or high-throughput transcription factor functional studies from the CHEA Transcription Factor Targets 2022 dataset.
	ClinVar Gene-Phenotype Associations 2025	phenotypes associated with TONSL gene from the curated ClinVar Gene-Phenotype Associations 2025 dataset.
	CM4AI U2OS Cell Map Protein Localization Assemblies	assemblies containing TONSL protein from integrated AP-MS and IF data from the CM4AI U2OS Cell Map Protein Localization Assemblies dataset.
	CMAP Signatures of Differentially Expressed Genes for Small Molecules	small molecule perturbations changing expression of TONSL gene from the CMAP Signatures of Differentially Expressed Genes for Small Molecules dataset.
	COMPARTMENTS Curated Protein Localization Evidence Scores	cellular components containing TONSL protein from the COMPARTMENTS Curated Protein Localization Evidence Scores dataset.
	COMPARTMENTS Curated Protein Localization Evidence Scores 2025	cellular components containing TONSL protein from the COMPARTMENTS Curated Protein Localization Evidence Scores 2025 dataset.
	COMPARTMENTS Experimental Protein Localization Evidence Scores	cellular components containing TONSL protein in low- or high-throughput protein localization assays from the COMPARTMENTS Experimental Protein Localization Evidence Scores dataset.
	COMPARTMENTS Text-mining Protein Localization Evidence Scores	cellular components co-occuring with TONSL protein in abstracts of biomedical publications from the COMPARTMENTS Text-mining Protein Localization Evidence Scores dataset.
	COMPARTMENTS Text-mining Protein Localization Evidence Scores 2025	cellular components co-occuring with TONSL protein in abstracts of biomedical publications from the COMPARTMENTS Text-mining Protein Localization Evidence Scores 2025 dataset.
	COSMIC Cell Line Gene CNV Profiles	cell lines with high or low copy number of TONSL gene relative to other cell lines from the COSMIC Cell Line Gene CNV Profiles dataset.
	COSMIC Cell Line Gene Mutation Profiles	cell lines with TONSL gene mutations from the COSMIC Cell Line Gene Mutation Profiles dataset.
	CTD Gene-Chemical Interactions	chemicals interacting with TONSL gene/protein from the curated CTD Gene-Chemical Interactions dataset.
	CTD Gene-Disease Associations	diseases associated with TONSL gene/protein from the curated CTD Gene-Disease Associations dataset.
	DeepCoverMOA Drug Mechanisms of Action	small molecule perturbations with high or low expression of TONSL protein relative to other small molecule perturbations from the DeepCoverMOA Drug Mechanisms of Action dataset.
	DISEASES Text-mining Gene-Disease Association Evidence Scores	diseases co-occuring with TONSL gene in abstracts of biomedical publications from the DISEASES Text-mining Gene-Disease Assocation Evidence Scores dataset.
	DISEASES Text-mining Gene-Disease Association Evidence Scores 2025	diseases co-occuring with TONSL gene in abstracts of biomedical publications from the DISEASES Text-mining Gene-Disease Assocation Evidence Scores 2025 dataset.
	DisGeNET Gene-Disease Associations	diseases associated with TONSL gene in GWAS and other genetic association datasets from the DisGeNET Gene-Disease Associations dataset.
	DisGeNET Gene-Phenotype Associations	phenotypes associated with TONSL gene in GWAS and other genetic association datasets from the DisGeNET Gene-Phenoptype Associations dataset.
	ENCODE Histone Modification Site Profiles	histone modification site profiles with high histone modification abundance at TONSL gene from the ENCODE Histone Modification Site Profiles dataset.
	ENCODE Transcription Factor Binding Site Profiles	transcription factor binding site profiles with transcription factor binding evidence at the promoter of TONSL gene from the ENCODE Transcription Factor Binding Site Profiles dataset.
	ENCODE Transcription Factor Targets	transcription factors binding the promoter of TONSL gene in ChIP-seq datasets from the ENCODE Transcription Factor Targets dataset.
	ESCAPE Omics Signatures of Genes and Proteins for Stem Cells	PubMedIDs of publications reporting gene signatures containing TONSL from the ESCAPE Omics Signatures of Genes and Proteins for Stem Cells dataset.
	GAD Gene-Disease Associations	diseases associated with TONSL gene in GWAS and other genetic association datasets from the GAD Gene-Disease Associations dataset.
	GAD High Level Gene-Disease Associations	diseases associated with TONSL gene in GWAS and other genetic association datasets from the GAD High Level Gene-Disease Associations dataset.