RRAGC Gene

Name	Ras-related GTP binding C
Description	This gene encodes a member of the GTR/RAG GTP-binding protein family. The encoded protein is a monomeric guanine nucleotide-binding protein which forms a heterodimer with RRAGA and RRAGB and is primarily localized to the cytoplasm. The encoded protein promotes intracellular localization of the mTOR complex. Alternative splicing results in multiple transcript variants. [provided by RefSeq, Dec 2012]
Summary	{"type": "root", "children": [{"type": "p", "children": [{"type": "t", "text": "\nRagC, encoded by RRAGC, is a key component of the Rag GTPase heterodimer that mediates the recruitment and activation of mTOR complex 1 (mTORC1) in response to amino acid availability. In its functional role, RagC must be in a GDP‐bound state in concert with a GTP‐bound RagA to properly engage mTORC1 at the lysosomal surface, thereby acting as an essential nutrient sensor and signal transducer. Structural and biochemical studies have shown that this nucleotide-dependent configuration is critical for mTORC1 docking and subsequent signaling, ensuring that cellular growth is tightly regulated by nutrient status."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "1", "end_ref": "6"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nDetailed structural investigations have elucidated the mechanistic basis underlying the unique behavior of the Rag heterodimer. These studies reveal that RagC’s GDP-bound status is maintained by distinct interdomain interactions—such as key hydrogen bonds in the C-terminal roadblock domain—and that regulatory proteins like FLCN (with its FNIP partners) act as GAPs for RagC to enforce the proper nucleotide state. In addition, accessory components including lysosomal anchors and amino acid transporters (e.g., SLC38A9) as well as other regulatory factors like the mitochondrial threonyl-tRNA synthetase TARS2 have been implicated in modulating RagC’s activity. Such insights provide a molecular framework for understanding how the nucleotide state of RagC is sensed by mTORC1 components (for example, by the Raptor “claw”) to translate nutrient levels into appropriate cellular responses."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "1"}, {"type": "fg_fs", "start_ref": "7", "end_ref": "11"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nMutations in RRAGC have emerged as significant pathogenic events in human disease. Recurrent somatic mutations in follicular lymphoma and de novo gain-of-function missense changes (such as S75Y) identified in cases of pediatric cardiomyopathy illustrate that aberrant RagC activity renders mTORC1 signaling resistant to amino acid deprivation, leading to unchecked growth signaling. In these settings, the altered RagC function not only drives mTORC1 hyperactivation but also disrupts downstream processes such as TFEB phosphorylation, and may modulate additional pathways through effects on protein homeostasis and oncoprotein stability. Moreover, intercellular mechanisms—including the delivery of cancer-derived microRNAs that target RRAGC—further underscore the gene’s role in the complex metabolic reprogramming seen in tumor-stroma interactions."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "12", "end_ref": "18"}]}, {"type": "t", "text": "\n"}]}, {"type": "rg", "children": [{"type": "r", "ref": 1, "children": [{"type": "t", "text": "Zhi-Yang Tsun, Liron Bar-Peled, Lynne Chantranupong, et al. "}, {"type": "b", "children": [{"type": "t", "text": "The folliculin tumor suppressor is a GAP for the RagC/D GTPases that signal amino acid levels to mTORC1."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Mol Cell (2013)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1016/j.molcel.2013.09.016"}], "href": "https://doi.org/10.1016/j.molcel.2013.09.016"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "24095279"}], "href": "https://pubmed.ncbi.nlm.nih.gov/24095279"}]}, {"type": "r", "ref": 2, "children": [{"type": "t", "text": "Kuang Shen, Abigail Choe, David M Sabatini "}, {"type": "b", "children": [{"type": "t", "text": "Intersubunit Crosstalk in the Rag GTPase Heterodimer Enables mTORC1 to Respond Rapidly to Amino Acid Availability."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Mol Cell (2017)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1016/j.molcel.2017.09.026"}], "href": "https://doi.org/10.1016/j.molcel.2017.09.026"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "29056322"}], "href": "https://pubmed.ncbi.nlm.nih.gov/29056322"}]}, {"type": "r", "ref": 3, "children": [{"type": "t", "text": "Ming-Yuan Su, Kyle L Morris, Do Jin Kim, et al. "}, {"type": "b", "children": [{"type": "t", "text": "Hybrid Structure of the RagA/C-Ragulator mTORC1 Activation Complex."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Mol Cell (2017)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1016/j.molcel.2017.10.016"}], "href": "https://doi.org/10.1016/j.molcel.2017.10.016"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "29107538"}], "href": "https://pubmed.ncbi.nlm.nih.gov/29107538"}]}, {"type": "r", "ref": 4, "children": [{"type": "t", "text": "Kacper B Rogala, Xin Gu, Jibril F Kedir, et al. "}, {"type": "b", "children": [{"type": "t", "text": "Structural basis for the docking of mTORC1 on the lysosomal surface."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Science (2019)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1126/science.aay0166"}], "href": "https://doi.org/10.1126/science.aay0166"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "31601708"}], "href": "https://pubmed.ncbi.nlm.nih.gov/31601708"}]}, {"type": "r", "ref": 5, "children": [{"type": "t", "text": "Madhanagopal Anandapadamanaban, Glenn R Masson, Olga Perisic, et al. 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"}, {"type": "b", "children": [{"type": "t", "text": "Euchromatin histone methyltransferase II (EHMT2) regulates the expression of ras-related GTP binding C (RRAGC) protein."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "BMB Rep (2020)"}]}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "32684241"}], "href": "https://pubmed.ncbi.nlm.nih.gov/32684241"}]}, {"type": "r", "ref": 8, "children": [{"type": "t", "text": "Simon A Fromm, Rosalie E Lawrence, James H Hurley "}, {"type": "b", "children": [{"type": "t", "text": "Structural mechanism for amino acid-dependent Rag GTPase nucleotide state switching by SLC38A9."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Nat Struct Mol Biol (2020)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1038/s41594-020-0490-9"}], "href": "https://doi.org/10.1038/s41594-020-0490-9"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "32868926"}], "href": "https://pubmed.ncbi.nlm.nih.gov/32868926"}]}, {"type": "r", "ref": 9, "children": [{"type": "t", "text": "Shu Yang, Yingbiao Zhang, Chun-Yuan Ting, et al. 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"}, {"type": "b", "children": [{"type": "t", "text": "Structure of the lysosomal mTORC1-TFEB-Rag-Ragulator megacomplex."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Nature (2023)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1038/s41586-022-05652-7"}], "href": "https://doi.org/10.1038/s41586-022-05652-7"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "36697823"}], "href": "https://pubmed.ncbi.nlm.nih.gov/36697823"}]}, {"type": "r", "ref": 12, "children": [{"type": "t", "text": "Jessica Okosun, Rachel L Wolfson, Jun Wang, et al. "}, {"type": "b", "children": [{"type": "t", "text": "Recurrent mTORC1-activating RRAGC mutations in follicular lymphoma."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Nat Genet (2016)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1038/ng.3473"}], "href": "https://doi.org/10.1038/ng.3473"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "26691987"}], "href": "https://pubmed.ncbi.nlm.nih.gov/26691987"}]}, {"type": "r", "ref": 13, "children": [{"type": "t", "text": "Pamela A Long, Michael T Zimmermann, Maengjo Kim, et al. "}, {"type": "b", "children": [{"type": "t", "text": "De novo RRAGC mutation activates mTORC1 signaling in syndromic fetal dilated cardiomyopathy."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Hum Genet (2016)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1007/s00439-016-1685-3"}], "href": "https://doi.org/10.1007/s00439-016-1685-3"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "27234373"}], "href": "https://pubmed.ncbi.nlm.nih.gov/27234373"}]}, {"type": "r", "ref": 14, "children": [{"type": "t", "text": "Zhang Xiao Ying, Meiyan Jin, Luke F Peterson, et al. "}, {"type": "b", "children": [{"type": "t", "text": "Recurrent Mutations in the MTOR Regulator RRAGC in Follicular Lymphoma."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Clin Cancer Res (2016)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1158/1078-0432.CCR-16-0609"}], "href": "https://doi.org/10.1158/1078-0432.CCR-16-0609"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "27267853"}], "href": "https://pubmed.ncbi.nlm.nih.gov/27267853"}]}, {"type": "r", "ref": 15, "children": [{"type": "t", "text": "Sung-Hoon Kim, Jung-Hyun Choi, Peng Wang, et al. "}, {"type": "b", "children": [{"type": "t", "text": "Mitochondrial Threonyl-tRNA Synthetase TARS2 Is Required for Threonine-Sensitive mTORC1 Activation."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Mol Cell (2021)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1016/j.molcel.2020.11.036"}], "href": "https://doi.org/10.1016/j.molcel.2020.11.036"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "33340489"}], "href": "https://pubmed.ncbi.nlm.nih.gov/33340489"}]}, {"type": "r", "ref": 16, "children": [{"type": "t", "text": "Miranda Y Fong, Wei Yan, Majid Ghassemian, et al. "}, {"type": "b", "children": [{"type": "t", "text": "Cancer-secreted miRNAs regulate amino-acid-induced mTORC1 signaling and fibroblast protein synthesis."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "EMBO Rep (2021)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.15252/embr.202051239"}], "href": "https://doi.org/10.15252/embr.202051239"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "33345445"}], "href": "https://pubmed.ncbi.nlm.nih.gov/33345445"}]}, {"type": "r", "ref": 17, "children": [{"type": "t", "text": "Maengjo Kim, Linghui Lu, Alexey V Dvornikov, et al. "}, {"type": "b", "children": [{"type": "t", "text": "TFEB Overexpression, Not mTOR Inhibition, Ameliorates RagC"}, {"type": "a", "children": [{"type": "t", "text": "sup"}], "href": "sup"}, {"type": "t", "text": "S75Y"}, {"type": "a", "children": [{"type": "t", "text": "/sup"}], "href": "/sup"}, {"type": "t", "text": " Cardiomyopathy."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Int J Mol Sci (2021)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.3390/ijms22115494"}], "href": "https://doi.org/10.3390/ijms22115494"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "34071043"}], "href": "https://pubmed.ncbi.nlm.nih.gov/34071043"}]}, {"type": "r", "ref": 18, "children": [{"type": "t", "text": "Yin Shi, Shengfeng Xu, Natalie Y L Ngoi, et al. "}, {"type": "b", "children": [{"type": "t", "text": "Rag GTPases suppress PRL-3 degradation and predict poor clinical diagnosis of cancer patients with low PRL-3 mRNA expression."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Biochem Biophys Res Commun (2021)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1016/j.bbrc.2021.08.090"}], "href": "https://doi.org/10.1016/j.bbrc.2021.08.090"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "34482023"}], "href": "https://pubmed.ncbi.nlm.nih.gov/34482023"}]}]}]}
Synonyms	GTR2, RAGC, TIB929
Proteins	RRAGC_HUMAN
NCBI Gene ID	64121
API
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Predicted Functions
Co-expressed Genes
Expression in Tissues and Cell Lines

Functional Associations

RRAGC has 8,007 functional associations with biological entities spanning 9 categories (molecular profile, organism, chemical, functional term, phrase or reference, disease, phenotype or trait, structural feature, cell line, cell type or tissue, gene, protein or microRNA, sequence feature) extracted from 123 datasets.

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

If available, associations are ranked by standardized value

Harmonizome 3.0

RRAGC 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 RRAGC 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 RRAGC 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 RRAGC 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 RRAGC 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 RRAGC 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 RRAGC 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 RRAGC 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 RRAGC 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 RRAGC 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 RRAGC gene relative to other cell types and tissues from the BioGPS Mouse Cell Type and Tissue Gene Expression Profiles dataset.
	Carcinogenome Chemical Perturbation Carcinogenicity Signatures	small molecule perturbations changing expression of RRAGC gene from the Carcinogenome Chemical Perturbation Carcinogenicity Signatures dataset.
	CCLE Cell Line Gene CNV Profiles	cell lines with high or low copy number of RRAGC 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 RRAGC gene relative to other cell lines from the CCLE Cell Line Gene Expression Profiles dataset.
	CCLE Cell Line Proteomics	Cell lines associated with RRAGC protein from the CCLE Cell Line Proteomics dataset.
	CellMarker Gene-Cell Type Associations	cell types associated with RRAGC 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 RRAGC gene from the CHEA Transcription Factor Binding Site Profiles dataset.
	ChEA Transcription Factor Targets	transcription factors binding the promoter of RRAGC 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 RRAGC gene in low- or high-throughput transcription factor functional studies from the CHEA Transcription Factor Targets 2022 dataset.
	CM4AI KOLF21J CRISPRi Gene Perturbation Atlas	gene perturbations changing expression of RRAGC gene from the CM4AI KOLF21J CRISPRi Gene Perturbation Atlas dataset.
	CM4AI U2OS Cell Map Protein Localization Assemblies	assemblies containing RRAGC 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 RRAGC gene from the CMAP Signatures of Differentially Expressed Genes for Small Molecules dataset.
	COMPARTMENTS Curated Protein Localization Evidence Scores	cellular components containing RRAGC protein from the COMPARTMENTS Curated Protein Localization Evidence Scores dataset.
	COMPARTMENTS Curated Protein Localization Evidence Scores 2025	cellular components containing RRAGC protein from the COMPARTMENTS Curated Protein Localization Evidence Scores 2025 dataset.
	COMPARTMENTS Experimental Protein Localization Evidence Scores	cellular components containing RRAGC protein in low- or high-throughput protein localization assays from the COMPARTMENTS Experimental Protein Localization Evidence Scores dataset.
	COMPARTMENTS Experimental Protein Localization Evidence Scores 2025	cellular components containing RRAGC protein in low- or high-throughput protein localization assays from the COMPARTMENTS Experimental Protein Localization Evidence Scores 2025 dataset.
	COMPARTMENTS Text-mining Protein Localization Evidence Scores	cellular components co-occuring with RRAGC 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 RRAGC 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 RRAGC gene relative to other cell lines from the COSMIC Cell Line Gene CNV Profiles dataset.
	COSMIC Cell Line Gene Mutation Profiles	cell lines with RRAGC gene mutations from the COSMIC Cell Line Gene Mutation Profiles dataset.
	CTD Gene-Chemical Interactions	chemicals interacting with RRAGC gene/protein from the curated CTD Gene-Chemical Interactions dataset.
	CTD Gene-Disease Associations	diseases associated with RRAGC gene/protein from the curated CTD Gene-Disease Associations dataset.
	DeepCoverMOA Drug Mechanisms of Action	small molecule perturbations with high or low expression of RRAGC protein relative to other small molecule perturbations from the DeepCoverMOA Drug Mechanisms of Action dataset.
	DepMap CRISPR Gene Dependency	cell lines with fitness changed by RRAGC gene knockdown relative to other cell lines from the DepMap CRISPR Gene Dependency dataset.
	DISEASES Curated Gene-Disease Association Evidence Scores 2025	diseases involving RRAGC gene from the DISEASES Curated Gene-Disease Association Evidence Scores 2025 dataset.
	DISEASES Experimental Gene-Disease Association Evidence Scores 2025	diseases associated with RRAGC gene in GWAS datasets from the DISEASES Experimental Gene-Disease Assocation Evidence Scores 2025 dataset.
	DISEASES Text-mining Gene-Disease Association Evidence Scores	diseases co-occuring with RRAGC 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 RRAGC 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 RRAGC gene in GWAS and other genetic association datasets from the DisGeNET Gene-Disease Associations dataset.
	ENCODE Histone Modification Site Profiles	histone modification site profiles with high histone modification abundance at RRAGC 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 RRAGC gene from the ENCODE Transcription Factor Binding Site Profiles dataset.