{"gene":"REEP6","run_date":"2026-04-28T19:45:45","timeline":{"discoveries":[{"year":2014,"finding":"REEP6 expression in rod photoreceptors is regulated by the bZIP transcription factor NRL, which binds within Reep6 intron 1 to drive expression of the retina-specific isoform REEP6.1 via an intronic enhancer sequence.","method":"Chromatin immunoprecipitation (ChIP) assay identifying NRL binding in Reep6 intron 1; reporter assays in cultured cells; retinal explant transfections mapping the intronic enhancer; exon-specific Taqman assay and 5'-RACE for isoform identification","journal":"Human molecular genetics","confidence":"High","confidence_rationale":"Tier 1-2 — multiple orthogonal methods (ChIP, reporter assay, retinal explant transfection) in a single study with rigorous controls","pmids":["24691551"],"is_preprint":false},{"year":2016,"finding":"REEP6 is a member of the REEP/Yop1 family of ER-shaping proteins expressed in the retina; a retina-specific isoform REEP6.1 (containing 27 additional amino acids from exon 5) is specifically expressed in rod photoreceptors, and missense variants (p.Pro128Leu, p.Leu135Pro) and a frameshift in REEP6.1 destabilize the protein in cultured cells.","method":"Expression analysis in human 3D organoid optic cups; expression of mutant proteins in cultured cells; CRISPR-Cas9 knock-in mouse model with p.Leu135Pro variant showing progressive photoreceptor degeneration","journal":"American journal of human genetics","confidence":"High","confidence_rationale":"Tier 1-2 — orthogonal methods including organoid expression, cell-based protein stability assays, and in vivo knock-in mouse model","pmids":["27889058"],"is_preprint":false},{"year":2017,"finding":"REEP6 localizes to the inner segment and outer plexiform layer of rod photoreceptors; loss of REEP6 causes expansion of distal ER and increased mitochondria in rod inner segments, nearly undetectable expression of retinal guanylate cyclases GC1 and GC2, increased CHOP expression and caspase-12 activation indicating ER stress-mediated apoptosis, but does not affect rhodopsin, Rom1, or peripherin/rds trafficking.","method":"CRISPR/Cas9 Reep6 knockout mouse; transmission electron microscopy and 3View serial block-face scanning EM; electroretinography; immunohistochemistry; Western blot for GC1, GC2, CHOP, caspase-12","journal":"Human molecular genetics","confidence":"High","confidence_rationale":"Tier 1-2 — clean knockout with multiple orthogonal structural and molecular readouts; replicated in independent study","pmids":["28475715"],"is_preprint":false},{"year":2017,"finding":"REEP6 is detected in a subset of Clathrin-coated vesicles and physically interacts with the t-SNARE Syntaxin3, implicating it in cargo trafficking via clathrin-coated vesicles to selected membrane sites in rod photoreceptors.","method":"Reep6-/- mouse model; immunofluorescence co-localization of REEP6 with clathrin-coated vesicle markers; co-immunoprecipitation of REEP6 with Syntaxin3","journal":"Human molecular genetics","confidence":"High","confidence_rationale":"Tier 2 — reciprocal Co-IP with co-localization in a defined KO model with photoreceptor degeneration phenotype","pmids":["28369466"],"is_preprint":false},{"year":2016,"finding":"REEP6 interacts with CXCR1 (but not CXCR2), and overexpression of REEP6 enhances IL-8-stimulated cellular responses through CXCR1, while depletion impairs ligand-stimulated receptor internalization and β-arrestin2 intracellular clustering without affecting basal plasma membrane expression of CXCR1.","method":"Co-immunoprecipitation; overexpression and siRNA knockdown in cultured cells; β-arrestin2 clustering assay; ERK phosphorylation measurement; in vivo xenograft model","journal":"Scientific reports","confidence":"Medium","confidence_rationale":"Tier 2-3 — Co-IP plus functional cellular assays from a single lab","pmids":["27966653"],"is_preprint":false},{"year":2021,"finding":"The Reep6.1 isoform (containing exon 5) is the only isoform expressed in adult rod photoreceptors and is functionally distinct from Reep6.2; isoform-specific knockout (deletion of exon 5) causes rod degeneration equivalent to full knockout, and overexpression of Reep6.2 fails to rescue the knockout phenotype while Reep6.1 overexpression does rescue it.","method":"Isoform-specific knockout mouse (Reep6E5/E5); AAV-mediated overexpression rescue experiments in Reep6 knockout mice; electroretinography; histology","journal":"Human molecular genetics","confidence":"High","confidence_rationale":"Tier 1-2 — isoform-specific KO with rescue experiments using overexpression of each isoform, multiple orthogonal functional readouts","pmids":["34104971"],"is_preprint":false},{"year":2022,"finding":"REEP6 plays a role in β-adrenergic signal transduction in adipocytes by facilitating the expression and trafficking of adenylate cyclase 3 (ADCY3) to the plasma membrane; REEP6 KO reduces ADCY3 plasma membrane targeting and severely impairs protein kinase A-mediated signaling in brown adipose tissue.","method":"REEP6 knockout mouse model; adipocyte-specific in vitro knockdown in C3H10T1/2 and primary adipocytes; confocal and electron microscopy; Western blot; indirect calorimetry; glucose tolerance test; high-fat diet model","journal":"Metabolism: clinical and experimental","confidence":"High","confidence_rationale":"Tier 1-2 — in vivo KO with cell-autonomous validation in primary adipocytes, multiple orthogonal methods including imaging of ADCY3 membrane localization","pmids":["35150731"],"is_preprint":false},{"year":2022,"finding":"miR-672-5p directly regulates REEP6 expression in spinal dorsal horn neurons; downregulation of miR-672-5p increases REEP6 levels and contributes to neuronal hyperexcitability and neuropathic pain, as demonstrated by direct interaction between miR-672-5p and REEP6 mRNA 3'UTR.","method":"miRNA microarray; intrathecal injection of miR-672-5p agomir/antagomir; dual-luciferase reporter assay confirming miR-672-5p binding sites in REEP6 mRNA; electrophysiology (sEPSC recording); behavioral pain assays","journal":"Neurochemical research","confidence":"Medium","confidence_rationale":"Tier 2 — luciferase reporter assay for direct miRNA-mRNA interaction validated with in vivo functional experiments","pmids":["36064821"],"is_preprint":false},{"year":2026,"finding":"WTAP-mediated m6A methylation facilitates translation of REEP6 (along with PDE6B and RDH12) in rod photoreceptors through m6A deposition in the 3'UTR of Reep6 mRNA; loss of WTAP abolishes m6A modification and reduces REEP6 protein expression, contributing to retinal degeneration.","method":"Conditional Wtap knockout mouse; m6A sequencing; polysome profiling/ribosome association assays; AAV-mediated WTAP re-expression rescue; CAG-Wtap transgenic cross","journal":"Science China. Life sciences","confidence":"High","confidence_rationale":"Tier 1-2 — m6A epitranscriptomic mechanism with in vivo KO, rescue experiments, and mechanistic identification of 3'UTR m6A sites","pmids":["41796262"],"is_preprint":false},{"year":2025,"finding":"In vitro expression of REEP6 alters ER marker distribution and Golgi morphology; Reep6 knockout mice show reduced expression of multiple rod phototransduction proteins (PDE6, guanylate cyclases, rhodopsin, GRK1) at the transcript level, and activation of inflammatory pathways.","method":"Novel Reep6 knockout mouse (exons 2-5 deletion); in vitro REEP6 expression with ER/Golgi marker imaging; RNA-seq; Western blot","journal":"bioRxiv","confidence":"Medium","confidence_rationale":"Tier 2-3 — preprint with in vitro ER/Golgi morphology assay plus in vivo KO RNA-seq, single lab","pmids":["bio_10.1101_2025.03.02.641069"],"is_preprint":true}],"current_model":"REEP6 is a rod photoreceptor-enriched ER-shaping protein of the REEP/Yop1 family whose retina-specific isoform (REEP6.1) is transcriptionally driven by the NRL transcription factor and whose expression is post-transcriptionally regulated by WTAP-mediated m6A methylation; it localizes to the inner segment ER and a subset of Clathrin-coated vesicles where it interacts with Syntaxin3 to mediate trafficking of cargo (including guanylate cyclases GC1/GC2 and adenylate cyclase ADCY3 in adipocytes) to the plasma membrane, such that its loss causes ER stress, disrupted ER and Golgi morphology, loss of phototransduction proteins, and progressive rod photoreceptor degeneration."},"narrative":{"teleology":[{"year":2014,"claim":"Identifying how REEP6 expression is restricted to rod photoreceptors resolved the transcriptional logic linking NRL, a master rod-fate transcription factor, to ER-shaping gene expression via an intronic enhancer that drives the retina-specific REEP6.1 isoform.","evidence":"ChIP of NRL at Reep6 intron 1, reporter assays, retinal explant transfection, and 5′-RACE isoform mapping","pmids":["24691551"],"confidence":"High","gaps":["Whether other rod-enriched transcription factors co-regulate REEP6.1","Chromatin accessibility dynamics at the intronic enhancer during photoreceptor differentiation"]},{"year":2016,"claim":"Demonstration that human disease-associated REEP6 missense variants destabilize the protein and cause progressive photoreceptor degeneration in a knock-in mouse established REEP6 as a retinitis pigmentosa gene and linked protein stability to rod survival.","evidence":"Patient variant analysis; mutant protein expression in cultured cells; CRISPR knock-in mouse (p.Leu135Pro) with ERG and histology","pmids":["27889058"],"confidence":"High","gaps":["Structural basis for destabilization by P128L and L135P mutations","Whether protein stabilization strategies can slow degeneration"]},{"year":2017,"claim":"Knockout studies resolved the subcellular function of REEP6 by showing that it shapes inner segment ER, is required for GC1/GC2 expression, and localizes to clathrin-coated vesicles where it interacts with Syntaxin3 — establishing a model in which REEP6 links ER morphology to vesicular cargo trafficking in rods.","evidence":"CRISPR Reep6 KO mouse; TEM and serial block-face SEM; immunohistochemistry for GC1/GC2, CHOP, caspase-12; co-IP of REEP6 with Syntaxin3; co-localization with clathrin-coated vesicle markers","pmids":["28475715","28369466"],"confidence":"High","gaps":["Whether REEP6 directly shapes ER tubules like other REEP/Yop1 family members or acts primarily through vesicular trafficking","Mechanism by which GC1/GC2 are specifically lost while rhodopsin trafficking is unaffected","Whether the Syntaxin3 interaction is direct or bridged"]},{"year":2021,"claim":"Isoform-specific knockout and rescue experiments demonstrated that REEP6.1 (containing exon 5) is the sole functional isoform in adult rods and is non-redundant with REEP6.2, resolving a key question about isoform specificity.","evidence":"Reep6 exon 5-specific KO mouse; AAV-mediated overexpression of REEP6.1 vs. REEP6.2 in Reep6 KO retina; ERG; histology","pmids":["34104971"],"confidence":"High","gaps":["What structural feature of the exon 5-encoded 27 amino acids confers rod-specific function","Whether REEP6.2 has a function in other tissues"]},{"year":2022,"claim":"Discovery that REEP6 facilitates ADCY3 plasma membrane targeting in brown adipocytes extended its trafficking role beyond the retina and linked it to β-adrenergic signaling and energy metabolism.","evidence":"REEP6 KO mouse on high-fat diet; adipocyte knockdown; confocal and electron microscopy of ADCY3 localization; indirect calorimetry","pmids":["35150731"],"confidence":"High","gaps":["Whether REEP6 directly binds ADCY3 or acts through an intermediary","Generality of REEP6 trafficking function across additional GPCR-coupled effectors"]},{"year":2022,"claim":"Identification of miR-672-5p as a direct post-transcriptional repressor of REEP6 in dorsal horn neurons revealed a non-retinal regulatory axis and implicated REEP6 upregulation in neuropathic pain-associated neuronal hyperexcitability.","evidence":"Dual-luciferase reporter assay for miR-672-5p/REEP6 3′UTR; intrathecal agomir/antagomir injection; electrophysiology and behavioral pain assays","pmids":["36064821"],"confidence":"Medium","gaps":["Whether REEP6 upregulation is causally sufficient for hyperexcitability or is one of multiple miR-672-5p targets","Replication in independent pain models"]},{"year":2026,"claim":"Showing that WTAP-mediated m6A methylation of the Reep6 3′UTR promotes its translation established epitranscriptomic regulation as a mechanism controlling REEP6 protein levels in rod photoreceptors.","evidence":"Conditional Wtap KO mouse; m6A-seq; polysome profiling; AAV-mediated WTAP rescue","pmids":["41796262"],"confidence":"High","gaps":["Identity of the m6A reader(s) that mediate REEP6 translational enhancement","Relative contribution of transcriptional (NRL) versus epitranscriptomic (WTAP/m6A) regulation to total REEP6 protein levels"]},{"year":null,"claim":"It remains unknown whether REEP6 directly tubulates ER membranes like canonical REEP/Yop1 family members, what structural feature of the exon 5-encoded segment confers isoform-specific function, and how cargo selectivity (e.g., GC1/GC2 but not rhodopsin) is achieved at the molecular level.","evidence":"","pmids":[],"confidence":"High","gaps":["No in vitro reconstitution of REEP6 membrane-shaping activity","No high-resolution structure of REEP6 or the REEP6–Syntaxin3 complex","Mechanism of selective cargo recognition unresolved"]}],"mechanism_profile":{"molecular_activity":[{"term_id":"GO:0005198","term_label":"structural molecule activity","supporting_discovery_ids":[2,9]},{"term_id":"GO:0060090","term_label":"molecular adaptor activity","supporting_discovery_ids":[3,6]}],"localization":[{"term_id":"GO:0005783","term_label":"endoplasmic reticulum","supporting_discovery_ids":[2,9]},{"term_id":"GO:0031410","term_label":"cytoplasmic vesicle","supporting_discovery_ids":[3]}],"pathway":[{"term_id":"R-HSA-9609507","term_label":"Protein localization","supporting_discovery_ids":[3,6]},{"term_id":"R-HSA-5653656","term_label":"Vesicle-mediated transport","supporting_discovery_ids":[3,6]},{"term_id":"R-HSA-162582","term_label":"Signal Transduction","supporting_discovery_ids":[6]},{"term_id":"R-HSA-1852241","term_label":"Organelle biogenesis and maintenance","supporting_discovery_ids":[2,9]}],"complexes":[],"partners":["STX3","ADCY3","NRL","CXCR1"],"other_free_text":[]},"mechanistic_narrative":"REEP6 is a rod photoreceptor-enriched ER-shaping protein that facilitates the trafficking of transmembrane cargo from the endoplasmic reticulum to the plasma membrane, and whose loss causes ER stress, photoreceptor degeneration, and impaired signaling in multiple cell types. The retina-specific isoform REEP6.1, driven by NRL binding at an intronic enhancer, is the sole functional isoform in adult rods; isoform-specific deletion phenocopies full knockout, and only REEP6.1 overexpression rescues rod degeneration [PMID:24691551, PMID:34104971]. REEP6 localizes to the inner segment ER and a subset of clathrin-coated vesicles where it interacts with the t-SNARE Syntaxin3; its absence leads to near-complete loss of retinal guanylate cyclases GC1/GC2, ER expansion, and ER stress-mediated apoptosis in rods [PMID:28475715, PMID:28369466]. Beyond the retina, REEP6 promotes plasma membrane targeting of adenylate cyclase 3 (ADCY3) in brown adipocytes, coupling ER-to-surface trafficking to β-adrenergic/PKA signaling and thermogenesis [PMID:35150731]."},"prefetch_data":{"uniprot":{"accession":"Q96HR9","full_name":"Receptor expression-enhancing protein 6","aliases":["Polyposis locus protein 1-like 1"],"length_aa":211,"mass_kda":23.4,"function":"Required for correct function and survival of retinal photoreceptors (PubMed:27889058). Required for retinal development (By similarity). In rod photoreceptors, facilitates stability and/or trafficking of guanylate cyclases and is required to maintain endoplasmic reticulum and mitochondrial homeostasis (By similarity). May play a role in clathrin-coated intracellular vesicle trafficking of proteins from the endoplasmic reticulum to the retinal rod plasma membrane (By similarity)","subcellular_location":"Endoplasmic reticulum membrane; Cytoplasmic vesicle, clathrin-coated vesicle membrane","url":"https://www.uniprot.org/uniprotkb/Q96HR9/entry"},"depmap":{"release":"DepMap","has_data":true,"is_common_essential":false,"resolved_as":"","url":"https://depmap.org/portal/gene/REEP6","classification":"Not Classified","n_dependent_lines":5,"n_total_lines":1208,"dependency_fraction":0.0041390728476821195},"opencell":{"profiled":true,"resolved_as":"","ensg_id":"ENSG00000115255","cell_line_id":"CID001525","localizations":[{"compartment":"er","grade":3}],"interactors":[{"gene":"RTN3","stoichiometry":0.2}],"url":"https://opencell.sf.czbiohub.org/target/CID001525","total_profiled":1310},"omim":[{"mim_id":"617304","title":"RETINITIS PIGMENTOSA 77; RP77","url":"https://www.omim.org/entry/617304"},{"mim_id":"609346","title":"RECEPTOR EXPRESSION-ENHANCING PROTEIN 6; REEP6","url":"https://www.omim.org/entry/609346"},{"mim_id":"268000","title":"RETINITIS PIGMENTOSA; RP","url":"https://www.omim.org/entry/268000"},{"mim_id":"162080","title":"NEURAL RETINA LEUCINE ZIPPER; NRL","url":"https://www.omim.org/entry/162080"}],"hpa":{"profiled":true,"resolved_as":"","reliability":"Supported","locations":[{"location":"Endoplasmic reticulum","reliability":"Supported"},{"location":"Acrosome","reliability":"Additional"}],"tissue_specificity":"Tissue enhanced","tissue_distribution":"Detected in many","driving_tissues":[{"tissue":"intestine","ntpm":204.5},{"tissue":"liver","ntpm":523.4},{"tissue":"testis","ntpm":363.6}],"url":"https://www.proteinatlas.org/search/REEP6"},"hgnc":{"alias_symbol":["DP1L1","FLJ25383","Yip2f","TB1"],"prev_symbol":["C19orf32"]},"alphafold":{"accession":"Q96HR9","domains":[{"cath_id":"-","chopping":"2-51","consensus_level":"medium","plddt":82.2656,"start":2,"end":51},{"cath_id":"1.10.287","chopping":"54-125","consensus_level":"medium","plddt":72.9435,"start":54,"end":125}],"viewer_url":"https://alphafold.ebi.ac.uk/entry/Q96HR9","model_url":"https://alphafold.ebi.ac.uk/files/AF-Q96HR9-F1-model_v6.cif","pae_url":"https://alphafold.ebi.ac.uk/files/AF-Q96HR9-F1-predicted_aligned_error_v6.png","plddt_mean":73.69},"mouse_models":{"mgi_url":"https://www.informatics.jax.org/marker/summary?nomen=REEP6","jax_strain_url":"https://www.jax.org/strain/search?query=REEP6"},"sequence":{"accession":"Q96HR9","fasta_url":"https://rest.uniprot.org/uniprotkb/Q96HR9.fasta","uniprot_url":"https://www.uniprot.org/uniprotkb/Q96HR9/entry","alphafold_viewer_url":"https://alphafold.ebi.ac.uk/entry/Q96HR9"}},"corpus_meta":[{"pmid":"21946354","id":"PMC_21946354","title":"Identification of a functional transposon insertion in the maize domestication gene tb1.","date":"2011","source":"Nature genetics","url":"https://pubmed.ncbi.nlm.nih.gov/21946354","citation_count":543,"is_preprint":false},{"pmid":"16642024","id":"PMC_16642024","title":"A distant upstream enhancer at the maize domestication gene tb1 has pleiotropic effects on plant and inflorescent architecture.","date":"2006","source":"Nature genetics","url":"https://pubmed.ncbi.nlm.nih.gov/16642024","citation_count":279,"is_preprint":false},{"pmid":"14701910","id":"PMC_14701910","title":"Pattern of diversity in the genomic region near the maize domestication gene tb1.","date":"2003","source":"Proceedings of the National Academy of Sciences of the United States of America","url":"https://pubmed.ncbi.nlm.nih.gov/14701910","citation_count":186,"is_preprint":false},{"pmid":"16754863","id":"PMC_16754863","title":"Phylogenetic analysis of the \"ECE\" (CYC/TB1) clade reveals duplications predating the core eudicots.","date":"2006","source":"Proceedings of the National Academy of Sciences of the United States of America","url":"https://pubmed.ncbi.nlm.nih.gov/16754863","citation_count":151,"is_preprint":false},{"pmid":"27889058","id":"PMC_27889058","title":"Mutations in REEP6 Cause Autosomal-Recessive Retinitis Pigmentosa.","date":"2016","source":"American journal of human genetics","url":"https://pubmed.ncbi.nlm.nih.gov/27889058","citation_count":103,"is_preprint":false},{"pmid":"22101417","id":"PMC_22101417","title":"Evolution and diversification of the CYC/TB1 gene family in Asteraceae--a comparative study in Gerbera (Mutisieae) and sunflower (Heliantheae).","date":"2011","source":"Molecular biology and evolution","url":"https://pubmed.ncbi.nlm.nih.gov/22101417","citation_count":99,"is_preprint":false},{"pmid":"28475715","id":"PMC_28475715","title":"REEP6 deficiency leads to retinal degeneration through disruption of ER homeostasis and protein trafficking.","date":"2017","source":"Human molecular genetics","url":"https://pubmed.ncbi.nlm.nih.gov/28475715","citation_count":38,"is_preprint":false},{"pmid":"21799845","id":"PMC_21799845","title":"Cereal domestication and evolution of branching: evidence for soft selection in the Tb1 orthologue of pearl millet (Pennisetum glaucum [L.] R. 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