{"gene":"REEP6","run_date":"2026-06-10T06:43:36","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 rod-specific isoform REEP6.1 through an intronic enhancer sequence. Chromatin immunoprecipitation identified NRL binding at this site, and reporter assays in cultured cells and retinal explants confirmed NRL-directed REEP6.1 expression.","method":"Chromatin immunoprecipitation (ChIP), reporter assay, retinal explant transfection, 5'-RACE, exon-specific Taqman assay","journal":"Human molecular genetics","confidence":"High","confidence_rationale":"Tier 2 / Moderate — reciprocal ChIP plus reporter assay in two systems (cultured cells and retinal explants), single lab but multiple orthogonal methods","pmids":["24691551"],"is_preprint":false},{"year":2014,"finding":"Knockdown of Reep6 in mouse and zebrafish resulted in death of retinal cells, establishing a required role for REEP6 in retinal cell survival.","method":"siRNA knockdown in mouse retina; morpholino knockdown in zebrafish","journal":"Human molecular genetics","confidence":"Medium","confidence_rationale":"Tier 2 / Moderate — loss-of-function with defined cellular phenotype (cell death) in two model organisms, single lab","pmids":["24691551"],"is_preprint":false},{"year":2016,"finding":"REEP6 is a member of the REEP/Yop1 family of ER-shaping proteins and is expressed in the retina as a retina-specific isoform (REEP6.1). Missense variants p.Pro128Leu and p.Leu135Pro and a REEP6.1-specific frameshift mutant expressed in cultured cells destabilize the protein. A knock-in mouse (p.Leu135Pro) shows progressive rod photoreceptor degeneration and dysfunction.","method":"Human 3D organoid optic cups; CRISPR-Cas9 knock-in mouse; overexpression in cultured cells; electroretinography; histology","journal":"American journal of human genetics","confidence":"High","confidence_rationale":"Tier 2 / Strong — multiple orthogonal methods (organoids, knock-in mouse, cell expression assays, ERG), replicated across families and models","pmids":["27889058"],"is_preprint":false},{"year":2017,"finding":"In rod photoreceptors, REEP6 localizes to the inner segment and outer plexiform layer. REEP6 knockout mice show expansion of the distal ER and increased mitochondria number in rods, severely reduced expression of retinal guanylate cyclases GC1 and GC2 (near-undetectable), and ER stress markers (elevated CHOP and activated caspase-12), implicating REEP6 in maintaining cGMP homeostasis through facilitating the stability/trafficking of guanylate cyclases and ER/mitochondrial homeostasis. Rhodopsin, Rom1, and peripherin/rds trafficking were unaffected.","method":"CRISPR/Cas9 knockout mouse; transmission electron microscopy; 3View serial block-face scanning EM; immunofluorescence localization; electroretinography; Western blot for GC1, GC2, CHOP, caspase-12","journal":"Human molecular genetics","confidence":"High","confidence_rationale":"Tier 2 / Strong — CRISPR KO mouse with multiple orthogonal structural and biochemical readouts, defined cellular mechanism, single lab but rigorous","pmids":["28475715"],"is_preprint":false},{"year":2017,"finding":"REEP6 is detected in a subset of Clathrin-coated vesicles in rod photoreceptors and interacts with the t-SNARE Syntaxin3. Loss of REEP6 causes aberrant accumulation of vacuole-like structures at the apical inner segment and reduction in selected rod phototransduction proteins.","method":"Co-immunoprecipitation; immunofluorescence colocalization with Clathrin; Reep6 knockout mouse; electroretinography","journal":"Human molecular genetics","confidence":"Medium","confidence_rationale":"Tier 2–3 / Moderate — Co-IP for Syntaxin3 interaction plus KO phenotype, single lab, two orthogonal methods","pmids":["28369466"],"is_preprint":false},{"year":2016,"finding":"REEP6 interacts with CXCR1 (but not CXCR2) and, together with REEP5, facilitates ligand-stimulated endocytosis of CXCR1 and intracellular clustering of β-arrestin2. Depletion of REEP5/REEP6 impairs receptor internalization and reduces IL-8-stimulated ERK phosphorylation and actin polymerization without affecting CXCR1 plasma membrane expression.","method":"Co-immunoprecipitation; siRNA knockdown; overexpression; β-arrestin2 clustering assay; ERK phosphorylation Western blot; in vivo xenograft model","journal":"Scientific reports","confidence":"Medium","confidence_rationale":"Tier 2–3 / Moderate — Co-IP plus functional knockdown with multiple readouts, single lab","pmids":["27966653"],"is_preprint":false},{"year":2021,"finding":"The retina-specific isoform Reep6.1 (containing exon 5-encoded 27 aa) has rod-specific functions that cannot be substituted by the canonical isoform Reep6.2. Exon-5-specific knockout (Reep6E5/E5) produces rod degeneration comparable to full Reep6 knockout, and overexpression of Reep6.2 fails to rescue this phenotype whereas overexpression of Reep6.1 does rescue it.","method":"Isoform-specific knockout mouse (exon 5 deletion); AAV-mediated overexpression rescue experiment; electroretinography; histology","journal":"Human molecular genetics","confidence":"High","confidence_rationale":"Tier 2 / Strong — isoform-specific KO plus rescue experiment with both isoforms, multiple orthogonal readouts, single lab","pmids":["34104971"],"is_preprint":false},{"year":2022,"finding":"REEP6 plays a cell-autonomous role in β-adrenergic signal transduction in adipocytes by facilitating the trafficking of adenylate cyclase 3 (ADCY3) to the plasma membrane. REEP6 KO reduces ADCY3 plasma membrane targeting, severely reduces protein kinase A-mediated signaling in brown adipose tissue, and reduces mitochondrial mass. These defects were reproduced in cultured adipocytes, confirming cell autonomy.","method":"REEP6 knockout mouse; high-fat diet model; siRNA knockdown in C3H10T1/2 adipocytes and primary adipocytes; indirect calorimetry; confocal and electron microscopy; Western blot; mitochondrial DNA analysis; plasma membrane fractionation for ADCY3","journal":"Metabolism: clinical and experimental","confidence":"High","confidence_rationale":"Tier 2 / Strong — KO mouse plus cell-autonomous knockdown, plasma membrane fractionation for ADCY3, multiple orthogonal metabolic readouts, single lab","pmids":["35150731"],"is_preprint":false},{"year":2022,"finding":"miR-672-5p directly regulates REEP6 expression in spinal dorsal horn neurons (validated by dual-luciferase reporter assay targeting the REEP6 mRNA 3'UTR). Upregulation of REEP6 in dorsal horn neurons contributes to neuronal hyperexcitability and mechanical allodynia in bortezomib-induced neuropathic pain.","method":"miRNA microarray; PCR; dual-luciferase reporter assay; intrathecal injection of miR-672-5p agomir/antagomir; sEPSC recording; REEP6 overexpression/knockdown in vivo","journal":"Neurochemical research","confidence":"Medium","confidence_rationale":"Tier 2–3 / Moderate — luciferase reporter plus in vivo functional experiments, single lab","pmids":["36064821"],"is_preprint":false},{"year":2005,"finding":"The REEP6 ortholog Dp1l1 (TB2-like 1) localizes to the cytoplasm in a punctate pattern in retinal ganglion cells, consistent with a role in intracellular membrane trafficking, as determined by immunohistochemistry and intracellular localization analyses.","method":"In situ hybridization; immunohistochemistry; intracellular localization analysis; radiation hybrid mapping","journal":"Investigative ophthalmology & visual science","confidence":"Medium","confidence_rationale":"Tier 3 / Moderate — direct localization experiment in two assays, foundational characterization of the ortholog, single lab","pmids":["15728532"],"is_preprint":false},{"year":2026,"finding":"WTAP-mediated m6A methylation facilitates translation of REEP6 (and PDE6B, RDH12) in rod photoreceptors by depositing m6A modifications in the 3'UTR of Reep6 mRNA. Loss of Wtap abolished WTAP-mediated methyltransferase complex stability, reduced m6A levels genome-wide, and caused epigenetic silencing of Reep6 with consequent rod degeneration; AAV-mediated reintroduction of WTAP partially restored REEP6 expression and mitigated degeneration.","method":"Conditional Wtap knockout mouse; m6A-seq; Western blot; AAV gene therapy rescue; ERG","journal":"Science China. Life sciences","confidence":"Medium","confidence_rationale":"Tier 2 / Moderate — KO mouse with m6A-seq and rescue experiment, single lab, multiple readouts","pmids":["41796262"],"is_preprint":false},{"year":2025,"finding":"In vitro expression of REEP6 alters ER marker expression and Golgi morphology. In an independent Reep6 knockout mouse, guanylate cyclases are not absent but are reduced ~one-third, rhodopsin and GRK1 are also reduced, PDE6 trafficking is normal (contrasting a prior report), and RNA-seq shows reduced phototransduction gene transcription and activated inflammation pathways, suggesting that REEP6 maintains ER and Golgi function required for expression of membrane phototransduction proteins.","method":"CRISPR/Cas9 Reep6 knockout mouse; in vitro REEP6 overexpression with ER/Golgi marker imaging; Western blot; RNA-seq; electroretinography","journal":"bioRxiv","confidence":"Medium","confidence_rationale":"Tier 2–3 / Moderate — independent KO model plus in vitro ER/Golgi morphology assay and RNA-seq, preprint not peer-reviewed","pmids":["bio_10.1101_2025.03.02.641069"],"is_preprint":true}],"current_model":"REEP6 is a transmembrane ER-shaping protein of the REEP/Yop1 family whose rod-specific isoform (REEP6.1), transcriptionally activated by the NRL transcription factor via an intronic enhancer and translationally promoted by WTAP-mediated m6A modification, localizes to the inner segment of rod photoreceptors where it maintains ER and Golgi homeostasis, facilitates the stability/trafficking of guanylate cyclases (GC1/GC2) and adenylate cyclase 3, participates in Clathrin-coated vesicle trafficking through interaction with Syntaxin3, and supports CXCR1-mediated endocytosis in non-retinal cells; loss of REEP6 causes ER stress, reduced expression of membrane phototransduction proteins, and progressive rod photoreceptor degeneration."},"narrative":{"mechanistic_narrative":"REEP6 is a transmembrane member of the REEP/Yop1 family of ER-shaping proteins that maintains endoplasmic reticulum and Golgi homeostasis required for the stability and trafficking of membrane signaling enzymes, and is essential for rod photoreceptor survival [PMID:27889058, PMID:28475715]. In the retina, expression is driven by the rod transcription factor NRL, which binds an enhancer within Reep6 intron 1 to activate the rod-specific isoform REEP6.1 [PMID:24691551]; this isoform carries an exon-5-encoded 27-residue segment whose function cannot be substituted by the canonical REEP6.2 isoform, as exon-5-specific knockout phenocopies full knockout and only REEP6.1 rescues the resulting degeneration [PMID:34104971]. Within rods, REEP6 localizes to the inner segment, and its loss expands the distal ER, increases mitochondrial number, triggers ER stress (CHOP, caspase-12), and reduces the guanylate cyclases GC1/GC2 along with other membrane phototransduction proteins, linking REEP6 to cGMP homeostasis and rod viability [PMID:28475715, PMID:bio_10.1101_2025.03.02.641069]. REEP6 participates in vesicular trafficking, residing in a subset of clathrin-coated vesicles and interacting with the t-SNARE Syntaxin3 [PMID:28369466]. Beyond the retina, REEP6 facilitates plasma-membrane trafficking of signaling effectors more broadly: it promotes adenylate cyclase 3 (ADCY3) delivery to support β-adrenergic/PKA signaling in adipocytes [PMID:35150731] and supports CXCR1-mediated endocytosis and β-arrestin2 clustering [PMID:27966653]. Destabilizing missense and frameshift variants cause progressive rod photoreceptor degeneration in patients and a knock-in mouse model [PMID:27889058].","teleology":[{"year":2005,"claim":"Established the first cellular context for the REEP6 ortholog, placing it in intracellular membrane trafficking before its retinal role was known.","evidence":"In situ hybridization, immunohistochemistry and intracellular localization of the ortholog Dp1l1 in retinal ganglion cells","pmids":["15728532"],"confidence":"Medium","gaps":["No molecular partners or trafficking step defined","Ortholog rather than human REEP6 directly characterized"]},{"year":2014,"claim":"Defined how rod-restricted REEP6 expression is achieved and showed it is required for retinal cell survival, connecting a transcriptional program to a functional necessity.","evidence":"ChIP and reporter assays identifying NRL binding in Reep6 intron 1; siRNA/morpholino knockdown in mouse and zebrafish","pmids":["24691551"],"confidence":"High","gaps":["Knockdown phenotype does not reveal the molecular function of REEP6","Mechanism of cell death not resolved"]},{"year":2016,"claim":"Linked REEP6 to human retinal disease and to the ER-shaping REEP/Yop1 family, showing pathogenic variants destabilize the protein and cause rod degeneration.","evidence":"Patient missense/frameshift variants expressed in cells, human organoid optic cups, and a p.Leu135Pro knock-in mouse with ERG/histology","pmids":["27889058"],"confidence":"High","gaps":["Did not identify the trafficking substrates or partners affected","ER-shaping activity inferred from family membership, not directly assayed"]},{"year":2016,"claim":"Demonstrated a non-retinal trafficking role, implicating REEP6 in receptor endocytosis and downstream signaling.","evidence":"Co-IP, siRNA depletion of REEP5/REEP6, β-arrestin2 clustering, ERK phosphorylation and xenograft assays for CXCR1","pmids":["27966653"],"confidence":"Medium","gaps":["Single-lab Co-IP without reciprocal structural validation","Mechanism by which REEP6 promotes internalization not defined"]},{"year":2017,"claim":"Established a concrete molecular mechanism in rods: REEP6 maintains ER/mitochondrial homeostasis and is required for guanylate cyclase stability and cGMP homeostasis.","evidence":"CRISPR knockout mouse with EM, immunofluorescence, ERG, and Western blots for GC1/GC2, CHOP, caspase-12","pmids":["28475715"],"confidence":"High","gaps":["Whether REEP6 directly chaperones GCs or acts indirectly via ER stress unresolved","Discrepancy with later reports on degree of GC loss"]},{"year":2017,"claim":"Placed REEP6 in clathrin-coated vesicle trafficking and identified a SNARE partner, suggesting a vesicular route for delivery of phototransduction proteins.","evidence":"Co-IP with Syntaxin3, clathrin colocalization, and knockout mouse phenotyping","pmids":["28369466"],"confidence":"Medium","gaps":["Single Co-IP for Syntaxin3 without reciprocal validation","Functional consequence of the Syntaxin3 interaction not directly tested"]},{"year":2021,"claim":"Showed the rod-specific isoform REEP6.1 carries the essential activity, defining isoform specialization as functionally non-redundant.","evidence":"Exon-5-specific knockout mouse and AAV rescue with REEP6.1 versus REEP6.2","pmids":["34104971"],"confidence":"High","gaps":["Molecular property conferred by the exon-5 segment not defined","Why REEP6.2 cannot substitute remains mechanistically unexplained"]},{"year":2022,"claim":"Generalized REEP6's trafficking role to a second effector and tissue, showing it delivers ADCY3 to the plasma membrane for β-adrenergic signaling.","evidence":"Knockout mouse and adipocyte knockdown with plasma membrane fractionation for ADCY3 and metabolic/PKA readouts","pmids":["35150731"],"confidence":"High","gaps":["Direct physical interaction between REEP6 and ADCY3 not shown","Whether the same mechanism operates in rods for GCs not established"]},{"year":2022,"claim":"Identified an upstream post-transcriptional regulator of REEP6 in a neuronal pain context, broadening its regulatory landscape.","evidence":"miR-672-5p luciferase reporter targeting the REEP6 3'UTR plus in vivo agomir/antagomir and electrophysiology","pmids":["36064821"],"confidence":"Medium","gaps":["Mechanistic role of REEP6 in neuronal excitability not defined","Relevance to retinal/adipocyte functions unclear"]},{"year":2026,"claim":"Connected an epitranscriptomic mechanism to REEP6 dosage, showing WTAP-mediated m6A in the 3'UTR promotes REEP6 translation and that its loss causes rod degeneration.","evidence":"Conditional Wtap knockout mouse, m6A-seq, Western blot, and AAV rescue with ERG","pmids":["41796262"],"confidence":"Medium","gaps":["Direct effect on REEP6 separated from co-regulated PDE6B/RDH12 not isolated","Single lab, partial rescue only"]},{"year":2025,"claim":"Independent knockout reframed the ER/Golgi-maintenance model, showing reduced (not absent) guanylate cyclases and broadly reduced phototransduction transcription with inflammation activation.","evidence":"Independent CRISPR knockout mouse, in vitro ER/Golgi marker imaging, RNA-seq and ERG (preprint)","pmids":["bio_10.1101_2025.03.02.641069"],"confidence":"Medium","gaps":["Preprint, not peer-reviewed","Conflicts with prior report on GC absence and PDE6 trafficking","Direct cause of transcriptional downregulation unresolved"]},{"year":null,"claim":"Whether REEP6 directly binds the membrane proteins whose trafficking it supports (GCs, ADCY3, rhodopsin/GRK1) or acts indirectly by shaping ER/Golgi and vesicle compartments remains the central open mechanistic question.","evidence":"","pmids":[],"confidence":"Medium","gaps":["No structure of REEP6 or its complexes","No demonstrated direct cargo interaction","ER-shaping activity not biochemically reconstituted"]}],"mechanism_profile":{"molecular_activity":[{"term_id":"GO:0005198","term_label":"structural molecule activity","supporting_discovery_ids":[2,3]},{"term_id":"GO:0060090","term_label":"molecular adaptor activity","supporting_discovery_ids":[4,7]}],"localization":[{"term_id":"GO:0005783","term_label":"endoplasmic reticulum","supporting_discovery_ids":[2,3,11]},{"term_id":"GO:0005794","term_label":"Golgi apparatus","supporting_discovery_ids":[11]},{"term_id":"GO:0031410","term_label":"cytoplasmic vesicle","supporting_discovery_ids":[4]},{"term_id":"GO:0005829","term_label":"cytosol","supporting_discovery_ids":[9]}],"pathway":[{"term_id":"R-HSA-5653656","term_label":"Vesicle-mediated transport","supporting_discovery_ids":[4,7]},{"term_id":"R-HSA-9709957","term_label":"Sensory Perception","supporting_discovery_ids":[3,6]},{"term_id":"R-HSA-162582","term_label":"Signal Transduction","supporting_discovery_ids":[5,7]}],"complexes":[],"partners":["STX3","CXCR1","REEP5"],"other_free_text":[]}},"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; 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