{"entryType":"UniProtKB reviewed (Swiss-Prot)","primaryAccession":"Q9VBX1","secondaryAccessions":["A8E6Q0"],"uniProtkbId":"NEMF_DROME","entryAudit":{"firstPublicDate":"2011-05-03","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2010-11-30","entryVersion":149,"sequenceVersion":2},"annotationScore":5.0,"organism":{"scientificName":"Drosophila melanogaster","commonName":"Fruit fly","taxonId":7227,"lineage":["Eukaryota","Metazoa","Ecdysozoa","Arthropoda","Altocrustacea","Allotriocarida","Hexapoda","Insecta","Pterygota","Neoptera","Eumetabola","Endopterygota","Aparaglossata","Panorpida","Diptera","Brachycera","Muscomorpha","Ephydroidea","Drosophilidae","Drosophila","Sophophora"]},"proteinExistence":"1: Evidence at protein level","proteinDescription":{"recommendedName":{"fullName":{"evidences":[{"evidenceCode":"ECO:0000305"}],"value":"Ribosome quality control complex subunit NEMF homolog"}},"alternativeNames":[{"fullName":{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"16103875"}],"value":"Protein Caliban"}}]},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"16103875"},{"evidenceCode":"ECO:0000312","source":"FlyBase","id":"FBgn0259152"}],"value":"Clbn"},"orfNames":[{"evidences":[{"evidenceCode":"ECO:0000312","source":"FlyBase","id":"FBgn0259152"}],"value":"CG11847"}]}],"comments":[{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"O60524"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16103875"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22964637"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"30231800"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"31378462"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"33057338"}],"value":"Key component of the ribosome quality control complex (RQC), a ribosome-associated complex that mediates the extraction of incompletely synthesized nascent chains from stalled ribosomes as well as their ubiquitin-mediated proteasomal degradation (By similarity). Thereby, frees 60S subunit ribosomes from the stalled translation complex and prevents the accumulation of nascent polypeptide chains that are potentially toxic for the cell (By similarity). Within the RQC complex, Clbn/NEMF specifically binds stalled 60S ribosomal subunits by recognizing an exposed, nascent chain-conjugated tRNA moiety (PubMed:31378462). Following binding to stalled 60S ribosomal subunits, Clbn/NEMF mediates CAT tailing by recruiting alanine-charged tRNA to the A-site and directing the elongation of stalled nascent chains independently of mRNA or 40S subunits, leading to non-templated C-terminal alanine extensions (CAT tails) (By similarity). On mitochondrial surface, plays a role in mitochondrial-stress induced translational termination impairment and protein carboxyl terminal extension (MISTERMINATE) (PubMed:31378462). Plays a role in regulating nuclear transport possibly through directly binding to both emb and cargo proteins (PubMed:16103875). Plays a role in the regulation of G1-to-S cell cycle transition (PubMed:30231800). Regulates S phase checkpoint by antagonizing E2F1 activity (PubMed:30231800). Together with hid and tefu/ATM, plays a role in DNA damage-induced apoptosis through both p53-dependent and -independent activity (PubMed:22964637). Plays an essential role in the regulation of mitochondrial structure and redox state in enterocytes which is essential for the control of intestinal stem cells proliferation and intestinal homeostasis (PubMed:33057338)"}],"commentType":"FUNCTION"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"O60524"},{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"Q12532"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16103875"}],"value":"Component of the ribosome quality control complex (RQC), composed of at least the E3 ubiquitin ligase l(3)76BDr/LTN1 and Clbn/NEMF associated with the 60S ribosomal subunit. The complex probably also contains TCF25 as well as TER94/VCP and its ubiquitin-binding cofactors (By similarity). Interacts (via its C-terminus) with pros (via its homeobox) (PubMed:16103875). Interacts (via its N-terminus) with emb (PubMed:16103875)"}],"commentType":"SUBUNIT"},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000305"}],"value":"Nucleus","id":"SL-0191"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"33057338"}],"value":"Cytoplasm","id":"SL-0086"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"33057338"}],"value":"Mitochondrion outer membrane","id":"SL-0172"},"topology":{"evidences":[{"evidenceCode":"ECO:0000305"}],"value":"Peripheral membrane protein","id":"SL-9903"},"orientation":{"evidences":[{"evidenceCode":"ECO:0000305"}],"value":"Cytoplasmic side","id":"SL-9910"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"33057338"}],"value":"Expressed in enterocytes (at protein level)"}],"commentType":"TISSUE SPECIFICITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16103875"}],"value":"Expressed ubiquitously throughout embryonic development"}],"commentType":"DEVELOPMENTAL STAGE"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22964637"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"30231800"}],"value":"Up-regulated in response to DNA-damage induced by ionizing radiation"}],"commentType":"INDUCTION"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16103875"}],"value":"The N-terminal domain contains a nuclear export signal. The C-terminal domain may interact with cargo proteins"}],"commentType":"DOMAIN"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16103875"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22964637"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"30231800"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"31378462"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"33057338"}],"value":"Mutant flies are viable and fertile (PubMed:16103875). Shows small, developmental delays and shortened lifespans (PubMed:22964637, PubMed:33057338). Shows dysfunctional intestinal barrier with increased number of intestinal stem cells (ISC) and enteroblasts (EB) (PubMed:33057338). Results in enterocytes (EC) with fragmented mitochondria and defective depolarization (PubMed:33057338). Results in aberrant activation of JAK-STAT signaling pathway (PubMed:33057338). Shows hypersensitization to ionizing irradiation with melanotic masses, defective S phase checkpoint and decreased cell death by apoptosis (PubMed:22964637, PubMed:30231800). In neuroblasts, pros protein does not show relocalization to the nucleus (PubMed:16103875). After ionizing irradiation, simultaneous knockout of Clbn/NEMF and p53 shows similar number of DNA breaks per nucleus but absence of irradiation-induced cell death compared to single knockout (PubMed:22964637). Simultaneous knockout of Clbn/NEMF and hid has similar disregulation of apoptosis to single mutants upon ionizing irradiation (PubMed:22964637). Simultaneous knockout Clbn/NEMF and tefu increases the formation of spontaneous tumors (PubMed:22964637). Simultaneous knockout of Clbn and Pink1 rescues ATP levels, wing posture defects and impaired neuronal numbers observed in Pink1 single mutants (PubMed:31378462). RNAi-mediated knockdown results in reduced lifespan (PubMed:33057338). RNAi-mediated knockdown in EC results in increased number of progenitor cells and EE cells (PubMed:33057338). RNAi-mediated knockdown in ISC and EB, or EE cells, has no significant effect on the number of ISC, EB and EE cells (PubMed:33057338). RNAi-mediated knockdown in the smooth muscle surrounding the gut has no effect on the number of ISC, EB and EE cells (PubMed:33057338). Simultaneous RNAi-mediated knockdown of Clbn and Pink1 rescues muscular and mitochondrial morphology defects present in Pink1 knockdown single mutants (PubMed:31378462)"}],"commentType":"DISRUPTION PHENOTYPE"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16103875"}],"value":"Belongs to the NEMF family"}],"commentType":"SIMILARITY"}],"features":[{"type":"Chain","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":992,"modifier":"EXACT"}},"description":"Ribosome quality control complex subunit NEMF 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