{"entryType":"UniProtKB reviewed (Swiss-Prot)","primaryAccession":"Q96JB5","secondaryAccessions":["B7Z6N4","D3DTU1","D3DTU2","F5H3I5","Q53FA2","Q9H3F8","Q9H8G0","Q9HBR9"],"uniProtkbId":"CK5P3_HUMAN","entryAudit":{"firstPublicDate":"2003-06-16","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2003-06-16","entryVersion":198,"sequenceVersion":2},"annotationScore":5.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"lineage":["Eukaryota","Metazoa","Chordata","Craniata","Vertebrata","Euteleostomi","Mammalia","Eutheria","Euarchontoglires","Primates","Haplorrhini","Catarrhini","Hominidae","Homo"]},"proteinExistence":"1: Evidence at protein level","proteinDescription":{"recommendedName":{"fullName":{"evidences":[{"evidenceCode":"ECO:0000305"}],"value":"CDK5 regulatory subunit-associated protein 3"}},"alternativeNames":[{"fullName":{"evidences":[{"evidenceCode":"ECO:0000305","source":"PubMed","id":"10721722"}],"value":"CDK5 activator-binding protein C53"}},{"fullName":{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"20164180"}],"value":"LXXLL/leucine-zipper-containing ARF-binding protein"}},{"fullName":{"evidences":[{"evidenceCode":"ECO:0000312","source":"EMBL","id":"AAK69655.1"}],"value":"Protein HSF-27"}}]},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"30635284"},{"evidenceCode":"ECO:0000312","source":"HGNC","id":"HGNC:18673"}],"value":"CDK5RAP3"},"synonyms":[{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"20164180"}],"value":"IC53"},{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"20164180"}],"value":"LZAP"}],"orfNames":[{"value":"MSTP016"},{"value":"OK/SW-cl.114"},{"value":"PP1553"}]}],"comments":[{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"Q99LM2"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15790566"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16173922"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17785205"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19223857"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"20228063"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21283629"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23152784"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23478299"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"30635284"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"32851973"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"36121123"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"36543799"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"37595036"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"38383785"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"38383789"}],"value":"Substrate adapter of E3 ligase complexes mediating ufmylation, the covalent attachment of the ubiquitin-like modifier UFM1 to substrate proteins, and which is involved in various processes, such as ribosome recycling and reticulophagy (also called ER-phagy) (PubMed:23152784, PubMed:30635284, PubMed:32851973, PubMed:36121123, PubMed:36543799, PubMed:37595036, PubMed:38383785, PubMed:38383789). As part of the UREL complex, plays a key role in ribosome recycling by promoting mono-ufmylation of RPL26/uL24 subunit of the 60S ribosome (PubMed:38383785, PubMed:38383789). Ufmylation of RPL26/uL24 occurs on free 60S ribosomes following ribosome dissociation: it weakens the junction between post-termination 60S subunits and SEC61 translocons, promoting release and recycling of the large ribosomal subunit from the endoplasmic reticulum membrane (PubMed:38383785, PubMed:38383789). Ufmylation of RPL26/uL24 and subsequent 60S ribosome recycling either take place after normal termination of translation or after ribosome stalling during cotranslational translocation at the endoplasmic reticulum (PubMed:32851973, PubMed:37595036, PubMed:38383785, PubMed:38383789). Within the UREL complex, CDK5RAP3 acts as a substrate adapter that constrains UFL1 ligase activity to mono-ufmylate RPL26/uL24 at 'Lys-134' (PubMed:36121123, PubMed:38383785, PubMed:38383789). The UREL complex is also involved in reticulophagy in response to endoplasmic reticulum stress by promoting ufmylation of proteins such as CYB5R3, thereby promoting lysosomal degradation of ufmylated proteins (PubMed:36543799). Also acts as a regulator of transcription: negatively regulates NF-kappa-B-mediated gene transcription through the control of RELA phosphorylation (PubMed:17785205, PubMed:20228063). Also regulates mitotic G2/M transition checkpoint and mitotic G2 DNA damage checkpoint (PubMed:15790566, PubMed:19223857). Through its interaction with CDKN2A/ARF and MDM2 may induce MDM2-dependent p53/TP53 ubiquitination, stabilization and activation in the nucleus, thereby promoting G1 cell cycle arrest and inhibition of cell proliferation (PubMed:16173922). May also play a role in the rupture of the nuclear envelope during apoptosis (PubMed:23478299). May regulate MAPK14 activity by regulating its dephosphorylation by PPM1D/WIP1 (PubMed:21283629). Required for liver development (By similarity)"}],"commentType":"FUNCTION"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21971960"}],"value":"(Microbial infection) May be negatively regulated by hepatitis B virus large envelope protein mutant pre-s2 to promote mitotic entry"}],"commentType":"FUNCTION"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"Q9JLH7"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12737517"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15790566"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16173922"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"17785205"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19223857"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"20164180"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"20228063"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21283629"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21465471"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"32851973"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"36121123"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"36543799"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"36762703"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"37595036"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"38383785"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"38383789"}],"value":"Substrate adapter component of the UFM1 ribosome E3 ligase (UREL) complex, composed of UFL1, DDRGK1 and CDK5RAP3 (PubMed:20164180, PubMed:20228063, PubMed:32851973, PubMed:36121123, PubMed:36543799, PubMed:37595036, PubMed:38383785, PubMed:38383789). Interaction with UFL1 anchors CDK5RAP3 in the cytoplasm, preventing its translocation to the nucleus which allows expression of the CCND1 cyclin and progression of cells through the G1/S transition (By similarity). Interacts with ATG8 family proteins MAP1LC3A, MAP1LC3B, GABARAP, GABARAPL1 and GABARAPL2 (PubMed:32851973, PubMed:36543799, PubMed:36762703). Interacts with CDK5R1; competes with CDK5RAP1 and CDK5RAP2 (PubMed:12737517). Interacts with RELA (PubMed:17785205). Interacts with CHEK1; may negatively regulate CHEK1 and thereby stimulate entry into mitosis (PubMed:19223857). Interacts with CDKN2A/ARF and MDM2; forms a ternary complex involved in regulation of p53/TP53 (PubMed:16173922). Interacts with MAPK14 (PubMed:21283629). Interacts with CCNB1 (PubMed:15790566). Interacts with TUBG1; may regulate CDK5RAP3 in mitotic G2/M transition checkpoint (PubMed:21465471)"}],"commentType":"SUBUNIT"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21971960"}],"value":"(Microbial infection) Interacts with hepatitis B virus large envelope protein mutant pre-s2; promotes mitotic entry"}],"commentType":"SUBUNIT"},{"commentType":"INTERACTION","interactions":[{"interactantOne":{"uniProtKBAccession":"Q96JB5","intActId":"EBI-718818"},"interactantTwo":{"uniProtKBAccession":"O75344","geneName":"FKBP6","intActId":"EBI-744771"},"numberOfExperiments":12,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"Q96JB5","intActId":"EBI-718818"},"interactantTwo":{"uniProtKBAccession":"Q04206","geneName":"RELA","intActId":"EBI-73886"},"numberOfExperiments":4,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"Q96JB5","intActId":"EBI-718818"},"interactantTwo":{"uniProtKBAccession":"Q9Y5M8","geneName":"SRPRB","intActId":"EBI-355393"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"Q96JB5","intActId":"EBI-718818"},"interactantTwo":{"uniProtKBAccession":"Q9UJ04","geneName":"TSPYL4","intActId":"EBI-308511"},"numberOfExperiments":6,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"Q96JB5","intActId":"EBI-718818"},"interactantTwo":{"uniProtKBAccession":"Q9Y3C8","geneName":"UFC1","intActId":"EBI-1045733"},"numberOfExperiments":7,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"Q96JB5","intActId":"EBI-718818"},"interactantTwo":{"uniProtKBAccession":"O94874","geneName":"UFL1","intActId":"EBI-1048088"},"numberOfExperiments":12,"organismDiffer":false}]},{"commentType":"SUBCELLULAR LOCATION","note":{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23478299"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"38383785"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"38383789"}],"value":"Tethered to the endoplasmic reticulum membrane as part of the UFM1 ribosome E3 ligase (UREL) complex (PubMed:38383785, PubMed:38383789). Colocalizes and associates with microtubules (PubMed:23478299)"}]},"subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"36543799"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"38383785"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"38383789"}],"value":"Endoplasmic reticulum membrane","id":"SL-0097"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15790566"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16173922"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19223857"}],"value":"Cytoplasm","id":"SL-0086"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16173922"}],"value":"Nucleus","id":"SL-0191"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19223857"}],"value":"Cytoplasm, cytoskeleton, microtubule organizing center, centrosome","id":"SL-0048"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23478299"}],"value":"Cytoplasm, cytoskeleton","id":"SL-0090"}}]},{"commentType":"ALTERNATIVE PRODUCTS","events":["Alternative splicing"],"isoforms":[{"name":{"value":"1"},"isoformIds":["Q96JB5-1"],"isoformSequenceStatus":"Displayed"},{"name":{"value":"2"},"synonyms":[{"value":"IC53"}],"isoformIds":["Q96JB5-2"],"sequenceIds":["VSP_007566","VSP_007567"],"isoformSequenceStatus":"Described"},{"name":{"value":"3"},"synonyms":[{"value":"IC53-2"}],"isoformIds":["Q96JB5-3"],"sequenceIds":["VSP_007568"],"isoformSequenceStatus":"Described"},{"name":{"value":"4"},"isoformIds":["Q96JB5-4"],"sequenceIds":["VSP_055646"],"isoformSequenceStatus":"Described"}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10721722"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12054757"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12737517"}],"value":"Ubiquitously expressed (PubMed:10721722, PubMed:12054757). Expressed in heart, brain, placenta, lung, liver, skeletal muscle, kidney and pancreas. Isoform 3 is expressed in kidney, liver, skeletal muscle and placenta (PubMed:12737517)"}],"commentType":"TISSUE SPECIFICITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"32851973"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"36762703"}],"value":"The shuffled ATG8-binding motifs mediate interaction with both ATG8 family protein and UFM1"}],"commentType":"DOMAIN"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"38383789"}],"value":"The RPL10a-binding domain (RBD) anchors the UREL complex onto the ribosome via association with RPL10a/ul1"}],"commentType":"DOMAIN"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"Q9JLH7"}],"value":"May be phosphorylated by CDK5"}],"commentType":"PTM"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"20228063"}],"value":"Ubiquitinated. Probably triggers proteasomal degradation and is negatively regulated by UFL1"}],"commentType":"PTM"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"Q99LM2"}],"value":"May be ufmylated"}],"commentType":"PTM"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"23478299"}],"value":"Cleaved by caspases early during apoptosis, the resulting peptides may play a role in rupture of the nuclear envelope"}],"commentType":"PTM"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000305"}],"value":"Due to an intron retention"}],"commentType":"MISCELLANEOUS","molecule":"Isoform 2"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000305"}],"value":"Due to intron retention"}],"commentType":"MISCELLANEOUS","molecule":"Isoform 3"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000305"}],"value":"Belongs to the CDK5RAP3 family"}],"commentType":"SIMILARITY"},{"commentType":"SEQUENCE CAUTION","sequenceCautionType":"Frameshift","sequence":"AAK69655.1","evidences":[{"evidenceCode":"ECO:0000305"}]}],"features":[{"type":"Chain","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":506,"modifier":"EXACT"}},"description":"CDK5 regulatory subunit-associated protein 3","featureId":"PRO_0000220516"},{"type":"Region","location":{"start":{"value":269,"modifier":"EXACT"},"end":{"value":506,"modifier":"EXACT"}},"description":"Required for interaction with UFL1 and mediates interaction with CHEK1","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19223857"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"20228063"}]},{"type":"Region","location":{"start":{"value":355,"modifier":"EXACT"},"end":{"value":370,"modifier":"EXACT"}},"description":"RPL10a-binding domain (RBD)","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"38383789"}]},{"type":"Motif","location":{"start":{"value":267,"modifier":"EXACT"},"end":{"value":270,"modifier":"EXACT"}},"description":"Shuffled ATG8-binding motif 1","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"32851973"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"36762703"}]},{"type":"Motif","location":{"start":{"value":292,"modifier":"EXACT"},"end":{"value":295,"modifier":"EXACT"}},"description":"Shuffled ATG8-binding motif 2","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"32851973"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"36762703"}]},{"type":"Motif","location":{"start":{"value":310,"modifier":"EXACT"},"end":{"value":313,"modifier":"EXACT"}},"description":"Shuffled ATG8-binding motif 3","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"32851973"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"36762703"}]},{"type":"Cross-link","location":{"start":{"value":450,"modifier":"EXACT"},"end":{"value":450,"modifier":"EXACT"}},"description":"Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO2)","evidences":[{"evidenceCode":"ECO:0007744","source":"PubMed","id":"25772364"}]},{"type":"Alternative sequence","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":225,"modifier":"EXACT"}},"description":"in isoform 3","evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"12737517"}],"featureId":"VSP_007568","alternativeSequence":{}},{"type":"Alternative sequence","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":87,"modifier":"EXACT"}},"description":"in isoform 2","evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"12054757"}],"featureId":"VSP_007566","alternativeSequence":{}},{"type":"Alternative sequence","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":2,"modifier":"EXACT"}},"description":"in isoform 4","evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"14702039"}],"featureId":"VSP_055646","alternativeSequence":{"originalSequence":"ME","alternativeSequences":["MRRQSMTSATRDLHTALDGKATQGGKK"]}},{"type":"Alternative sequence","location":{"start":{"value":88,"modifier":"EXACT"},"end":{"value":111,"modifier":"EXACT"}},"description":"in isoform 2","evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"12054757"}],"featureId":"VSP_007567","alternativeSequence":{"originalSequence":"RYSSQRMKDWQEIIALYEKDNTYL","alternativeSequences":["MCVHPGACLPHVGVSWAEFPGHFS"]}},{"type":"Natural variant","location":{"start":{"value":324,"modifier":"EXACT"},"end":{"value":324,"modifier":"EXACT"}},"description":"in dbSNP:rs35054799","featureCrossReferences":[{"database":"dbSNP","id":"rs35054799"}],"featureId":"VAR_048688","alternativeSequence":{"originalSequence":"L","alternativeSequences":["V"]}},{"type":"Mutagenesis","location":{"start":{"value":217,"modifier":"EXACT"},"end":{"value":217,"modifier":"EXACT"}},"description":"Abolished interaction with UFL1; 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