{"results":[{"entryType":"UniProtKB reviewed (Swiss-Prot)","primaryAccession":"P15336","secondaryAccessions":["A1L3Z2","A4D7U4","A4D7U5","A4D7V1","D3DPE9","G8JLM5","Q13000","Q3B7B7","Q4ZFU9","Q53RY2","Q8TAR1","Q96JT8"],"uniProtkbId":"ATF2_HUMAN","entryAudit":{"firstPublicDate":"1990-04-01","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2008-11-25","entryVersion":245,"sequenceVersion":4},"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":{"value":"Cyclic AMP-dependent transcription factor ATF-2"},"shortNames":[{"value":"cAMP-dependent transcription factor ATF-2"}]},"alternativeNames":[{"fullName":{"value":"Activating transcription factor 2"}},{"fullName":{"value":"Cyclic AMP-responsive element-binding protein 2"},"shortNames":[{"value":"CREB-2"},{"value":"cAMP-responsive element-binding protein 2"}]},{"fullName":{"value":"HB16"}},{"fullName":{"value":"cAMP response element-binding protein CRE-BP1"}}]},"genes":[{"geneName":{"value":"ATF2"},"synonyms":[{"value":"CREB2"},{"value":"CREBP1"}]}],"comments":[{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"P16951"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"11932306"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15105425"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15916964"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18397884"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22304920"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"2529117"},{"evidenceCode":"ECO:0000303","source":"PubMed","id":"18677098"}],"value":"Transcriptional activator which regulates the transcription of various genes, including those involved in anti-apoptosis, cell growth, and DNA damage response (PubMed:11932306, PubMed:15105425, PubMed:15916964, PubMed:22304920, PubMed:2529117). Dependent on its binding partner, binds to CRE (cAMP response element) consensus sequences (5'-TGACGTCA-3') or to AP-1 (activator protein 1) consensus sequences (5'-TGACTCA-3') (PubMed:11932306, PubMed:2529117). In the nucleus, contributes to global transcription and the DNA damage response, in addition to specific transcriptional activities that are related to cell development, proliferation and death (PubMed:22304920). In liver, acts as a regulator of glucose metabolism downstream of OR5V1 signaling, which promotes ATF2 activation, leading to expression of carnitine palmitoyltransferase CPT1A (By similarity). In the cytoplasm, interacts with and perturbs HK1- and VDAC1-containing complexes at the mitochondrial outer membrane, thereby impairing mitochondrial membrane potential, inducing mitochondrial leakage and promoting cell death (PubMed:22304920). The phosphorylated form (mediated by ATM) plays a role in the DNA damage response and is involved in the ionizing radiation (IR)-induced S phase checkpoint control and in the recruitment of the MRN complex into the IR-induced foci (IRIF) (PubMed:15916964). In concert with CUL3 and RBX1, promotes the degradation of KAT5 thereby attenuating its ability to acetylate and activate ATM (PubMed:18397884). Can elicit oncogenic or tumor suppressor activities depending on the tissue or cell type (PubMed:18677098)"}],"commentType":"FUNCTION"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"P16951"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"11932306"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15916964"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18397884"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22275354"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22304920"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"2529117"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"9748258"}],"value":"Binds DNA as a dimer and can form a homodimer in the absence of DNA (PubMed:11932306, PubMed:2529117). Can form a heterodimer with JUN (PubMed:22275354). Heterodimerization is essential for its transcriptional activity (PubMed:22275354). Binds through its N-terminal region to UTF1 which acts as a coactivator of ATF2 transcriptional activity (PubMed:9748258). Interacts with the HK1/VDAC1 complex (PubMed:22304920). Interacts with NBN, MRE11, XPO1, KAT5 and CUL3 (PubMed:15916964, PubMed:18397884). Binds through its N-terminal region to UTF1 which acts as a coactivator of ATF2 transcriptional activity (By similarity)"}],"commentType":"SUBUNIT"},{"commentType":"INTERACTION","interactions":[{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"P15336","geneName":"ATF2","intActId":"EBI-1170906"},"numberOfExperiments":2,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"P18847","geneName":"ATF3","intActId":"EBI-712767"},"numberOfExperiments":5,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"P18848","geneName":"ATF4","intActId":"EBI-492498"},"numberOfExperiments":2,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"P17544","geneName":"ATF7","intActId":"EBI-765623"},"numberOfExperiments":5,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"O14867","geneName":"BACH1","intActId":"EBI-1263541"},"numberOfExperiments":4,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"Q8N9N5-2","geneName":"BANP","intActId":"EBI-11524452"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"Q16520","geneName":"BATF","intActId":"EBI-749503"},"numberOfExperiments":2,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"Q9NR55","geneName":"BATF3","intActId":"EBI-10312707"},"numberOfExperiments":5,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"P53567","geneName":"CEBPG","intActId":"EBI-740209"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"Q7L2Z9","geneName":"CENPQ","intActId":"EBI-2350265"},"numberOfExperiments":5,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"Q99966","geneName":"CITED1","intActId":"EBI-2624951"},"numberOfExperiments":2,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"P35638","geneName":"DDIT3","intActId":"EBI-742651"},"numberOfExperiments":2,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"Q96B26","geneName":"EXOSC8","intActId":"EBI-371922"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"P01100","geneName":"FOS","intActId":"EBI-852851"},"numberOfExperiments":15,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"Q6FG41","geneName":"FOS","intActId":"EBI-10198738"},"numberOfExperiments":3,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"P53539","geneName":"FOSB","intActId":"EBI-2806743"},"numberOfExperiments":5,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"P15407","geneName":"FOSL1","intActId":"EBI-744510"},"numberOfExperiments":4,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"P15408","geneName":"FOSL2","intActId":"EBI-3893419"},"numberOfExperiments":8,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"P56524","geneName":"HDAC4","intActId":"EBI-308629"},"numberOfExperiments":2,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"P05412","geneName":"JUN","intActId":"EBI-852823"},"numberOfExperiments":23,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"P17275","geneName":"JUNB","intActId":"EBI-748062"},"numberOfExperiments":7,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"P17535","geneName":"JUND","intActId":"EBI-2682803"},"numberOfExperiments":7,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"P15336","intActId":"EBI-1170906"},"interactantTwo":{"uniProtKBAccession":"Q9BVG8","geneName":"KIFC3","intActId":"EBI-2125614"},"numberOfExpe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LOCATION","note":{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15916964"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22275354"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22304920"}],"value":"Shuttles between the cytoplasm and the nucleus and heterodimerization with JUN is essential for the nuclear localization (PubMed:22275354). Localization to the cytoplasm is observed under conditions of cellular stress and in disease states (PubMed:22304920). Localizes at the mitochondrial outer membrane in response to genotoxic stress (PubMed:22304920). Phosphorylation at Thr-52 is required for its nuclear localization and negatively regulates its mitochondrial localization (PubMed:22304920). Co-localizes with the MRN complex in the IR-induced foci (IRIF) (PubMed:15916964)"}]},"subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15105425"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15916964"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18397884"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19176525"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21098032"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22275354"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22304920"}],"value":"Nucleus","id":"SL-0191"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22275354"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22304920"}],"value":"Cytoplasm","id":"SL-0086"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22304920"}],"value":"Mitochondrion outer membrane","id":"SL-0172"}}]},{"commentType":"ALTERNATIVE PRODUCTS","events":["Alternative splicing"],"isoforms":[{"name":{"value":"1"},"isoformIds":["P15336-1"],"isoformSequenceStatus":"Displayed"},{"name":{"value":"2"},"isoformIds":["P15336-2"],"sequenceIds":["VSP_000587","VSP_000588"],"isoformSequenceStatus":"Described"},{"name":{"value":"3"},"isoformIds":["P15336-3"],"sequenceIds":["VSP_045161"],"isoformSequenceStatus":"Described"},{"name":{"value":"4"},"isoformIds":["P15336-4"],"sequenceIds":["VSP_046959"],"isoformSequenceStatus":"Described"},{"name":{"value":"5"},"isoformIds":["P15336-5"],"sequenceIds":["VSP_046960"],"isoformSequenceStatus":"Described"},{"name":{"value":"6"},"isoformIds":["P15336-6"],"sequenceIds":["VSP_046960","VSP_047593"],"isoformSequenceStatus":"Described"},{"name":{"value":"7"},"isoformIds":["P15336-7"],"sequenceIds":["VSP_046960","VSP_047594","VSP_047595"],"isoformSequenceStatus":"Described"},{"name":{"value":"8"},"synonyms":[{"value":"ATF2-small"}],"isoformIds":["P15336-8"],"sequenceIds":["VSP_047593"],"isoformSequenceStatus":"Described"}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"2529117"}],"value":"Ubiquitously expressed, with more abundant expression in the brain"}],"commentType":"TISSUE SPECIFICITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22275354"}],"value":"The nuclear export signal 1 (N-NES) negatively regulates its nuclear localization and transcriptional activity"}],"commentType":"DOMAIN"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"P16951"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10821277"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"12110590"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15105425"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"15916964"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"21098032"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8855261"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"9430721"}],"value":"Phosphorylation of Thr-69 by MAPK14 and MAPK11, and at Thr-71 by MAPK1/ERK2, MAPK3/ERK1, MAPK11, MAPK12 and MAPK14 in response to external stimulus like insulin causes increased transcriptional activity (PubMed:12110590, PubMed:9430721). Phosphorylated at Thr-69 and Thr-71 by PKA, promoting transcriptional activity (By similarity). Phosphorylated by PLK3 following hyperosmotic stress (PubMed:21098032). Also phosphorylated and activated by JNK and CaMK4 (PubMed:8855261). ATM-mediated phosphorylation at Ser-490 and Ser-498 stimulates its function in DNA damage response (PubMed:15916964). Phosphorylation at Ser-62, Thr-73 and Ser-121 activates its transcriptional activity (PubMed:15105425). Phosphorylation at Thr-69 or Thr-71 enhances acetylation of histones H2B and H4 (PubMed:10821277)"}],"commentType":"PTM"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000305"}],"value":"Belongs to the bZIP family. ATF subfamily"}],"commentType":"SIMILARITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"10821277"},{"evidenceCode":"ECO:0000305"}],"value":"Was reported to have histone acetyltransferase (HAT) activity, specifically towards histones H2B and H4 in vitro (PubMed:10821277). 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