{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T01:35:32Z","timestamp":1791423332702,"version":"4.3.3"},"reference-count":81,"publisher":"Springer Science and Business Media LLC","issue":"3","license":[{"start":{"date-parts":[[2009,12,10]],"date-time":"2009-12-10T00:00:00Z","timestamp":1260403200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"},{"start":{"date-parts":[[2009,12,10]],"date-time":"2009-12-10T00:00:00Z","timestamp":1260403200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["EMBO J"],"published-print":{"date-parts":[[2010,2,3]]},"DOI":"10.1038\/emboj.2009.372","type":"journal-article","created":{"date-parts":[[2009,12,10]],"date-time":"2009-12-10T09:07:40Z","timestamp":1260436060000},"page":"574-585","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":118,"title":["53BP1 promotes ATM activity through direct interactions with the MRN complex"],"prefix":"10.1038","volume":"29","author":[{"given":"Ji\u2010Hoon","family":"Lee","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Aaron A","family":"Goodarzi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Penny A","family":"Jeggo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tanya T","family":"Paull","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2009,12,10]]},"reference":[{"key":"BFEMBJ2009372_CR1","doi-asserted-by":"crossref","first-page":"1854","DOI":"10.4161\/cc.4.12.2282","volume":"4","author":"MM Adams","year":"2005","unstructured":"Adams MM, Wang B, Xia Z, Morales JC, Lu X, Donehower LA, Bochar DA, Elledge SJ, Carpenter PB (2005) 53BP1 oligomerization is independent of its methylation by PRMT1. Cell Cycle 4: 1854\u20131861","journal-title":"Cell Cycle"},{"key":"BFEMBJ2009372_CR2","doi-asserted-by":"crossref","first-page":"1719","DOI":"10.1128\/MCB.21.5.1719-1729.2001","volume":"21","author":"L Anderson","year":"2001","unstructured":"Anderson L, Henderson C, Adachi Y (2001) Phosphorylation and rapid relocalization of 53BP1 to nuclear foci upon DNA damage. Mol Cell Biol 21: 1719\u20131729","journal-title":"Mol Cell Biol"},{"key":"BFEMBJ2009372_CR3","doi-asserted-by":"crossref","first-page":"647","DOI":"10.1016\/j.molcel.2007.01.028","volume":"25","author":"V Bhaskara","year":"2007","unstructured":"Bhaskara V, Dupre A, Lengsfeld B, Hopkins BB, Chan A, Lee JH, Zhang X, Gautier J, Zakian VA, Paull TT (2007) Rad50 adenylate kinase activity regulates DNA tethering by Mre11\/Rad50 complexes. Mol Cell 25: 647\u2013661","journal-title":"Mol Cell"},{"key":"BFEMBJ2009372_CR4","doi-asserted-by":"crossref","first-page":"1834","DOI":"10.4161\/cc.4.12.2250","volume":"4","author":"FM Boisvert","year":"2005","unstructured":"Boisvert FM, Rhie A, Richard S, Doherty AJ (2005) The GAR motif of 53BP1 is arginine methylated by PRMT1 and is necessary for 53BP1 DNA binding activity. Cell Cycle 4: 1834\u20131841","journal-title":"Cell Cycle"},{"key":"BFEMBJ2009372_CR5","doi-asserted-by":"crossref","first-page":"68","DOI":"10.1096\/fasebj.11.1.9034168","volume":"11","author":"P Bork","year":"1997","unstructured":"Bork P, Hofmann K, Bucher P, Neuwald AF, Altschul SF, Koonin EV (1997) A superfamily of conserved domains in DNA damage\u2010responsive cell cycle checkpoint proteins. FASEB J 11: 68\u201376","journal-title":"FASEB J"},{"key":"BFEMBJ2009372_CR6","doi-asserted-by":"crossref","first-page":"1361","DOI":"10.1016\/j.cell.2006.10.043","volume":"127","author":"MV Botuyan","year":"2006","unstructured":"Botuyan MV, Lee J, Ward IM, Kim JE, Thompson JR, Chen J, Mer G (2006) Structural basis for the methylation state\u2010specific recognition of histone H4\u2010K20 by 53BP1 and Crb2 in DNA repair. Cell 127: 1361\u20131373","journal-title":"Cell"},{"key":"BFEMBJ2009372_CR7","doi-asserted-by":"crossref","first-page":"466","DOI":"10.1038\/nature04866","volume":"442","author":"AL Bredemeyer","year":"2006","unstructured":"Bredemeyer AL, Sharma GG, Huang CY, Helmink BA, Walker LM, Khor KC, Nuskey B, Sullivan KE, Pandita TK, Bassing CH, Sleckman BP (2006) ATM stabilizes DNA double\u2010strand\u2010break complexes during V(D)J recombination. Nature 442: 466\u2013470","journal-title":"Nature"},{"key":"BFEMBJ2009372_CR8","doi-asserted-by":"crossref","first-page":"5214","DOI":"10.1128\/MCB.21.15.5214-5222.2001","volume":"21","author":"G Buscemi","year":"2001","unstructured":"Buscemi G, Savio C, Zannini L, Micciche F, Masnada D, Nakanishi M, Tauchi H, Komatsu K, Mizutani S, Khanna K, Chen P, Concannon P, Chessa L, Delia D (2001) Chk2 activation dependence on Nbs1 after DNA damage. Mol Cell Biol 21: 5214\u20135222","journal-title":"Mol Cell Biol"},{"key":"BFEMBJ2009372_CR9","doi-asserted-by":"crossref","first-page":"477","DOI":"10.1016\/S0092-8674(00)81175-7","volume":"93","author":"JP Carney","year":"1998","unstructured":"Carney JP, Maser RS, Olivares H, Davis EM, Le Beau M, Yates III JR, Hays L, Morgan WF, Petrini JH (1998) The hMre11\/hRad50 protein complex and Nijmegen breakage syndrome: linkage of double\u2010strand break repair to the cellular DNA damage response. Cell 93: 477\u2013486","journal-title":"Cell"},{"key":"BFEMBJ2009372_CR10","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1038\/nrm805","volume":"3","author":"D D'Amours","year":"2002","unstructured":"D'Amours D, Jackson SP (2002) The Mre11 complex: at the crossroads of dna repair and checkpoint signalling. Nat Rev Mol Cell Biol 3: 317\u2013327","journal-title":"Nat Rev Mol Cell Biol"},{"key":"BFEMBJ2009372_CR11","doi-asserted-by":"crossref","first-page":"749","DOI":"10.1083\/jcb.200612047","volume":"176","author":"D Deckbar","year":"2007","unstructured":"Deckbar D, Birraux J, Krempler A, Tchouandong L, Beucher A, Walker S, Stiff T, Jeggo P, Lobrich M (2007) Chromosome breakage after G2 checkpoint release. J Cell Biol 176: 749\u2013755","journal-title":"J Cell Biol"},{"key":"BFEMBJ2009372_CR12","doi-asserted-by":"crossref","first-page":"914","DOI":"10.1038\/nature08196","volume":"460","author":"Y Deng","year":"2009","unstructured":"Deng Y, Guo X, Ferguson DO, Chang S (2009) Multiple roles for MRE11 at uncapped telomeres. Nature 460: 914\u2013918","journal-title":"Nature"},{"key":"BFEMBJ2009372_CR13","doi-asserted-by":"crossref","first-page":"3863","DOI":"10.1093\/emboj\/cdf383","volume":"21","author":"DJ Derbyshire","year":"2002","unstructured":"Derbyshire DJ, Basu BP, Serpell LC, Joo WS, Date T, Iwabuchi K, Doherty AJ (2002) Crystal structure of human 53BP1 BRCT domains bound to p53 tumour suppressor. EMBO J 21: 3863\u20133872","journal-title":"EMBO J"},{"key":"BFEMBJ2009372_CR14","doi-asserted-by":"crossref","first-page":"1003","DOI":"10.1084\/jem.20070319","volume":"204","author":"S Difilippantonio","year":"2007","unstructured":"Difilippantonio S, Celeste A, Kruhlak MJ, Lee Y, Difilippantonio MJ, Feigenbaum L, Jackson SP, McKinnon PJ, Nussenzweig A (2007) Distinct domains in Nbs1 regulate irradiation\u2010induced checkpoints and apoptosis. J Exp Med 204: 1003\u20131011","journal-title":"J Exp Med"},{"key":"BFEMBJ2009372_CR15","doi-asserted-by":"crossref","first-page":"529","DOI":"10.1038\/nature07476","volume":"456","author":"S Difilippantonio","year":"2008","unstructured":"Difilippantonio S, Gapud E, Wong N, Huang CY, Mahowald G, Chen HT, Kruhlak MJ, Callen E, Livak F, Nussenzweig MC, Sleckman BP, Nussenzweig A (2008) 53BP1 facilitates long\u2010range DNA end\u2010joining during V(D)J recombination. Nature 456: 529\u2013533","journal-title":"Nature"},{"key":"BFEMBJ2009372_CR16","doi-asserted-by":"crossref","first-page":"524","DOI":"10.1038\/nature07433","volume":"456","author":"N Dimitrova","year":"2008","unstructured":"Dimitrova N, Chen YC, Spector DL, de Lange T (2008) 53BP1 promotes non\u2010homologous end joining of telomeres by increasing chromatin mobility. Nature 456: 524\u2013528","journal-title":"Nature"},{"key":"BFEMBJ2009372_CR17","doi-asserted-by":"crossref","first-page":"808","DOI":"10.1038\/nsmb.1639","volume":"16","author":"M Dinkelmann","year":"2009","unstructured":"Dinkelmann M, Spehalski E, Stoneham T, Buis J, Wu Y, Sekiguchi JM, Ferguson DO (2009) Multiple functions of MRN in end\u2010joining pathways during isotype class switching. Nat Struct Mol Biol 16: 808\u2013813","journal-title":"Nat Struct Mol Biol"},{"key":"BFEMBJ2009372_CR18","doi-asserted-by":"crossref","first-page":"998","DOI":"10.1038\/ncb892","volume":"4","author":"Jr RA DiTullio","year":"2002","unstructured":"DiTullio Jr RA, Mochan TA, Venere M, Bartkova J, Sehested M, Bartek J, Halazonetis TD (2002) 53BP1 functions in an ATM\u2010dependent checkpoint pathway that is constitutively activated in human cancer. Nat Cell Biol 4: 998\u20131002","journal-title":"Nat Cell Biol"},{"key":"BFEMBJ2009372_CR19","doi-asserted-by":"crossref","first-page":"31251","DOI":"10.1074\/jbc.M405372200","volume":"279","author":"M Fabbro","year":"2004","unstructured":"Fabbro M, Savage K, Hobson K, Deans AJ, Powell SN, McArthur GA, Khanna KK (2004) BRCA1\u2010BARD1 complexes are required for p53Ser\u201015 phosphorylation and a G1\/S arrest following ionizing radiation\u2010induced DNA damage. J Biol Chem 279: 31251\u201331258","journal-title":"J Biol Chem"},{"key":"BFEMBJ2009372_CR20","doi-asserted-by":"crossref","first-page":"605","DOI":"10.1038\/nature03442","volume":"434","author":"J Falck","year":"2005","unstructured":"Falck J, Coates J, Jackson SP (2005) Conserved modes of recruitment of ATM, ATR and DNA\u2010PKcs to sites of DNA damage. Nature 434: 605\u2013611","journal-title":"Nature"},{"key":"BFEMBJ2009372_CR21","doi-asserted-by":"crossref","first-page":"2860","DOI":"10.1093\/emboj\/cdg274","volume":"22","author":"N Foray","year":"2003","unstructured":"Foray N, Marot D, Gabriel A, Randrianarison V, Carr AM, Perricaudet M, Ashworth A, Jeggo P (2003) A subset of ATM\u2010 and ATR\u2010dependent phosphorylation events requires the BRCA1 protein. EMBO J 22: 2860\u20132871","journal-title":"EMBO J"},{"key":"BFEMBJ2009372_CR22","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1038\/75508","volume":"25","author":"M Gatei","year":"2000","unstructured":"Gatei M, Young D, Cerosaletti KM, Desai\u2010Mehta A, Spring K, Kozlov S, Lavin MF, Gatti RA, Concannon P, Khanna K (2000) ATM\u2010dependent phosphorylation of nibrin in response to radiation exposure. Nat Genet 25: 115\u2013119","journal-title":"Nat Genet"},{"key":"BFEMBJ2009372_CR23","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1016\/j.molcel.2008.05.017","volume":"31","author":"AA Goodarzi","year":"2008","unstructured":"Goodarzi AA, Noon AT, Deckbar D, Ziv Y, Shiloh Y, Lobrich M, Jeggo PA (2008) ATM signaling facilitates repair of DNA double\u2010strand breaks associated with heterochromatin. Mol Cell 31: 167\u2013177","journal-title":"Mol Cell"},{"key":"BFEMBJ2009372_CR24","doi-asserted-by":"crossref","first-page":"569","DOI":"10.1042\/BST0370569","volume":"37","author":"AA Goodarzi","year":"2009","unstructured":"Goodarzi AA, Noon AT, Jeggo PA (2009) The impact of heterochromatin on DSB repair. Biochem Soc Trans 37(Pt 3): 569\u2013576","journal-title":"Biochem Soc Trans"},{"key":"BFEMBJ2009372_CR25","doi-asserted-by":"crossref","first-page":"5864","DOI":"10.1038\/sj.onc.1209874","volume":"25","author":"K Gudmundsdottir","year":"2006","unstructured":"Gudmundsdottir K, Ashworth A (2006) The roles of BRCA1 and BRCA2 and associated proteins in the maintenance of genomic stability. Oncogene 25: 5864\u20135874","journal-title":"Oncogene"},{"key":"BFEMBJ2009372_CR26","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1016\/S0092-8674(00)81626-8","volume":"95","author":"JE Haber","year":"1998","unstructured":"Haber JE (1998) The many interfaces of Mre11. Cell 95: 583\u2013586","journal-title":"Cell"},{"key":"BFEMBJ2009372_CR27","doi-asserted-by":"crossref","first-page":"669","DOI":"10.1084\/jem.20081326","volume":"206","author":"BA Helmink","year":"2009","unstructured":"Helmink BA, Bredemeyer AL, Lee BS, Huang CY, Sharma GG, Walker LM, Bednarski JJ, Lee WL, Pandita TK, Bassing CH, Sleckman BP (2009) MRN complex function in the repair of chromosomal Rag\u2010mediated DNA double\u2010strand breaks. J Exp Med 206: 669\u2013679","journal-title":"J Exp Med"},{"key":"BFEMBJ2009372_CR28","doi-asserted-by":"crossref","first-page":"406","DOI":"10.1038\/nature03114","volume":"432","author":"Y Huyen","year":"2004","unstructured":"Huyen Y, Zgheib O, Ditullio Jr RA, Gorgoulis VG, Zacharatos P, Petty TJ, Sheston EA, Mellert HS, Stavridi ES, Halazonetis TD (2004) Methylated lysine 79 of histone H3 targets 53BP1 to DNA double\u2010strand breaks. Nature 432: 406\u2013411","journal-title":"Nature"},{"key":"BFEMBJ2009372_CR29","doi-asserted-by":"crossref","first-page":"6098","DOI":"10.1073\/pnas.91.13.6098","volume":"91","author":"K Iwabuchi","year":"1994","unstructured":"Iwabuchi K, Bartel PL, Li B, Marraccino R, Fields S (1994) Two cellular proteins that bind to wild\u2010type but not mutant p53. Proc Natl Acad Sci USA 91: 6098\u20136102","journal-title":"Proc Natl Acad Sci USA"},{"key":"BFEMBJ2009372_CR30","doi-asserted-by":"crossref","first-page":"36487","DOI":"10.1074\/jbc.M304066200","volume":"278","author":"K Iwabuchi","year":"2003","unstructured":"Iwabuchi K, Basu BP, Kysela B, Kurihara T, Shibata M, Guan D, Cao Y, Hamada T, Imamura K, Jeggo PA, Date T, Doherty AJ (2003) Potential role for 53BP1 in DNA end\u2010joining repair through direct interaction with DNA. J Biol Chem 278: 36487\u201336495","journal-title":"J Biol Chem"},{"key":"BFEMBJ2009372_CR31","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1101\/gad.959202","volume":"16","author":"WS Joo","year":"2002","unstructured":"Joo WS, Jeffrey PD, Cantor SB, Finnin MS, Livingston DM, Pavletich NP (2002) Structure of the 53BP1 BRCT region bound to p53 and its comparison to the Brca1 BRCT structure. Genes Dev 16: 583\u2013593","journal-title":"Genes Dev"},{"key":"BFEMBJ2009372_CR32","doi-asserted-by":"crossref","first-page":"55562","DOI":"10.1074\/jbc.M411021200","volume":"279","author":"PA Jowsey","year":"2004","unstructured":"Jowsey PA, Doherty AJ, Rouse J (2004) Human PTIP facilitates ATM\u2010mediated activation of p53 and promotes cellular resistance to ionizing radiation. J Biol Chem 279: 55562\u201355569","journal-title":"J Biol Chem"},{"key":"BFEMBJ2009372_CR33","doi-asserted-by":"crossref","first-page":"1423","DOI":"10.1101\/gad.1200304","volume":"18","author":"R Kitagawa","year":"2004","unstructured":"Kitagawa R, Bakkenist CJ, McKinnon PJ, Kastan MB (2004) Phosphorylation of SMC1 is a critical downstream event in the ATM\u2010NBS1\u2010BRCA1 pathway. Genes Dev 18: 1423\u20131438","journal-title":"Genes Dev"},{"key":"BFEMBJ2009372_CR34","doi-asserted-by":"crossref","first-page":"266","DOI":"10.1038\/ng0796-266","volume":"13","author":"EV Koonin","year":"1996","unstructured":"Koonin EV, Altschul SF, Bork P (1996) BRCA1 protein products. Functional motifs. Nat Genet 13: 266\u2013268","journal-title":"Nat Genet"},{"key":"BFEMBJ2009372_CR35","doi-asserted-by":"crossref","first-page":"759","DOI":"10.1038\/nrm2514","volume":"9","author":"MF Lavin","year":"2008","unstructured":"Lavin MF (2008) Ataxia\u2010telangiectasia: from a rare disorder to a paradigm for cell signalling and cancer. Nat Rev Mol Cell Biol 9: 759\u2013769","journal-title":"Nat Rev Mol Cell Biol"},{"key":"BFEMBJ2009372_CR36","first-page":"123","volume":"569","author":"MF Lavin","year":"2005","unstructured":"Lavin MF, Birrell G, Chen P, Kozlov S, Scott S, Gueven N (2005) ATM signaling and genomic stability in response to DNA damage. Mutat Res 569: 123\u2013132","journal-title":"Mutat Res"},{"key":"BFEMBJ2009372_CR37","doi-asserted-by":"crossref","first-page":"45171","DOI":"10.1074\/jbc.M308705200","volume":"278","author":"J\u2010H Lee","year":"2003","unstructured":"Lee J\u2010H, Ghirlando R, Bhaskara V, Hoffmeyer MR, Gu J, Paull TT (2003) Regulation of Mre11\/Rad50 by Nbs1: effects on nucleotide\u2010dependent DNA binding and association with ATLD mutant complexes. J Biol Chem 278: 45171\u201345181","journal-title":"J Biol Chem"},{"key":"BFEMBJ2009372_CR38","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1126\/science.1091496","volume":"304","author":"JH Lee","year":"2004","unstructured":"Lee JH, Paull TT (2004) Direct activation of the ATM protein kinase by the Mre11\/Rad50\/Nbs1 complex. Science 304: 93\u201396","journal-title":"Science"},{"key":"BFEMBJ2009372_CR39","doi-asserted-by":"crossref","first-page":"551","DOI":"10.1126\/science.1108297","volume":"308","author":"JH Lee","year":"2005","unstructured":"Lee JH, Paull TT (2005) ATM activation by DNA double\u2010strand breaks through the Mre11\u2010Rad50\u2010Nbs1 complex. Science 308: 551\u2013554","journal-title":"Science"},{"key":"BFEMBJ2009372_CR40","doi-asserted-by":"crossref","first-page":"529","DOI":"10.1016\/S0076-6879(06)08033-5","volume":"408","author":"JH Lee","year":"2006","unstructured":"Lee JH, Paull TT (2006) Purification and biochemical characterization of ataxia\u2010telangiectasia mutated and Mre11\/Rad50\/Nbs1. Methods Enzymol 408: 529\u2013539","journal-title":"Methods Enzymol"},{"key":"BFEMBJ2009372_CR41","doi-asserted-by":"crossref","first-page":"613","DOI":"10.1038\/35007091","volume":"404","author":"DS Lim","year":"2000","unstructured":"Lim DS, Kim ST, Xu B, Maser RS, Lin J, Petrini JH, Kastan MB (2000) ATM phosphorylates p95\/nbs1 in an S\u2010phase checkpoint pathway. Nature 404: 613\u2013617","journal-title":"Nature"},{"key":"BFEMBJ2009372_CR42","doi-asserted-by":"crossref","first-page":"481","DOI":"10.1038\/ni1067","volume":"5","author":"JP Manis","year":"2004","unstructured":"Manis JP, Morales JC, Xia Z, Kutok JL, Alt FW, Carpenter PB (2004) 53BP1 links DNA damage\u2010response pathways to immunoglobulin heavy chain class\u2010switch recombination. Nat Immunol 5: 481\u2013487","journal-title":"Nat Immunol"},{"key":"BFEMBJ2009372_CR43","doi-asserted-by":"crossref","first-page":"636","DOI":"10.1126\/science.1088877","volume":"302","author":"IA Manke","year":"2003","unstructured":"Manke IA, Lowery DM, Nguyen A, Yaffe MB (2003) BRCT repeats as phosphopeptide\u2010binding modules involved in protein targeting. Science 302: 636\u2013639","journal-title":"Science"},{"key":"BFEMBJ2009372_CR44","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1038\/86920","volume":"27","author":"RS Maser","year":"2001","unstructured":"Maser RS, Zinkel R, Petrini JH (2001) An alternative mode of translation permits production of a variant NBS1 protein from the common Nijmegen breakage syndrome allele. Nat Genet 27: 417\u2013421","journal-title":"Nat Genet"},{"key":"BFEMBJ2009372_CR45","doi-asserted-by":"crossref","first-page":"1160","DOI":"10.1126\/science.1140321","volume":"316","author":"S Matsuoka","year":"2007","unstructured":"Matsuoka S, Ballif BA, Smogorzewska A, McDonald III ER, Hurov KE, Luo J, Bakalarski CE, Zhao Z, Solimini N, Lerenthal Y, Shiloh Y, Gygi SP, Elledge SJ (2007) ATM and ATR substrate analysis reveals extensive protein networks responsive to DNA damage. Science 316: 1160\u20131166","journal-title":"Science"},{"key":"BFEMBJ2009372_CR46","first-page":"8586","volume":"63","author":"TA Mochan","year":"2003","unstructured":"Mochan TA, Venere M, DiTullio Jr RA, Halazonetis TD (2003) 53BP1 and NFBD1\/MDC1\u2010Nbs1 function in parallel interacting pathways activating ataxia\u2010telangiectasia mutated (ATM) in response to DNA damage. Cancer Res 63: 8586\u20138591","journal-title":"Cancer Res"},{"key":"BFEMBJ2009372_CR47","doi-asserted-by":"crossref","first-page":"945","DOI":"10.1016\/j.dnarep.2004.03.017","volume":"3","author":"TA Mochan","year":"2004","unstructured":"Mochan TA, Venere M, DiTullio Jr RA, Halazonetis TD (2004) 53BP1, an activator of ATM in response to DNA damage. DNA Repair (Amst) 3: 945\u2013952","journal-title":"DNA Repair (Amst)"},{"key":"BFEMBJ2009372_CR48","doi-asserted-by":"crossref","first-page":"937","DOI":"10.1016\/j.jmb.2003.11.026","volume":"335","author":"G Moncalian","year":"2004","unstructured":"Moncalian G, Lengsfeld B, Bhaskara V, Hopfner KP, Karcher A, Alden E, Tainer JA, Paull TT (2004) The rad50 signature motif: essential to ATP binding and biological function. J Mol Biol 335: 937\u2013951","journal-title":"J Mol Biol"},{"key":"BFEMBJ2009372_CR49","doi-asserted-by":"crossref","first-page":"14971","DOI":"10.1074\/jbc.M212484200","volume":"278","author":"JC Morales","year":"2003","unstructured":"Morales JC, Xia Z, Lu T, Aldrich MB, Wang B, Rosales C, Kellems RE, Hittelman WN, Elledge SJ, Carpenter PB (2003) Role for the BRCA1 C\u2010terminal repeats (BRCT) protein 53BP1 in maintaining genomic stability. J Biol Chem 278: 14971\u201314977","journal-title":"J Biol Chem"},{"key":"BFEMBJ2009372_CR50","doi-asserted-by":"crossref","first-page":"5312","DOI":"10.1093\/nar\/gkm493","volume":"35","author":"IM Munoz","year":"2007","unstructured":"Munoz IM, Jowsey PA, Toth R, Rouse J (2007) Phospho\u2010epitope binding by the BRCT domains of hPTIP controls multiple aspects of the cellular response to DNA damage. Nucleic Acids Res 35: 5312\u20135322","journal-title":"Nucleic Acids Res"},{"key":"BFEMBJ2009372_CR51","doi-asserted-by":"crossref","first-page":"1957","DOI":"10.1101\/gad.1099003","volume":"17","author":"D Nakada","year":"2003","unstructured":"Nakada D, Matsumoto K, Sugimoto K (2003) ATM\u2010related Tel1 associates with double\u2010strand breaks through an Xrs2\u2010dependent mechanism. Genes Dev 17: 1957\u20131962","journal-title":"Genes Dev"},{"key":"BFEMBJ2009372_CR52","doi-asserted-by":"crossref","first-page":"913","DOI":"10.1038\/ncb879","volume":"4","author":"K Nakanishi","year":"2002","unstructured":"Nakanishi K, Taniguchi T, Ranganathan V, New HV, Moreau LA, Stotsky M, Mathew CG, Kastan MB, Weaver DT, D\u2032Andrea AD (2002) Interaction of FANCD2 and NBS1 in the DNA damage response. Nat Cell Biol 4: 913\u2013920","journal-title":"Nat Cell Biol"},{"key":"BFEMBJ2009372_CR53","doi-asserted-by":"crossref","first-page":"436","DOI":"10.1016\/j.dnarep.2009.01.013","volume":"8","author":"S Panier","year":"2009","unstructured":"Panier S, Durocher D (2009) Regulatory ubiquitylation in response to DNA double\u2010strand breaks. DNA Repair (Amst) 8: 436\u2013443","journal-title":"DNA Repair (Amst)"},{"key":"BFEMBJ2009372_CR54","doi-asserted-by":"crossref","first-page":"6086","DOI":"10.1073\/pnas.111125998","volume":"98","author":"TT Paull","year":"2001","unstructured":"Paull TT, Cortez D, Bowers B, Elledge SJ, Gellert M (2001) Direct DNA binding by Brca1. Proc Natl Acad Sci USA 98: 6086\u20136091","journal-title":"Proc Natl Acad Sci USA"},{"key":"BFEMBJ2009372_CR55","doi-asserted-by":"crossref","first-page":"6409","DOI":"10.1073\/pnas.110144297","volume":"97","author":"TT Paull","year":"2000","unstructured":"Paull TT, Gellert M (2000) A mechanistic basis for Mre11\u2010directed DNA joining at microhomologies. Proc Natl Acad Sci USA 97: 6409\u20136414","journal-title":"Proc Natl Acad Sci USA"},{"key":"BFEMBJ2009372_CR56","doi-asserted-by":"crossref","first-page":"613","DOI":"10.1083\/jcb.153.3.613","volume":"153","author":"I Rappold","year":"2001","unstructured":"Rappold I, Iwabuchi K, Date T, Chen J (2001) Tumor suppressor p53 binding protein 1 (53BP1) is involved in DNA damage\u2010signaling pathways. J Cell Biol 153: 613\u2013620","journal-title":"J Cell Biol"},{"key":"BFEMBJ2009372_CR57","doi-asserted-by":"crossref","first-page":"715","DOI":"10.1016\/j.molcel.2004.10.029","volume":"16","author":"E Riballo","year":"2004","unstructured":"Riballo E, Kuhne M, Rief N, Doherty A, Smith GC, Recio MJ, Reis C, Dahm K, Fricke A, Krempler A, Parker AR, Jackson SP, Gennery A, Jeggo PA, Lobrich M (2004) A pathway of double\u2010strand break rejoining dependent upon ATM, Artemis, and proteins locating to gamma\u2010H2AX foci. Mol Cell 16: 715\u2013724","journal-title":"Mol Cell"},{"key":"BFEMBJ2009372_CR58","doi-asserted-by":"crossref","first-page":"1381","DOI":"10.1083\/jcb.151.7.1381","volume":"151","author":"LB Schultz","year":"2000","unstructured":"Schultz LB, Chehab NH, Malikzay A, Halazonetis TD (2000) p53 binding protein 1 (53BP1) is an early participant in the cellular response to DNA double\u2010strand breaks. J Cell Biol 151: 1381\u20131390","journal-title":"J Cell Biol"},{"key":"BFEMBJ2009372_CR59","doi-asserted-by":"crossref","first-page":"405","DOI":"10.1016\/S1097-2765(04)00238-2","volume":"14","author":"EN Shiozaki","year":"2004","unstructured":"Shiozaki EN, Gu L, Yan N, Shi Y (2004) Structure of the BRCT repeats of BRCA1 bound to a BACH1 phosphopeptide: implications for signaling. Mol Cell 14: 405\u2013412","journal-title":"Mol Cell"},{"key":"BFEMBJ2009372_CR60","doi-asserted-by":"crossref","first-page":"551","DOI":"10.1016\/S0960-9822(99)80242-5","volume":"9","author":"J Soulier","year":"1999","unstructured":"Soulier J, Lowndes NF (1999) The BRCT domain of the S. cerevisiae checkpoint protein Rad9 mediates a Rad9\u2010Rad9 interaction after DNA damage. Curr Biol 9: 551\u2013554","journal-title":"Curr Biol"},{"key":"BFEMBJ2009372_CR61","doi-asserted-by":"crossref","first-page":"577","DOI":"10.1016\/S0092-8674(00)81547-0","volume":"99","author":"GS Stewart","year":"1999","unstructured":"Stewart GS, Maser RS, Stankovic T, Bressan DA, Kaplan MI, Jaspers NG, Raams A, Byrd PJ, Petrini JH, Taylor AM (1999) The DNA double\u2010strand break repair gene hMre11 is mutated in individuals with an ataxia\u2010telangiectasia\u2010like disorder. Cell 99: 577\u2013587","journal-title":"Cell"},{"key":"BFEMBJ2009372_CR62","doi-asserted-by":"crossref","first-page":"218","DOI":"10.1038\/nature05740","volume":"447","author":"TH Stracker","year":"2007","unstructured":"Stracker TH, Morales M, Couto SS, Hussein H, Petrini JH (2007) The carboxy terminus of NBS1 is required for induction of apoptosis by the MRE11 complex. Nature 447: 218\u2013221","journal-title":"Nature"},{"key":"BFEMBJ2009372_CR63","doi-asserted-by":"crossref","first-page":"845","DOI":"10.1016\/j.dnarep.2004.03.014","volume":"3","author":"TH Stracker","year":"2004","unstructured":"Stracker TH, Theunissen JW, Morales M, Petrini JH (2004) The Mre11 complex and the metabolism of chromosome breaks: the importance of communicating and holding things together. DNA Repair (Amst) 3: 845\u2013854","journal-title":"DNA Repair (Amst)"},{"key":"BFEMBJ2009372_CR64","doi-asserted-by":"crossref","first-page":"630","DOI":"10.1128\/MMBR.66.4.630-670.2002","volume":"66","author":"LS Symington","year":"2002","unstructured":"Symington LS (2002) Role of RAD52 epistasis group genes in homologous recombination and double\u2010strand break repair. Microbiol Mol Biol Rev 66: 630\u2013670","journal-title":"Microbiol Mol Biol Rev"},{"key":"BFEMBJ2009372_CR65","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1016\/j.molcel.2008.12.022","volume":"33","author":"T Usui","year":"2009","unstructured":"Usui T, Foster SS, Petrini JH (2009) Maintenance of the DNA\u2010damage checkpoint requires DNA\u2010damage\u2010induced mediator protein oligomerization. Mol Cell 33: 147\u2013159","journal-title":"Mol Cell"},{"key":"BFEMBJ2009372_CR66","doi-asserted-by":"crossref","first-page":"467","DOI":"10.1016\/S0092-8674(00)81174-5","volume":"93","author":"R Varon","year":"1998","unstructured":"Varon R, Vissinga C, Platzer M, Cerosaletti KM, Chrzanowska KH, Saar K, Beckmann G, Seemanova E, Cooper PR, Nowak NJ, Stumm M, Weemaes CM, Gatti RA, Wilson RK, Digweed M, Rosenthal A, Sperling K, Concannon P, Reis A (1998) Nibrin, a novel DNA double\u2010strand break repair protein, is mutated in Nijmegen breakage syndrome. Cell 93: 467\u2013476","journal-title":"Cell"},{"key":"BFEMBJ2009372_CR67","doi-asserted-by":"crossref","first-page":"1435","DOI":"10.1126\/science.1076182","volume":"298","author":"B Wang","year":"2002","unstructured":"Wang B, Matsuoka S, Carpenter PB, Elledge SJ (2002) 53BP1, a mediator of the DNA damage checkpoint. Science 298: 1435\u20131438","journal-title":"Science"},{"key":"BFEMBJ2009372_CR68","doi-asserted-by":"crossref","first-page":"38472","DOI":"10.1074\/jbc.M607577200","volume":"281","author":"I Ward","year":"2006","unstructured":"Ward I, Kim JE, Minn K, Chini CC, Mer G, Chen J (2006) The tandem BRCT domain of 53BP1 is not required for its repair function. J Biol Chem 281: 38472\u201338477","journal-title":"J Biol Chem"},{"key":"BFEMBJ2009372_CR69","doi-asserted-by":"crossref","first-page":"2556","DOI":"10.1128\/MCB.23.7.2556-2563.2003","volume":"23","author":"IM Ward","year":"2003","unstructured":"Ward IM, Minn K, van Deursen J, Chen J (2003) p53 binding protein 53BP1 is required for DNA damage responses and tumor suppression in mice. Mol Cell Biol 23: 2556\u20132563","journal-title":"Mol Cell Biol"},{"key":"BFEMBJ2009372_CR70","doi-asserted-by":"crossref","first-page":"459","DOI":"10.1083\/jcb.200403021","volume":"165","author":"IM Ward","year":"2004","unstructured":"Ward IM, Reina\u2010San\u2010Martin B, Olaru A, Minn K, Tamada K, Lau JS, Cascalho M, Chen L, Nussenzweig A, Livak F, Nussenzweig MC, Chen J (2004) 53BP1 is required for class switch recombination. J Cell Biol 165: 459\u2013464","journal-title":"J Cell Biol"},{"key":"BFEMBJ2009372_CR71","doi-asserted-by":"crossref","first-page":"477","DOI":"10.1038\/35013089","volume":"405","author":"X Wu","year":"2000","unstructured":"Wu X, Ranganathan V, Weisman DS, Heine WF, Ciccone DN, O\u2032Neill TB, Crick KE, Pierce KA, Lane WS, Rathbun G, Livingston DM, Weaver DT (2000) ATM phosphorylation of Nijmegen breakage syndrome protein is required in a DNA damage response. Nature 405: 477\u2013482","journal-title":"Nature"},{"key":"BFEMBJ2009372_CR72","doi-asserted-by":"crossref","first-page":"2708","DOI":"10.1074\/jbc.M007665200","volume":"276","author":"Z Xia","year":"2001","unstructured":"Xia Z, Morales JC, Dunphy WG, Carpenter PB (2001) Negative cell cycle regulation and DNA damage\u2010inducible phosphorylation of the BRCT protein 53BP1. J Biol Chem 276: 2708\u20132718","journal-title":"J Biol Chem"},{"key":"BFEMBJ2009372_CR73","doi-asserted-by":"crossref","first-page":"1045","DOI":"10.1016\/j.molcel.2007.12.005","volume":"28","author":"A Xie","year":"2007","unstructured":"Xie A, Hartlerode A, Stucki M, Odate S, Puget N, Kwok A, Nagaraju G, Yan C, Alt FW, Chen J, Jackson SP, Scully R (2007) Distinct roles of chromatin\u2010associated proteins MDC1 and 53BP1 in mammalian double\u2010strand break repair. Mol Cell 28: 1045\u20131057","journal-title":"Mol Cell"},{"key":"BFEMBJ2009372_CR74","doi-asserted-by":"crossref","first-page":"814","DOI":"10.1038\/nsmb.1640","volume":"16","author":"A Xie","year":"2009","unstructured":"Xie A, Kwok A, Scully R (2009) Role of mammalian Mre11 in classical and alternative nonhomologous end joining. Nat Struct Mol Biol 16: 814\u2013818","journal-title":"Nat Struct Mol Biol"},{"key":"BFEMBJ2009372_CR75","doi-asserted-by":"crossref","first-page":"3445","DOI":"10.1128\/MCB.21.10.3445-3450.2001","volume":"21","author":"B Xu","year":"2001","unstructured":"Xu B, Kim S, Kastan MB (2001) Involvement of Brca1 in S\u2010phase and G(2)\u2010phase checkpoints after ionizing irradiation. Mol Cell Biol 21: 3445\u20133450","journal-title":"Mol Cell Biol"},{"key":"BFEMBJ2009372_CR76","first-page":"4588","volume":"62","author":"B Xu","year":"2002","unstructured":"Xu B, O\u2032Donnell AH, Kim ST, Kastan MB (2002) Phosphorylation of serine 1387 in Brca1 is specifically required for the Atm\u2010mediated S\u2010phase checkpoint after ionizing irradiation. Cancer Res 62: 4588\u20134591","journal-title":"Cancer Res"},{"key":"BFEMBJ2009372_CR77","doi-asserted-by":"crossref","first-page":"571","DOI":"10.1101\/gad.970702","volume":"16","author":"PT Yazdi","year":"2002","unstructured":"Yazdi PT, Wang Y, Zhao S, Patel N, Lee EY, Qin J (2002) SMC1 is a downstream effector in the ATM\/NBS1 branch of the human S\u2010phase checkpoint. Genes Dev 16: 571\u2013582","journal-title":"Genes Dev"},{"key":"BFEMBJ2009372_CR78","doi-asserted-by":"crossref","first-page":"5363","DOI":"10.1128\/MCB.25.13.5363-5379.2005","volume":"25","author":"Z You","year":"2005","unstructured":"You Z, Chahwan C, Bailis J, Hunter T, Russell P (2005) ATM activation and its recruitment to damaged DNA require binding to the C terminus of Nbs1. Mol Cell Biol 25: 5363\u20135379","journal-title":"Mol Cell Biol"},{"key":"BFEMBJ2009372_CR79","doi-asserted-by":"crossref","first-page":"639","DOI":"10.1126\/science.1088753","volume":"302","author":"X Yu","year":"2003","unstructured":"Yu X, Chini CC, He M, Mer G, Chen J (2003) The BRCT domain is a phospho\u2010protein binding domain. Science 302: 639\u2013642","journal-title":"Science"},{"key":"BFEMBJ2009372_CR80","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1016\/j.radonc.2005.06.026","volume":"76","author":"O Zgheib","year":"2005","unstructured":"Zgheib O, Huyen Y, DiTullio Jr RA, Snyder A, Venere M, Stavridi ES, Halazonetis TD (2005) ATM signaling and 53BP1. Radiother Oncol 76: 119\u2013122","journal-title":"Radiother Oncol"},{"key":"BFEMBJ2009372_CR81","doi-asserted-by":"crossref","first-page":"473","DOI":"10.1038\/35013083","volume":"405","author":"S Zhao","year":"2000","unstructured":"Zhao S, Weng YC, Yuan SS, Lin YT, Hsu HC, Lin SC, Gerbino E, Song MH, Zdzienicka MZ, Gatti RA, Shay JW, Ziv Y, Shiloh Y, Lee EY (2000) Functional link between ataxia\u2010telangiectasia and Nijmegen breakage syndrome gene products. Nature 405: 473\u2013477","journal-title":"Nature"}],"container-title":["The EMBO Journal"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.wiley.com\/onlinelibrary\/tdm\/v1\/articles\/10.1038%2Femboj.2009.372","content-type":"unspecified","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1038\/emboj.2009.372.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1038\/emboj.2009.372","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1038\/emboj.2009.372.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,5,20]],"date-time":"2026-05-20T00:02:27Z","timestamp":1779235347000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1038\/emboj.2009.372"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2009,12,10]]},"references-count":81,"journal-issue":{"issue":"3","published-print":{"date-parts":[[2010,2,3]]}},"alternative-id":["BFEMBJ2009372"],"URL":"https:\/\/doi.org\/10.1038\/emboj.2009.372","relation":{},"ISSN":["0261-4189","1460-2075"],"issn-type":[{"value":"0261-4189","type":"print"},{"value":"1460-2075","type":"electronic"}],"subject":[],"published":{"date-parts":[[2009,12,10]]},"assertion":[{"value":"17 August 2009","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"11 November 2009","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"10 December 2009","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare that they have no conflict of interest.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Conflict of Interest"}},{"value":"This content has been made available to all.","name":"free","label":"Free to read"}]}}