{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,31]],"date-time":"2026-03-31T22:12:06Z","timestamp":1774995126860,"version":"3.50.1"},"reference-count":45,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2005,12,20]],"date-time":"2005-12-20T00:00:00Z","timestamp":1135036800000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/www.springer.com\/tdm"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Nat Struct Mol Biol"],"published-print":{"date-parts":[[2006,1]]},"DOI":"10.1038\/nsmb1040","type":"journal-article","created":{"date-parts":[[2005,12,20]],"date-time":"2005-12-20T15:09:05Z","timestamp":1135091345000},"page":"35-43","source":"Crossref","is-referenced-by-count":89,"title":["Structure of Saccharomyces cerevisiae DNA polymerase epsilon by cryo\u2013electron microscopy"],"prefix":"10.1038","volume":"13","author":[{"given":"Francisco J","family":"Asturias","sequence":"first","affiliation":[]},{"given":"Iris K","family":"Cheung","sequence":"additional","affiliation":[]},{"given":"Nasim","family":"Sabouri","sequence":"additional","affiliation":[]},{"given":"Olga","family":"Chilkova","sequence":"additional","affiliation":[]},{"given":"Daniel","family":"Wepplo","sequence":"additional","affiliation":[]},{"given":"Erik","family":"Johansson","sequence":"additional","affiliation":[]}],"member":"297","published-online":{"date-parts":[[2005,12,20]]},"reference":[{"key":"BFnsmb1040_CR1","first-page":"RE3","volume":"2003","author":"PV Shcherbakova","year":"2003","unstructured":"Shcherbakova, P.V., Bebenek, K. & Kunkel, T.A. Functions of eukaryotic DNA polymerases. Sci. Aging Knowledge Environ. [online] 2003, RE3 (2003).","journal-title":"Sci. Aging Knowledge Environ. [online]"},{"key":"BFnsmb1040_CR2","doi-asserted-by":"publisher","first-page":"137","DOI":"10.1016\/S0065-3233(04)69005-X","volume":"69","author":"K Bebenek","year":"2004","unstructured":"Bebenek, K. & Kunkel, T.A. Functions of DNA polymerases. Adv. Protein Chem. 69, 137\u2013165 (2004).","journal-title":"Adv. Protein Chem."},{"key":"BFnsmb1040_CR3","doi-asserted-by":"publisher","first-page":"54","DOI":"10.1016\/S0959-440X(98)80010-9","volume":"8","author":"CA Brautigam","year":"1998","unstructured":"Brautigam, C.A. & Steitz, T.A. Structural and functional insights provided by crystal structures of DNA polymerases and their substrate complexes. Curr. Opin. Struct. Biol. 8, 54\u201363 (1998).","journal-title":"Curr. Opin. Struct. Biol."},{"key":"BFnsmb1040_CR4","doi-asserted-by":"publisher","first-page":"4978","DOI":"10.1073\/pnas.081088798","volume":"98","author":"S Waga","year":"2001","unstructured":"Waga, S., Masuda, T., Takisawa, H. & Sugino, A. DNA polymerase epsilon is required for coordinated and efficient chromosomal DNA replication in Xenopus egg extracts. Proc. Natl. Acad. Sci. USA 98, 4978\u20134983 (2001).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"BFnsmb1040_CR5","doi-asserted-by":"publisher","first-page":"179","DOI":"10.1111\/j.1356-9597.2004.00716.x","volume":"9","author":"T Fukui","year":"2004","unstructured":"Fukui, T. et al. Distinct roles of DNA polymerases delta and epsilon at the replication fork in Xenopus egg extracts. Genes Cells 9, 179\u2013191 (2004).","journal-title":"Genes Cells"},{"key":"BFnsmb1040_CR6","doi-asserted-by":"publisher","first-page":"2809","DOI":"10.1128\/MCB.20.8.2809-2817.2000","volume":"20","author":"H Masumoto","year":"2000","unstructured":"Masumoto, H., Sugino, A. & Araki, H. Dpb11 controls the association between DNA polymerases alpha and epsilon and the autonomously replicating sequence region of budding yeast. Mol. Cell. Biol. 20, 2809\u20132817 (2000).","journal-title":"Mol. Cell. Biol."},{"key":"BFnsmb1040_CR7","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1016\/S0092-8674(01)80009-X","volume":"91","author":"OM Aparicio","year":"1997","unstructured":"Aparicio, O.M., Weinstein, D.M. & Bell, S.P. Components and dynamics of DNA replication complexes in S. cerevisiae: redistribution of MCM proteins and Cdc45p during S phase. Cell 91, 59\u201369 (1997).","journal-title":"Cell"},{"key":"BFnsmb1040_CR8","doi-asserted-by":"crossref","first-page":"717","DOI":"10.1093\/genetics\/142.3.717","volume":"142","author":"PV Shcherbakova","year":"1996","unstructured":"Shcherbakova, P.V. & Pavlov, Y.I. 3\u2032 \u2192 5\u2032 exonucleases of DNA polymerases epsilon and delta correct base analog induced DNA replication errors on opposite DNA strands in Saccharomyces cerevisiae. Genetics 142, 717\u2013726 (1996).","journal-title":"Genetics"},{"key":"BFnsmb1040_CR9","doi-asserted-by":"publisher","first-page":"405","DOI":"10.1006\/jmbi.2000.3744","volume":"299","author":"R Karthikeyan","year":"2000","unstructured":"Karthikeyan, R. et al. Evidence from mutational specificity studies that yeast DNA polymerases delta and epsilon replicate different DNA strands at an intracellular replication fork. J. Mol. Biol. 299, 405\u2013419 (2000).","journal-title":"J. Mol. Biol."},{"key":"BFnsmb1040_CR10","doi-asserted-by":"publisher","first-page":"14082","DOI":"10.1074\/jbc.M211818200","volume":"278","author":"O Chilkova","year":"2003","unstructured":"Chilkova, O., Jonsson, B.H. & Johansson, E. The quaternary structure of DNA polymerase epsilon from Saccharomyces cerevisiae. J. Biol. Chem. 278, 14082\u201314086 (2003).","journal-title":"J. Biol. Chem."},{"key":"BFnsmb1040_CR11","doi-asserted-by":"publisher","first-page":"43824","DOI":"10.1074\/jbc.M108842200","volume":"276","author":"E Johansson","year":"2001","unstructured":"Johansson, E., Majka, J. & Burgers, P.M. Structure of DNA polymerase delta from Saccharomyces cerevisiae. J. Biol. Chem. 276, 43824\u201343828 (2001).","journal-title":"J. Biol. Chem."},{"key":"BFnsmb1040_CR12","doi-asserted-by":"publisher","first-page":"5122","DOI":"10.1073\/pnas.091095198","volume":"98","author":"YH Jin","year":"2001","unstructured":"Jin, Y.H. et al. The 3\u2032 \u2192 5\u2032 exonuclease of DNA polymerase delta can substitute for the 5\u2032 flap endonuclease Rad27\/Fen1 in processing Okazaki fragments and preventing genome instability. Proc. Natl. Acad. Sci. USA 98, 5122\u20135127 (2001).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"BFnsmb1040_CR13","doi-asserted-by":"publisher","first-page":"1618","DOI":"10.1074\/jbc.M209801200","volume":"278","author":"R Ayyagari","year":"2003","unstructured":"Ayyagari, R., Gomes, X.V., Gordenin, D.A. & Burgers, P.M. Okazaki fragment maturation in yeast. I. Distribution of functions between FEN1 AND DNA2. J. Biol. Chem. 278, 1618\u20131625 (2003).","journal-title":"J. Biol. Chem."},{"key":"BFnsmb1040_CR14","doi-asserted-by":"publisher","first-page":"1626","DOI":"10.1074\/jbc.M209803200","volume":"278","author":"YH Jin","year":"2003","unstructured":"Jin, Y.H., Ayyagari, R., Resnick, M.A., Gordenin, D.A. & Burgers, P.M. Okazaki fragment maturation in yeast. II. Cooperation between the polymerase and 3\u2032-5\u2032-exonuclease activities of Pol delta in the creation of a ligatable nick. J. Biol. Chem. 278, 1626\u20131633 (2003).","journal-title":"J. Biol. Chem."},{"key":"BFnsmb1040_CR15","doi-asserted-by":"publisher","first-page":"2764","DOI":"10.1101\/gad.1252304","volume":"18","author":"P Garg","year":"2004","unstructured":"Garg, P., Stith, C.M., Sabouri, N., Johansson, E. & Burgers, P.M. Idling by DNA polymerase delta maintains a ligatable nick during lagging-strand DNA replication. Genes Dev. 18, 2764\u20132773 (2004).","journal-title":"Genes Dev."},{"key":"BFnsmb1040_CR16","doi-asserted-by":"crossref","first-page":"4072","DOI":"10.1016\/S0021-9258(19)39704-2","volume":"265","author":"RK Hamatake","year":"1990","unstructured":"Hamatake, R.K. et al. Purification and characterization of DNA polymerase II from the yeast Saccharomyces cerevisiae. Identification of the catalytic core and a possible holoenzyme form of the enzyme. J. Biol. Chem. 265, 4072\u20134083 (1990).","journal-title":"J. Biol. Chem."},{"key":"BFnsmb1040_CR17","doi-asserted-by":"publisher","first-page":"9473","DOI":"10.1073\/pnas.88.21.9473","volume":"88","author":"A Morrison","year":"1991","unstructured":"Morrison, A., Bell, J.B., Kunkel, T.A. & Sugino, A. Eukaryotic DNA polymerase amino acid sequence required for 3\u2032\u2014-5\u2032 exonuclease activity. Proc. Natl. Acad. Sci. USA 88, 9473\u20139477 (1991).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"BFnsmb1040_CR18","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1016\/0092-8674(95)90448-4","volume":"80","author":"TA Navas","year":"1995","unstructured":"Navas, T.A., Zhou, Z. & Elledge, S.J. DNA polymerase epsilon links the DNA replication machinery to the S phase checkpoint. Cell 80, 29\u201339 (1995).","journal-title":"Cell"},{"key":"BFnsmb1040_CR19","doi-asserted-by":"publisher","first-page":"679","DOI":"10.1016\/S1097-2765(00)80361-5","volume":"3","author":"T Kesti","year":"1999","unstructured":"Kesti, T., Flick, K., Keranen, S., Syvaoja, J.E. & Wittenberg, C. DNA polymerase epsilon catalytic domains are dispensable for DNA replication, DNA repair, and cell viability. Mol. Cell 3, 679\u2013685 (1999).","journal-title":"Mol. Cell"},{"key":"BFnsmb1040_CR20","doi-asserted-by":"publisher","first-page":"14245","DOI":"10.1074\/jbc.M313289200","volume":"279","author":"T Kesti","year":"2004","unstructured":"Kesti, T., McDonald, W.H., Yates, J.R. III & Wittenberg, C. Cell cycle-dependent phosphorylation of the DNA polymerase epsilon subunit, Dpb2, by the Cdc28 cyclin-dependent protein kinase. J. Biol. Chem. 279, 14245\u201314255 (2004).","journal-title":"J. Biol. Chem."},{"key":"BFnsmb1040_CR21","doi-asserted-by":"publisher","first-page":"4601","DOI":"10.1073\/pnas.88.11.4601","volume":"88","author":"H Araki","year":"1991","unstructured":"Araki, H., Hamatake, R.K., Johnston, L.H. & Sugino, A. DPB2, the gene encoding DNA polymerase II subunit B, is required for chromosome replication in Saccharomyces cerevisiae. Proc. Natl. Acad. Sci. USA 88, 4601\u20134605 (1991).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"BFnsmb1040_CR22","doi-asserted-by":"publisher","first-page":"4867","DOI":"10.1093\/nar\/19.18.4867","volume":"19","author":"H Araki","year":"1991","unstructured":"Araki, H. et al. Cloning DPB3, the gene encoding the third subunit of DNA polymerase II of Saccharomyces cerevisiae. Nucleic Acids Res. 19, 4867\u20134872 (1991).","journal-title":"Nucleic Acids Res."},{"key":"BFnsmb1040_CR23","doi-asserted-by":"publisher","first-page":"3846","DOI":"10.1093\/nar\/28.20.3846","volume":"28","author":"T Ohya","year":"2000","unstructured":"Ohya, T., Maki, S., Kawasaki, Y. & Sugino, A. Structure and function of the fourth subunit (Dpb4p) of DNA polymerase epsilon in Saccharomyces cerevisiae. Nucleic Acids Res. 28, 3846\u20133852 (2000).","journal-title":"Nucleic Acids Res."},{"key":"BFnsmb1040_CR24","doi-asserted-by":"publisher","first-page":"23247","DOI":"10.1074\/jbc.M002548200","volume":"275","author":"Y Li","year":"2000","unstructured":"Li, Y., Pursell, Z.F. & Linn, S. Identification and cloning of two histone fold motif-containing subunits of HeLa DNA polymerase epsilon. J. Biol. Chem. 275, 23247\u201323252 (2000).","journal-title":"J. Biol. Chem."},{"key":"BFnsmb1040_CR25","doi-asserted-by":"publisher","first-page":"217","DOI":"10.1128\/MCB.24.1.217-227.2004","volume":"24","author":"T Iida","year":"2004","unstructured":"Iida, T. & Araki, H. Noncompetitive counteractions of DNA polymerase epsilon and ISW2\/yCHRAC for epigenetic inheritance of telomere position effect in Saccharomyces cerevisiae. Mol. Cell. Biol. 24, 217\u2013227 (2004).","journal-title":"Mol. Cell. Biol."},{"key":"BFnsmb1040_CR26","doi-asserted-by":"publisher","first-page":"873","DOI":"10.1046\/j.1365-2443.2003.00683.x","volume":"8","author":"T Tsubota","year":"2003","unstructured":"Tsubota, T., Maki, S., Kubota, H., Sugino, A. & Maki, H. Double-stranded DNA binding properties of Saccharomyces cerevisiae DNA polymerase epsilon and of the Dpb3p-Dpb4p subassembly. Genes Cells 8, 873\u2013888 (2003).","journal-title":"Genes Cells"},{"key":"BFnsmb1040_CR27","doi-asserted-by":"crossref","unstructured":"Frank, J. Three-Dimensional Electron Microscopy of Macromolecular Assemblies 342 (Academic Press, San Diego, USA, 1996).","DOI":"10.1016\/B978-012265040-6\/50002-3"},{"key":"BFnsmb1040_CR28","doi-asserted-by":"publisher","first-page":"359","DOI":"10.1002\/jemt.1060090405","volume":"9","author":"M Radermacher","year":"1988","unstructured":"Radermacher, M. The three-dimensional reconstruction of single particles from random and non-random tilt series. J. Electron Microsc. Tech. 9, 359\u2013394 (1988).","journal-title":"J. Electron Microsc. Tech."},{"key":"BFnsmb1040_CR29","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1016\/0304-3991(87)90010-6","volume":"21","author":"M van Heel","year":"1987","unstructured":"van Heel, M. Similarity measures between images. Ultramicroscopy 21, 95\u2013100 (1987).","journal-title":"Ultramicroscopy"},{"key":"BFnsmb1040_CR30","doi-asserted-by":"publisher","first-page":"251","DOI":"10.1016\/0304-3991(94)90038-8","volume":"53","author":"P Penczek","year":"1994","unstructured":"Penczek, P., Grassucci, R.A. & Frank, J. The ribosome at improved resolution: new techniques for merging and orientation refinement in 3D cryo-electron microscopy of biological particles. Ultramicroscopy 53, 251\u2013270 (1994).","journal-title":"Ultramicroscopy"},{"key":"BFnsmb1040_CR31","doi-asserted-by":"publisher","first-page":"28816","DOI":"10.1074\/jbc.M002376200","volume":"275","author":"R Dua","year":"2000","unstructured":"Dua, R., Edwards, S., Levy, D.L. & Campbell, J.L. Subunit interactions within the Saccharomyces cerevisiae DNA polymerase epsilon (pol epsilon) complex. Demonstration of a dimeric pol epsilon. J. Biol. Chem. 275, 28816\u201328825 (2000).","journal-title":"J. Biol. Chem."},{"key":"BFnsmb1040_CR32","doi-asserted-by":"publisher","first-page":"50537","DOI":"10.1074\/jbc.M308515200","volume":"278","author":"RJ Kokoska","year":"2003","unstructured":"Kokoska, R.J., McCulloch, S.D. & Kunkel, T.A. The efficiency and specificity of apurinic\/apyrimidinic site bypass by human DNA polymerase eta and Sulfolobus solfataricus Dpo4. J. Biol. Chem. 278, 50537\u201350545 (2003).","journal-title":"J. Biol. Chem."},{"key":"BFnsmb1040_CR33","doi-asserted-by":"publisher","first-page":"4665","DOI":"10.1093\/nar\/gkh777","volume":"32","author":"SD McCulloch","year":"2004","unstructured":"McCulloch, S.D. et al. Enzymatic switching for efficient and accurate translesion DNA replication. Nucleic Acids Res. 32, 4665\u20134675 (2004).","journal-title":"Nucleic Acids Res."},{"key":"BFnsmb1040_CR34","doi-asserted-by":"publisher","first-page":"4759","DOI":"10.1074\/jbc.270.9.4759","volume":"270","author":"S Creighton","year":"1995","unstructured":"Creighton, S. & Goodman, M.F. Gel kinetic analysis of DNA polymerase fidelity in the presence of proofreading using bacteriophage T4 DNA polymerase. J. Biol. Chem. 270, 4759\u20134774 (1995).","journal-title":"J. Biol. Chem."},{"key":"BFnsmb1040_CR35","doi-asserted-by":"publisher","first-page":"232","DOI":"10.1016\/0076-6879(95)62021-4","volume":"262","author":"S Creighton","year":"1995","unstructured":"Creighton, S., Bloom, L.B. & Goodman, M.F. Gel fidelity assay measuring nucleotide misinsertion, exonucleolytic proofreading, and lesion bypass efficiencies. Methods Enzymol. 262, 232\u2013256 (1995).","journal-title":"Methods Enzymol."},{"key":"BFnsmb1040_CR36","doi-asserted-by":"publisher","first-page":"715","DOI":"10.1038\/nature01377","volume":"421","author":"SA Burgess","year":"2003","unstructured":"Burgess, S.A., Walker, M.L., Sakakibara, H., Knight, P.J. & Oiwa, K. Dynein structure and power stroke. Nature 421, 715\u2013718 (2003).","journal-title":"Nature"},{"key":"BFnsmb1040_CR37","doi-asserted-by":"publisher","first-page":"247","DOI":"10.1016\/j.jsb.2004.04.004","volume":"147","author":"SA Burgess","year":"2004","unstructured":"Burgess, S.A., Walker, M.L., Thirumurugan, K., Trinick, J. & Knight, P.J. Use of negative stain and single-particle image processing to explore dynamic properties of flexible macromolecules. J. Struct. Biol. 147, 247\u2013258 (2004).","journal-title":"J. Struct. Biol."},{"key":"BFnsmb1040_CR38","doi-asserted-by":"publisher","first-page":"724","DOI":"10.1002\/bies.10127","volume":"24","author":"P Cramer","year":"2002","unstructured":"Cramer, P. Common structural features of nucleic acid polymerases. Bioessays 24, 724\u2013729 (2002).","journal-title":"Bioessays"},{"key":"BFnsmb1040_CR39","doi-asserted-by":"crossref","first-page":"22698","DOI":"10.1016\/S0021-9258(18)54625-1","volume":"266","author":"PM Burgers","year":"1991","unstructured":"Burgers, P.M. Saccharomyces cerevisiae replication factor C. II. Formation and activity of complexes with the proliferating cell nuclear antigen and with DNA polymerases delta and epsilon. J. Biol. Chem. 266, 22698\u201322706 (1991).","journal-title":"J. Biol. Chem."},{"key":"BFnsmb1040_CR40","doi-asserted-by":"publisher","first-page":"8658","DOI":"10.1074\/jbc.M110615200","volume":"277","author":"J Fuss","year":"2002","unstructured":"Fuss, J. & Linn, S. Human DNA polymerase epsilon colocalizes with proliferating cell nuclear antigen and DNA replication late, but not early, in S phase. J. Biol. Chem. 277, 8658\u20138666 (2002).","journal-title":"J. Biol. Chem."},{"key":"BFnsmb1040_CR41","doi-asserted-by":"publisher","first-page":"1087","DOI":"10.1016\/S0092-8674(00)80296-2","volume":"89","author":"J Wang","year":"1997","unstructured":"Wang, J. et al. Crystal structure of a pol alpha family replication DNA polymerase from bacteriophage RB69. Cell 89, 1087\u20131099 (1997).","journal-title":"Cell"},{"key":"BFnsmb1040_CR42","doi-asserted-by":"publisher","first-page":"724","DOI":"10.1038\/nature02585","volume":"429","author":"GD Bowman","year":"2004","unstructured":"Bowman, G.D., O'Donnell, M. & Kuriyan, J. Structural analysis of a eukaryotic sliding DNA clamp-clamp loader complex. Nature 429, 724\u2013730 (2004).","journal-title":"Nature"},{"key":"BFnsmb1040_CR43","doi-asserted-by":"publisher","first-page":"187","DOI":"10.1007\/BF00267715","volume":"134","author":"GW Tischendorf","year":"1974","unstructured":"Tischendorf, G.W., Zeichhardt, H. & Stoffler, G. Determination of the location of proteins L14, L17, L18, L19, L22, L23 on the surface of the 5oS ribosomal subunit of Escherichia coli by immune electron microscopy. Mol. Gen. Genet. 134, 187\u2013208 (1974).","journal-title":"Mol. Gen. Genet."},{"key":"BFnsmb1040_CR44","doi-asserted-by":"crossref","first-page":"409","DOI":"10.1515\/9783111439846-018","volume-title":"Modern Methods in Protein Chemistry","author":"G Stoffler","year":"1983","unstructured":"Stoffler, G. & Stoffler-Meilicke, M. The ultrastructure of macromolecular complexes studied with antibodies. in Modern Methods in Protein Chemistry (ed. Tesche, H.) 409\u2013455 (De Gruyter, Berlin, 1983)."},{"key":"BFnsmb1040_CR45","doi-asserted-by":"publisher","first-page":"869","DOI":"10.1107\/S0907444994006396","volume":"50","author":"EA Merritt","year":"1994","unstructured":"Merritt, E.A. & Murphy, M.E. Raster3D Version 2.0. A program for photorealistic molecular graphics. Acta Crystallogr. D Biol. Crystallogr. 50, 869\u2013873 (1994).","journal-title":"Acta Crystallogr. D Biol. Crystallogr."}],"container-title":["Nature Structural &amp; Molecular Biology"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/www.nature.com\/articles\/nsmb1040.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"http:\/\/www.nature.com\/articles\/nsmb1040","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"http:\/\/www.nature.com\/articles\/nsmb1040.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,5,19]],"date-time":"2023-05-19T04:48:14Z","timestamp":1684471694000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/nsmb1040"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2005,12,20]]},"references-count":45,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2006,1]]}},"alternative-id":["BFnsmb1040"],"URL":"https:\/\/doi.org\/10.1038\/nsmb1040","relation":{},"ISSN":["1545-9993","1545-9985"],"issn-type":[{"value":"1545-9993","type":"print"},{"value":"1545-9985","type":"electronic"}],"subject":[],"published":{"date-parts":[[2005,12,20]]}}}