{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,25]],"date-time":"2026-10-25T09:20:57Z","timestamp":1792920057035,"version":"4.5.0"},"reference-count":48,"publisher":"Springer Science and Business Media LLC","issue":"9","license":[{"start":{"date-parts":[[2002,5,1]],"date-time":"2002-05-01T00:00:00Z","timestamp":1020211200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springer.com\/tdm"},{"start":{"date-parts":[[2002,5,1]],"date-time":"2002-05-01T00:00:00Z","timestamp":1020211200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springer.com\/tdm"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["EMBO J"],"published-print":{"date-parts":[[2002,5,1]]},"DOI":"10.1093\/emboj\/21.9.2055","type":"journal-article","created":{"date-parts":[[2002,7,26]],"date-time":"2002-07-26T18:55:22Z","timestamp":1027709722000},"page":"2055-2067","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":95,"title":["Three\u2010dimensional structure of the human transglutaminase 3 enzyme: binding of calcium ions changes structure for activation"],"prefix":"10.1038","volume":"21","author":[{"given":"Bijan","family":"Ahvazi","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hee Chul","family":"Kim","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sun\u2010Ho","family":"Kee","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zoltan","family":"Nemes","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Peter M.","family":"Steinert","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2002,5,1]]},"reference":[{"key":"BFEMBJ7594425_CR1","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1007\/BF00223514","volume":"85","author":"KE Achyuthan","year":"1989","unstructured":"Achyuthan KE, Mary A, Bhaerman R and Grenberg CS (1989) Enzymatic activity of guinea pig liver transglutaminase. Mol Cell Biochem, 85, 57\u201365.","journal-title":"Mol Cell Biochem"},{"key":"BFEMBJ7594425_CR2","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1016\/0014-5793(88)80928-1","volume":"239","author":"CM Bergamini","year":"1988","unstructured":"Bergamini CM (1988) GTP modulates calcium binding and cation\u2010induced conformational change in erythrocyte transglutaminase. FEBS Lett, 239, 255\u2013258.","journal-title":"FEBS Lett"},{"key":"BFEMBJ7594425_CR3","doi-asserted-by":"crossref","first-page":"192","DOI":"10.1107\/S0108768190011041","volume":"47","author":"NE Brese","year":"1991","unstructured":"Brese NE and O'Keeffe M (1991) Bond\u2013valence parameters for solids. Acta Crystallogr B, 47, 192\u2013197.","journal-title":"Acta Crystallogr B"},{"key":"BFEMBJ7594425_CR4","doi-asserted-by":"crossref","first-page":"585","DOI":"10.1107\/S0108767390002355","volume":"46","author":"AT Br\u00fcnger","year":"1990","unstructured":"Br\u00fcnger AT, Krukowski A and Erickson JW (1990) Slow\u2010cooling protocols for crystallographic refinement by simulated annealing. Acta Crystallogr A, 46, 585\u2013593.","journal-title":"Acta Crystallogr A"},{"key":"BFEMBJ7594425_CR5","doi-asserted-by":"crossref","first-page":"905","DOI":"10.1107\/S0907444998003254","volume":"54","author":"AT Br\u00fcnger","year":"1998","unstructured":"Br\u00fcnger AT et al. (1998) Crystallography and NMR system: a new software suite for macromolecular structure determination. Acta Crystallogr D, 54, 905\u2013921.","journal-title":"Acta Crystallogr D"},{"key":"BFEMBJ7594425_CR6","doi-asserted-by":"crossref","first-page":"26382","DOI":"10.1074\/jbc.270.44.26382","volume":"270","author":"E Candi","year":"1995","unstructured":"Candi E, Melino G, Mei G, Tarcsa E, Chung SI, Marekov LN and Steinert PM (1995) Biochemical, structural and transglutaminase substrate properties of human loricrin, the major epidermal cornified cell envelope protein. J Biol Chem, 270, 26382\u201326890.","journal-title":"J Biol Chem"},{"key":"BFEMBJ7594425_CR7","doi-asserted-by":"crossref","first-page":"7226","DOI":"10.1074\/jbc.274.11.7226","volume":"274","author":"E Candi","year":"1999","unstructured":"Candi E, Tarcsa E, Idler WW, Kartasova T, Marekov LN and Steinert PM (1999) Transglutaminase cross\u2010linking properties of the small proline\u2010rich 1 family of cornified cell envelope proteins. Integration with loricrin. J Biol Chem, 274, 7226\u20137237.","journal-title":"J Biol Chem"},{"key":"BFEMBJ7594425_CR8","doi-asserted-by":"crossref","unstructured":"CCP4 (1994) The CCP4 suite: programs for protein crystallography. Acta Crystallogr D, 50, 760\u2013763.","DOI":"10.1107\/S0907444994003112"},{"key":"BFEMBJ7594425_CR9","doi-asserted-by":"crossref","first-page":"8703","DOI":"10.1074\/jbc.275.12.8703","volume":"275","author":"YC Choi","year":"2000","unstructured":"Choi YC, Park GT, Kim TS, Sunwoo IN, Steinert PM and Kim SY (2000) Sporadic inclusion body myositis correlates with increased expression and cross\u2010linking by transglutaminases 1 and 2. J Biol Chem, 275, 8703\u20138710.","journal-title":"J Biol Chem"},{"key":"BFEMBJ7594425_CR10","doi-asserted-by":"crossref","first-page":"232","DOI":"10.1107\/S0907444999016352","volume":"56","author":"Z Dauter","year":"2000","unstructured":"Dauter Z, Dauter M and Rajashankar KR (2000) Novel approach to phasing proteins: derivatization by short cryo\u2010soaking with halides. Acta Crystallogr D, 56, 232\u2013237.","journal-title":"Acta Crystallogr D"},{"key":"BFEMBJ7594425_CR11","doi-asserted-by":"crossref","first-page":"358","DOI":"10.1016\/S0076-6879(85)13049-1","volume":"113","author":"JE Folk","year":"1985","unstructured":"Folk JE and Chung SI (1985) Transglutaminases. Methods Enzymol, 113, 358\u2013375.","journal-title":"Methods Enzymol"},{"key":"BFEMBJ7594425_CR12","doi-asserted-by":"crossref","first-page":"244","DOI":"10.1016\/0926-6593(66)90066-X","volume":"122","author":"JE Folk","year":"1966","unstructured":"Folk JE and Cole PW (1966) Transglutaminase: mechanistic features of the active site as determined by kinetic and inhibitor studies. Biochim Biophys Acta, 122, 244\u2013264.","journal-title":"Biochim Biophys Acta"},{"key":"BFEMBJ7594425_CR13","doi-asserted-by":"crossref","first-page":"4917","DOI":"10.1074\/jbc.274.8.4917","volume":"274","author":"BA Fox","year":"1999","unstructured":"Fox BA, Yee VC, Pedersen LC, Le Trong I, Bishop PD, Stenkamp RE and Teller DC (1999) Identification of the calcium binding site and a novel ytterbium site in blood coagulation factor XIII by X\u2010ray crystallography. J Biol Chem, 274, 4917\u20134923.","journal-title":"J Biol Chem"},{"key":"BFEMBJ7594425_CR14","doi-asserted-by":"crossref","first-page":"273","DOI":"10.1016\/0049-3848(83)90081-6","volume":"30","author":"JA Gladner","year":"1983","unstructured":"Gladner JA and Nossal R (1983) Effects of crosslinking on the rigidity and proteolytic susceptibility of human fibrin clots. Thromb Res, 30, 273\u2013288.","journal-title":"Thromb Res"},{"key":"BFEMBJ7594425_CR15","doi-asserted-by":"crossref","first-page":"3071","DOI":"10.1096\/fasebj.5.15.1683845","volume":"5","author":"CS Greenberg","year":"1991","unstructured":"Greenberg CS, Birckbichler PJ and Rice RH (1991) Transglut aminases: multifunctional cross\u2010linking enzymes that stabilize tissues. FASEB J, 5, 3071\u20133077.","journal-title":"FASEB J"},{"key":"BFEMBJ7594425_CR16","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1107\/S0108767390010224","volume":"47","author":"TA Jones","year":"1991","unstructured":"Jones TA, Zou JY, Cowan SW and Kjeldgaard M (1991) Improved methods for building protein models in electron density maps and the location of errors in these models. Acta Crystallogr A, 47, 110\u2013119.","journal-title":"Acta Crystallogr A"},{"key":"BFEMBJ7594425_CR17","doi-asserted-by":"crossref","first-page":"21971","DOI":"10.1016\/S0021-9258(18)45834-6","volume":"265","author":"HC Kim","year":"1990","unstructured":"Kim HC, Lewis MS, Gorman JJ, Park SC, Girard JE, Folk JE and Chung SI (1990) Protransglutaminase E from guinea pig skin. Isolation and partial characterization. J Biol Chem, 265, 21971\u201321978.","journal-title":"J Biol Chem"},{"key":"BFEMBJ7594425_CR18","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1006\/jsbi.2001.4384","volume":"135","author":"HC Kim","year":"2001","unstructured":"Kim HC, Nemes Z, Idler WW, Hyde CC, Steinert PM and Ahvazi B (2001) Crystallization and preliminary X\u2010ray analysis of human transglutaminase 3 from zymogen to active form. J Struct Biol, 135, 73\u201377.","journal-title":"J Struct Biol"},{"key":"BFEMBJ7594425_CR19","doi-asserted-by":"crossref","first-page":"12682","DOI":"10.1016\/S0021-9258(18)31442-X","volume":"268","author":"IG Kim","year":"1993","unstructured":"Kim IG, Gorman JJ, Park SC, Chung SI and Steinert PM (1993) The deduced sequence of the novel protransglutaminase E (TGase3) of human and mouse. J Biol Chem, 268, 12682\u201312690.","journal-title":"J Biol Chem"},{"key":"BFEMBJ7594425_CR20","doi-asserted-by":"crossref","first-page":"30715","DOI":"10.1074\/jbc.274.43.30715","volume":"274","author":"SY Kim","year":"1999","unstructured":"Kim SY, Grant P, Lee JH, Pant HC and Steinert PM (1999) Differential expression of multiple transglutaminases in human brain. Increased expression and cross\u2010linking by transglutaminases 1 and 2 in Alzheimer's disease. J Biol Chem, 274, 30715\u201330721.","journal-title":"J Biol Chem"},{"key":"BFEMBJ7594425_CR21","doi-asserted-by":"crossref","first-page":"842","DOI":"10.1107\/S0907444995016477","volume":"52","author":"GJ Kleywegt","year":"1996","unstructured":"Kleywegt GJ (1996) Use of non\u2010crystallographic symmetry in protein structure refinement. Acta Crystallogr D, 52, 842\u2013857.","journal-title":"Acta Crystallogr D"},{"key":"BFEMBJ7594425_CR22","doi-asserted-by":"crossref","first-page":"1557","DOI":"10.1016\/S0969-2126(97)00305-5","volume":"5","author":"GJ Kleywegt","year":"1997","unstructured":"Kleywegt GJ and Read RJ (1997) Not your average density. Structure, 5, 1557\u20131569.","journal-title":"Structure"},{"key":"BFEMBJ7594425_CR23","doi-asserted-by":"crossref","first-page":"946","DOI":"10.1107\/S0021889891004399","volume":"24","author":"PJ Kraulis","year":"1991","unstructured":"Kraulis PJ (1991) MOLSCRIPT\u2014a program to produce both detailed and schematic plots of protein structures. J Appl Crystallogr, 24, 946\u2013950.","journal-title":"J Appl Crystallogr"},{"key":"BFEMBJ7594425_CR24","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1107\/S0021889892009944","volume":"26","author":"RA Laskowski","year":"1992","unstructured":"Laskowski RA, MacArthur MW, Moss DS and Thornton JM (1992) PROCHECK: a program to check the stereochemical quality of protein structures. J Appl Crystallogr, 26, 283\u2013291.","journal-title":"J Appl Crystallogr"},{"key":"BFEMBJ7594425_CR26","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1007\/BF00240602","volume":"58","author":"L Lorand","year":"1984","unstructured":"Lorand L and Conrad SM (1984) Transglutaminases. Mol Cell Biochem, 58, 9\u201335.","journal-title":"Mol Cell Biochem"},{"key":"BFEMBJ7594425_CR25","doi-asserted-by":"crossref","first-page":"308","DOI":"10.1021\/bi00375a043","volume":"26","author":"L Lorand","year":"1987","unstructured":"Lorand L, Barnes N, Bruner\u2010Lorand JA, Hawkins M and Michalska M (1987) Inhibition of protein cross\u2010linking in Ca2+\u2010enriched human erythrocytes and activated platelets. Biochemistry, 26, 308\u2013313.","journal-title":"Biochemistry"},{"key":"BFEMBJ7594425_CR27","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1016\/0022-2836(68)90205-2","volume":"33","author":"BW Matthews","year":"1968","unstructured":"Matthews BW (1968) Solvent content of protein crystals. J Mol Biol, 33, 491\u2013497.","journal-title":"J Mol Biol"},{"key":"BFEMBJ7594425_CR28","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1007\/978-3-540-69185-3_9","volume":"24","author":"G Melino","year":"1998","unstructured":"Melino G, De Laurenzi V, Catani MV, Terrinoni A, Ciani B, Candi E, Marekov L and Steinert PM (1998) The cornified envelope: a model of cell death in the skin. Results Probl Cell Differ, 24, 175\u2013212.","journal-title":"Results Probl Cell Differ"},{"key":"BFEMBJ7594425_CR29","doi-asserted-by":"crossref","first-page":"505","DOI":"10.1016\/S0076-6879(97)77028-9","volume":"277","author":"EA Merritt","year":"1997","unstructured":"Merritt EA and Bacon DJ (1997) Raster3D: photorealistic molecular graphics. Methods Enzymol, 277, 505\u2013525.","journal-title":"Methods Enzymol"},{"key":"BFEMBJ7594425_CR30","doi-asserted-by":"crossref","first-page":"581","DOI":"10.1016\/S0076-6879(97)76079-8","volume":"276","author":"J Navaza","year":"1997","unstructured":"Navaza J and Saludjian P (1997) aMoRe: an automated molecular replacement program package. Methods Enzymol, 276, 581\u2013594.","journal-title":"Methods Enzymol"},{"key":"BFEMBJ7594425_CR31","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1038\/emm.1999.2","volume":"31","author":"Z Nemes","year":"1999","unstructured":"Nemes Z and Steinert PM (1999) Bricks and mortar of the epidermal barrier. Exp Mol Med, 31, 5\u201319.","journal-title":"Exp Mol Med"},{"key":"BFEMBJ7594425_CR32","doi-asserted-by":"crossref","first-page":"8402","DOI":"10.1073\/pnas.96.15.8402","volume":"96","author":"Z Nemes","year":"1999","unstructured":"Nemes Z, Marekov LN, Fesus L and Steinert PM (1999) A novel function for transglutaminase 1: attachment of long\u2010chain \u03c9\u2010hydroxyceramides to involucrin by ester bond formation. Proc Natl Acad Sci USA, 96, 8402\u20138407.","journal-title":"Proc Natl Acad Sci USA"},{"key":"BFEMBJ7594425_CR33","doi-asserted-by":"crossref","first-page":"12055","DOI":"10.1074\/jbc.M009862200","volume":"276","author":"K Noguchi","year":"2001","unstructured":"Noguchi K, Ishikawa K, Yokoyama K, Ohtsuka T, Nio N and Suzuki E (2001) Crystal structure of red sea bream transglutaminase. J Biol Chem, 276, 12055\u201312059.","journal-title":"J Biol Chem"},{"key":"BFEMBJ7594425_CR34","doi-asserted-by":"crossref","first-page":"307","DOI":"10.1016\/S0076-6879(97)76066-X","volume":"276","author":"Z Otwinowski","year":"1997","unstructured":"Otwinowski Z and Minor W (1997) Processing of X\u2010ray diffraction data collected in oscillation mode. Methods Enzymol, 276, 307\u2013326.","journal-title":"Methods Enzymol"},{"key":"BFEMBJ7594425_CR35","doi-asserted-by":"crossref","first-page":"1131","DOI":"10.1002\/pro.5560030720","volume":"3","author":"LC Pedersen","year":"1994","unstructured":"Pedersen LC, Yee VC, Bishop PD, Le Trong I, Teller DC and Stenkamp RE (1994) Transglutaminase factor XIII uses proteinase\u2010like catalytic triad to crosslink macromolecules. Protein Sci, 3, 1131\u20131135.","journal-title":"Protein Sci"},{"key":"BFEMBJ7594425_CR36","doi-asserted-by":"crossref","first-page":"892","DOI":"10.1126\/science.160.3830.892","volume":"160","author":"JJ Pisano","year":"1968","unstructured":"Pisano JJ, Finlayson JS and Peyton MP (1968) Cross\u2010link in fibrin polymerized by factor 13: \u03f5\u2010(\u03b3\u2010glutamyl)lysine. Science, 160, 892\u2013893.","journal-title":"Science"},{"key":"BFEMBJ7594425_CR37","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1016\/0092-8674(77)90059-9","volume":"11","author":"RH Rice","year":"1977","unstructured":"Rice RH and Green H (1977) The cornified envelope of terminally differentiated human epidermal keratinocytes consists of cross\u2010linked protein. Cell, 11, 417\u2013422.","journal-title":"Cell"},{"key":"BFEMBJ7594425_CR38","doi-asserted-by":"crossref","first-page":"1336","DOI":"10.1172\/JCI110885","volume":"71","author":"L Shen","year":"1983","unstructured":"Shen L and Lorand L (1983) Contribution of fibrin stabilization to clot strength. Supplementation of factor XIII\u2010deficient plasma with the purified zymogen. J Clin Invest, 71, 1336\u20131341.","journal-title":"J Clin Invest"},{"key":"BFEMBJ7594425_CR39","doi-asserted-by":"crossref","first-page":"17702","DOI":"10.1074\/jbc.270.30.17702","volume":"270","author":"PM Steinert","year":"1995","unstructured":"Steinert PM and Marekov LN (1995) The proteins elafin, filaggrin, keratin intermediate filaments, loricrin and small proline\u2010rich proteins 1 and 2 are isodipeptide cross\u2010linked components of the human epidermal cornified cell envelope. J Biol Chem, 270, 17702\u201317711.","journal-title":"J Biol Chem"},{"key":"BFEMBJ7594425_CR40","doi-asserted-by":"crossref","first-page":"26242","DOI":"10.1074\/jbc.271.42.26242","volume":"271","author":"PM Steinert","year":"1996","unstructured":"Steinert PM, Kim SY, Chung SI and Marekov LN (1996) The transglutaminase 1 enzyme is variably acylated by myristate and palmitate during differentiation in epidermal keratinocytes. J Biol Chem, 271, 26242\u201326250.","journal-title":"J Biol Chem"},{"key":"BFEMBJ7594425_CR41","doi-asserted-by":"crossref","first-page":"11758","DOI":"10.1074\/jbc.273.19.11758","volume":"273","author":"PM Steinert","year":"1998","unstructured":"Steinert PM, Kartasova T and Marekov LN (1998) Biochemical evidence that small proline\u2010rich proteins and trichohyalin function in epithelia by modulation of the biomechanical properties of their cornified cell envelopes. J Biol Chem, 273, 11758\u201311769.","journal-title":"J Biol Chem"},{"key":"BFEMBJ7594425_CR42","doi-asserted-by":"crossref","first-page":"916","DOI":"10.1038\/sj.cdd.4400568","volume":"6","author":"PM Steinert","year":"1999","unstructured":"Steinert PM, Candi E, Tarcsa E, Marekov LN, Sette M, Paci M, Ciani B, Guerrieri P and Melino G (1999) Transglutaminase crosslinking and structural studies of the human small proline rich 3 protein. Cell Death Differ, 6, 916\u2013930.","journal-title":"Cell Death Differ"},{"key":"BFEMBJ7594425_CR43","doi-asserted-by":"crossref","first-page":"27893","DOI":"10.1074\/jbc.272.44.27893","volume":"272","author":"E Tarcsa","year":"1997","unstructured":"Tarcsa E, Marekov LN, Andreoli J, Idler WW, Candi E, Chung SI and Steinert PM (1997) The fate of trichohyalin. Sequential post\u2010translational modifications by peptidyl\u2010arginine deiminase and transglutaminases. J Biol Chem, 272, 27893\u201327901.","journal-title":"J Biol Chem"},{"key":"BFEMBJ7594425_CR44","doi-asserted-by":"crossref","first-page":"23297","DOI":"10.1074\/jbc.273.36.23297","volume":"273","author":"E Tarcsa","year":"1998","unstructured":"Tarcsa E, Candi E, Kartasova T, Idler WW, Marekov LN and Steinert PM (1998) Structural and transglutaminase substrate properties of the small proline\u2010rich 2 family of cornified cell envelope proteins. J Biol Chem, 273, 23297\u201323303.","journal-title":"J Biol Chem"},{"key":"BFEMBJ7594425_CR45","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1016\/S0014-5793(98)00098-2","volume":"423","author":"MS Weiss","year":"1998","unstructured":"Weiss MS, Metzner HJ and Hilgenfeld R (1998) Two non\u2010proline cis peptide bonds may be important for factor XIII function. FEBS Lett, 423, 291\u2013296.","journal-title":"FEBS Lett"},{"key":"BFEMBJ7594425_CR46","doi-asserted-by":"crossref","first-page":"2041","DOI":"10.1271\/bbb.58.2041","volume":"58","author":"H Yasueda","year":"1994","unstructured":"Yasueda H, Kumazawa Y and Motoki M (1994) Purification and characterization of a tissue\u2010type transglutaminase from red sea bream (Pagrus major). Biosci Biotechnol Biochem, 58, 2041\u20132045.","journal-title":"Biosci Biotechnol Biochem"},{"key":"BFEMBJ7594425_CR47","doi-asserted-by":"crossref","first-page":"7296","DOI":"10.1073\/pnas.91.15.7296","volume":"91","author":"VC Yee","year":"1994","unstructured":"Yee VC, Pedersen LC, Le Trong I, Bishop PD, Stenkamp RE and Teller DC (1994) Three\u2010dimensional structure of a transglutaminase: human blood coagulation factor XIII. Proc Natl Acad Sci USA, 91, 7296\u2013300.","journal-title":"Proc Natl Acad Sci USA"},{"key":"BFEMBJ7594425_CR48","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1055\/s-2007-999035","volume":"22","author":"VC Yee","year":"1996","unstructured":"Yee VC, Le Trong I, Bishop PD, Pedersen LC, Stenkamp RE and Teller DC (1996) Structure and function studies of factor XIIIa by X\u2010ray crystallography. Semin Thromb Hemost, 22, 377\u2013384.","journal-title":"Semin Thromb Hemost"}],"container-title":["The EMBO Journal"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.wiley.com\/onlinelibrary\/tdm\/v1\/articles\/10.1093%2Femboj%2F21.9.2055","content-type":"unspecified","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1093\/emboj\/21.9.2055.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1093\/emboj\/21.9.2055\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full\/10.1093\/emboj\/21.9.2055","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1093\/emboj\/21.9.2055.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,5,20]],"date-time":"2026-05-20T18:56:52Z","timestamp":1779303412000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1093\/emboj\/21.9.2055"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2002,5,1]]},"references-count":48,"journal-issue":{"issue":"9","published-print":{"date-parts":[[2002,5,1]]}},"alternative-id":["BFEMBJ7594425"],"URL":"https:\/\/doi.org\/10.1093\/emboj\/21.9.2055","relation":{},"ISSN":["0261-4189","1460-2075"],"issn-type":[{"value":"0261-4189","type":"print"},{"value":"1460-2075","type":"electronic"}],"subject":[],"published":{"date-parts":[[2002,5,1]]},"assertion":[{"value":"28 January 2002","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"11 March 2002","order":2,"name":"revised","label":"Revised","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"11 March 2002","order":3,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"1 May 2002","order":4,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"This content has been made available to all.","name":"free","label":"Free to read"}]}}