{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T19:08:33Z","timestamp":1792782513799,"version":"4.5.0"},"reference-count":44,"publisher":"Springer Science and Business Media LLC","license":[{"start":{"date-parts":[[2026,9,25]],"date-time":"2026-09-25T00:00:00Z","timestamp":1790294400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"},{"start":{"date-parts":[[2026,9,25]],"date-time":"2026-09-25T00:00:00Z","timestamp":1790294400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"}],"funder":[{"name":"Ministerio de Economia y Competividad of Spain and by the Centre de Refer\u00e8ncia de R+D de Biotecnologia de la Generalitat de Catalunya.","award":["PID2021-125632OB-C22 and PID2024-159663OB-C21"],"award-info":[{"award-number":["PID2021-125632OB-C22 and PID2024-159663OB-C21"]}],"id":[{"id":"https:\/\/ror.org\/01bg62x04","id-type":"ROR","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Eur Biophys J"],"DOI":"10.1007\/s00249-026-01863-2","type":"journal-article","created":{"date-parts":[[2026,9,25]],"date-time":"2026-09-25T07:30:48Z","timestamp":1790321448000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["A hydrophobic vector association model of nucleosomes in chromatin"],"prefix":"10.1007","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-6559-7926","authenticated-orcid":false,"given":"\u00c1ngel","family":"Mozo-Villar\u00edas","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Enrique","family":"Querol","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Juan A.","family":"Cedano","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,9,25]]},"reference":[{"key":"1863_CR1","doi-asserted-by":"publisher","first-page":"e52","DOI":"10.1093\/nar\/gkab070","volume":"49","author":"Z Adhireksan","year":"2021","unstructured":"Adhireksan Z, Sharma D, Lee PL, Bao Q, Padavattan S, Shum WK, Davey GE, Davey CA (2021) Engineering nucleosomes for generating diverse chromatin assemblies. Nucleic Acids Res 49:e52. https:\/\/doi.org\/10.1093\/nar\/gkab070","journal-title":"Nucleic Acids Res"},{"key":"1863_CR2","doi-asserted-by":"publisher","unstructured":"Allahverdi A., Yang R., Korolev N., Fan Y., Davey C., Liu C-F., Nordenski\u00f6ld The effects of histone H4 tail on cation-induced chromatin folding and self-association (2011). Nucl. Ac. Res.. 39:1680\u20131691. https:\/\/https:\/\/doi.org\/10.1093\/nar\/gkq900","DOI":"10.1093\/nar\/gkq900"},{"key":"1863_CR3","doi-asserted-by":"publisher","first-page":"11170","DOI":"10.1073\/pnas.92.24.11170","volume":"92","author":"G Arents","year":"1995","unstructured":"Arents G, Moudrianakis EN (1995) The histone fold: a ubiquitous architectural motif utilized in DNA compaction and protein dimerization. Proc Natl Acad Sci U S A 92:11170\u201311174. https:\/\/doi.org\/10.1073\/pnas.92.24.11170","journal-title":"Proc Natl Acad Sci U S A"},{"key":"1863_CR4","doi-asserted-by":"publisher","first-page":"8783","DOI":"10.1093\/nar\/gks586","volume":"40","author":"A Bancaud","year":"2012","unstructured":"Bancaud A, Lavelle C, Huet S, Ellenberg J (2012) A fractal model for nuclear organization: current evidence and biological implications. Nucleic Acids Res 40:8783\u20138792. https:\/\/doi.org\/10.1093\/nar\/gks586","journal-title":"Nucleic Acids Res"},{"key":"1863_CR5","doi-asserted-by":"publisher","first-page":"3168","DOI":"10.1111\/febs.14495","volume":"285","author":"S Bhattacharyya","year":"2018","unstructured":"Bhattacharyya S, Mattiroli F, Luger K (2018) Archaeal DNA on the histone merry-go-round. FEBS J 285:3168\u20133174. https:\/\/doi.org\/10.1111\/febs.14495","journal-title":"FEBS J"},{"key":"1863_CR6","doi-asserted-by":"publisher","DOI":"10.1140\/epje\/s10189-023-00320-8","volume":"46","author":"J Cedano","year":"2023","unstructured":"Cedano J, Querol E, Mozo-Villar\u00edas A (2023) How hydrophobicity shapes the architecture of protein assemblies. Eur Phys J E 46:62. https:\/\/doi.org\/10.1140\/epje\/s10189-023-00320-8","journal-title":"Eur Phys J E"},{"key":"1863_CR7","doi-asserted-by":"publisher","first-page":"257","DOI":"10.1007\/s00249-025-01764-w","volume":"54","author":"J Cedano","year":"2025","unstructured":"Cedano J, Querol E, Mozo-Villar\u00edas A (2025) Amino acids hydrophobic properties in proteins are derived from their atomic polarities. Eur Biophys J 54:257\u2013265. https:\/\/doi.org\/10.1007\/s00249-025-01764-w","journal-title":"Eur Biophys J"},{"key":"1863_CR8","doi-asserted-by":"publisher","first-page":"489","DOI":"10.1007\/s13238-014-0080-x","volume":"5","author":"P Chen","year":"2014","unstructured":"Chen P, Zhu P, Li G (2014) New insights into the helical structure of 30-nm chromatin fibers. Protein Cell 5:489\u2013491. https:\/\/doi.org\/10.1007\/s13238-014-0080-x","journal-title":"Protein Cell"},{"key":"1863_CR9","doi-asserted-by":"publisher","first-page":"053001","DOI":"10.1088\/1478-3975\/ac0aff","volume":"18","author":"JR Dab\u00e1n","year":"2021","unstructured":"Dab\u00e1n JR (2021) Soft-matter properties of multilayer chromosomes. Phys Biol 18:053001. https:\/\/doi.org\/10.1088\/1478-3975\/ac0aff","journal-title":"Phys Biol"},{"key":"1863_CR10","doi-asserted-by":"publisher","unstructured":"Dab\u00e1n J.R. Rethinking Models of DNA Organization in Micrometer-Sized Chromosomes from the Perspective of the Nanoproperties of Chromatin Favoring a Multilayer Structure (2024). Small Struc. 5:2400203. https:\/\/https:\/\/doi.org\/10.1002\/sstr.202400203.","DOI":"10.1002\/sstr.202400203"},{"key":"1863_CR11","doi-asserted-by":"publisher","first-page":"1097","DOI":"10.1016\/S0022-2836(02)00386-8","volume":"319","author":"CA Davey","year":"2002","unstructured":"Davey CA, Sargent DF, Luger K, Maeder AW, Richmond TJ (2002) Solvent mediated interactions in the structure of the nucleosome core particle at 1.9\u00c5 resolution. J Mol Biol 319:1097\u20131113. https:\/\/doi.org\/10.1016\/S0022-2836(02)00386-8","journal-title":"J Mol Biol"},{"key":"1863_CR12","doi-asserted-by":"publisher","first-page":"493","DOI":"10.1038\/s41594-022-00768-w","volume":"29","author":"M Dombrowski","year":"2022","unstructured":"Dombrowski M, Engeholm M, Dienemann C, Dodonova S, Cramer P (2022) Histone H1 binding to nucleosome arrays depends on linker DNA length and trajectory. Nat Struct Mol Biol 29:493\u2013501. https:\/\/doi.org\/10.1038\/s41594-022-00768-w","journal-title":"Nat Struct Mol Biol"},{"key":"1863_CR13","doi-asserted-by":"publisher","first-page":"2051","DOI":"10.1021\/la502115g","volume":"31","author":"SH Donaldson","year":"2015","unstructured":"Donaldson SH, R\u00f8yne A, Kristiansen K, Rapp MV, Das S, Gebbie MA, Lee DW, Stock P, Valtiner M, Israelachvili J (2015) Developing a general interaction potential for hydrophobic and hydrophilic interactions. Langmuir 31:2051\u20132064. https:\/\/doi.org\/10.1021\/la502115g","journal-title":"Langmuir"},{"key":"1863_CR14","doi-asserted-by":"publisher","first-page":"321","DOI":"10.1146\/annurev-biophys-070317-032847","volume":"48","author":"B Fierz","year":"2019","unstructured":"Fierz B, Poirier MG (2019) Biophysics of chromatin dynamics. Annu Rev Biophys 48:321\u2013345. https:\/\/doi.org\/10.1146\/annurev-biophys-070317-032847","journal-title":"Annu Rev Biophys"},{"key":"1863_CR15","doi-asserted-by":"crossref","unstructured":"Finch J.T., Klug A. Solenoidal model for superstructure in chromatin (1976). Proc. Natl. Acad. Sci. (USA), 73:1879\u20131901","DOI":"10.1073\/pnas.73.6.1897"},{"key":"1863_CR16","doi-asserted-by":"publisher","first-page":"13317","DOI":"10.1073\/pnas.0903280106","volume":"106","author":"SA Grigoryev","year":"2009","unstructured":"Grigoryev SA, Arya G, Correll S, Woodcock CL, Schlick T (2009) Evidence for heteromorphic chromatin fibers from analysis of nucleosome interactions. Proc Natl Acad Sci U S A 106:13317\u201313322. https:\/\/doi.org\/10.1073\/pnas.0903280106","journal-title":"Proc Natl Acad Sci U S A"},{"key":"1863_CR17","doi-asserted-by":"publisher","first-page":"341","DOI":"10.1038\/300341a0","volume":"300","author":"J Israelachvili","year":"1982","unstructured":"Israelachvili J, Pashley R (1982) The hydrophobic interaction is long range, decaying exponentially with distance. Nature 300:341\u2013342. https:\/\/doi.org\/10.1038\/300341a0","journal-title":"Nature"},{"key":"1863_CR18","doi-asserted-by":"publisher","first-page":"20121022","DOI":"10.1098\/rsif.2012.1022","volume":"10","author":"AA Kalashnikova","year":"2013","unstructured":"Kalashnikova AA, Porter-Goff ME, Muthurajan UM, Luger K, Hansen JC (2013) The role of the nucleosome acidic patch in modulating higher order chromatin structure. J R Soc Interface 10:20121022. https:\/\/doi.org\/10.1098\/rsif.2012.1022","journal-title":"J R Soc Interface"},{"key":"1863_CR19","unstructured":"Kornberg R.D. The nucleosome:\u00a0historical perspective and current view (2007). Cold Spring Harbor Symp Quant. Biol., 72:1\u20135"},{"key":"1863_CR20","doi-asserted-by":"publisher","first-page":"865","DOI":"10.1126\/science.184.4139.865","volume":"184","author":"RD Kornberg","year":"1974","unstructured":"Kornberg RD, Thomas JO (1974) Chromatin structure: oligomers of the histones. Science 184:865\u2013868. https:\/\/doi.org\/10.1126\/science.184.4139.865","journal-title":"Science"},{"key":"1863_CR21","doi-asserted-by":"crossref","unstructured":"Kornberg R.D. Chromatin structure: a repeating unit of histones and DNA (1974). Science, 184:868\u2013871","DOI":"10.1126\/science.184.4139.868"},{"key":"1863_CR22","doi-asserted-by":"publisher","first-page":"289","DOI":"10.1126\/science.1181369","volume":"326","author":"E Lieberman-Aiden","year":"2009","unstructured":"Lieberman-Aiden E, van Berkum NL, Williams L, Imakaev M, Ragoczy T, Telling A, Amit I, Lajoie BR, Sabo PJ, Dorschner MO, Sandstrom R, Bernstein B, Bender MA, Groudine M, Gnirke A, Stamatoyannopoulos J, Mirny LA, Lander ES, Dekker J (2009) Comprehensive mapping of long-range interactions reveals folding principles of the human genome. Science 326:289\u2013293. https:\/\/doi.org\/10.1126\/science.1181369","journal-title":"Science"},{"key":"1863_CR23","doi-asserted-by":"publisher","first-page":"251","DOI":"10.1038\/38444","volume":"389","author":"K Luger","year":"1997","unstructured":"Luger K, M\u00e4der AW, Richmond RK, Sargent DF, Richmond TJ (1997) Crystal structure of the nucleosome core particle at 2.8\u00c5 resolution. Nature 389:251\u2013260. https:\/\/doi.org\/10.1038\/38444","journal-title":"Nature"},{"key":"1863_CR24","doi-asserted-by":"publisher","first-page":"291","DOI":"10.1016\/j.ceb.2010.03.001","volume":"22","author":"K Maeshima","year":"2010","unstructured":"Maeshima K, Hihara S, Eltsov M (2010) Chromatin structure: does the 30-nm fibre exist in vivo? Curr Opin Cell Biol 22:291\u2013297. https:\/\/doi.org\/10.1016\/j.ceb.2010.03.001","journal-title":"Curr Opin Cell Biol"},{"key":"1863_CR25","doi-asserted-by":"publisher","first-page":"609","DOI":"10.1126\/science.aaj1849","volume":"357","author":"F Mattiroli","year":"2017","unstructured":"Mattiroli F, Bhattacharyya S, Dyer PN, White AE, Sandman K, Burkhart BW, Byrne KR, Lee T, Ahn NG, Santangelo TJ, Reeve JN, Luger K (2017) Structure of histone-base chromatin in Archaea. Science 357:609\u2013612. https:\/\/doi.org\/10.1126\/science.aaj1849","journal-title":"Science"},{"key":"1863_CR26","doi-asserted-by":"publisher","first-page":"54","DOI":"10.1016\/j.sbi.2015.11.014","volume":"37","author":"R McGinty","year":"2016","unstructured":"McGinty R, Tan S (2016) Recognition of the nucleosome by chromatin factors and enzymes. Curr Opin Struct Biol 37:54\u201361. https:\/\/doi.org\/10.1016\/j.sbi.2015.11.014","journal-title":"Curr Opin Struct Biol"},{"key":"1863_CR27","doi-asserted-by":"publisher","first-page":"15739","DOI":"10.1073\/pnas.0606422103","volume":"103","author":"E Meyer","year":"2006","unstructured":"Meyer E, Rosenberg K, Israelachvili J (2006) Recent progress in understanding hydrophobic interactions. Proc Natl Acad Sci U S A 103:15739\u201315746. https:\/\/doi.org\/10.1073\/pnas.0606422103","journal-title":"Proc Natl Acad Sci U S A"},{"key":"1863_CR28","doi-asserted-by":"publisher","first-page":"951","DOI":"10.1007\/s00249-021-01557-x","volume":"50","author":"A Mozo-Villar\u00edas","year":"2021","unstructured":"Mozo-Villar\u00edas A, Cedano J, Querol E (2021) The importance of hydrophobic interactions in the structure of transcription systems. Eur Biophys J 50:951\u2013961. https:\/\/doi.org\/10.1007\/s00249-021-01557-x","journal-title":"Eur Biophys J"},{"key":"1863_CR29","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1017\/S0033583522000038","volume":"55","author":"A Mozo-Villar\u00edas","year":"2022","unstructured":"Mozo-Villar\u00edas A, Cedano J, Querol E (2022) Use of vector formalism in the analysis of hydrophobic and electric driving forces in biological assemblies. Quarterly Reviews of Biopysics 55:1\u201322. https:\/\/doi.org\/10.1017\/S0033583522000038","journal-title":"Quarterly Reviews of Biopysics"},{"key":"1863_CR30","doi-asserted-by":"publisher","first-page":"231","DOI":"10.1007\/s00249-025-01754-y","volume":"54","author":"A Mozo-Villar\u00edas","year":"2025","unstructured":"Mozo-Villar\u00edas A, Cedano J, Querol E (2025) The influence of the Debye\u2013H\u00fcckel factor in estimating the distance between interacting monomers in self\u2011assembling proteins. Eur Biophys J 54:231\u2013246. https:\/\/doi.org\/10.1007\/s00249-025-01754-y","journal-title":"Eur Biophys J"},{"key":"1863_CR31","doi-asserted-by":"publisher","first-page":"1644","DOI":"10.1038\/emboj.2012.35","volume":"31","author":"Y Nishino","year":"2012","unstructured":"Nishino Y, Eltsov M, Joti Y, Ito K, Takata H, Tajahashi Y, Hihara S, Frangakis AS, Imamoto N, Ishikawa T, Maeshima K (2012) Human mitotic chromosomes consist predominantly of irregularly folded nucleosome fibres without a 30-nm chromatin structure. The EMBO J 31:1644\u20131653","journal-title":"The EMBO J"},{"key":"1863_CR32","doi-asserted-by":"publisher","first-page":"330","DOI":"10.1126\/science.183.4122.330","volume":"183","author":"AL Olins","year":"1974","unstructured":"Olins AL, Olins DE (1974) Shperoid chromatin units (v bodies). Science 183:330\u2013332. https:\/\/doi.org\/10.1126\/science.183.4122.330","journal-title":"Science"},{"key":"1863_CR33","doi-asserted-by":"publisher","DOI":"10.1126\/science.aag0025","volume":"357","author":"HD Ou","year":"2017","unstructured":"Ou HD, Phan S, Deerinck TJ, Thor A, Ellisman MH, O\u2019Shea C (2017) ChromEMT: visualizing 3D chromatin structure and compaction in interphase and mitotic cells. Science 357:eaag0025. https:\/\/doi.org\/10.1126\/science.aag0025","journal-title":"Science"},{"key":"1863_CR34","doi-asserted-by":"publisher","first-page":"281","DOI":"10.1016\/0092-8674(75)90149-X","volume":"4","author":"P Oudet","year":"1975","unstructured":"Oudet P, Gross-Bellard M, Chambon P (1975) Electron microscopy and biochemical evidence that chromatin structureis a repeating unit. Cell 4:281\u2013300","journal-title":"Cell"},{"key":"1863_CR35","doi-asserted-by":"publisher","first-page":"2498","DOI":"10.1021\/jp2107523","volume":"116","author":"AJ Patel","year":"2012","unstructured":"Patel AJ, Varilly P, Jamadagni SN, Hagan MF, Chandler D, Garde S (2012) Sitting at the edge: how biomolecules use hydrophobicity to tune their interactions and function. J Phys Chem B 116:2498\u20132503. https:\/\/doi.org\/10.1021\/jp2107523","journal-title":"J Phys Chem B"},{"key":"1863_CR36","doi-asserted-by":"publisher","unstructured":"Robinson P.J., Rhodes D. Structure of the \u201830 nm\u2019 chromatin fibre: a key role for the linker histone (2006). Curr Opin. Struct. Biol. 16:336\u2013346. https:\/\/doi.https:\/\/doi.org\/10.1016\/j.sbi.2006.05.007.","DOI":"10.1016\/j.sbi.2006.05.007"},{"key":"1863_CR37","doi-asserted-by":"publisher","unstructured":"Rydberg B., Holley W.R., Mian I.S., Chatterjee A. Chromatin Conformation in Living Cell: Support for a Zig-Zag Model of the 30 nm Chromatin Fiber (1998). J. Mol. Biol., 284:71-84. https:\/\/doi.org\/10.1006\/jmbi.1998.2150","DOI":"10.1006\/jmbi.1998.2150"},{"key":"1863_CR38","doi-asserted-by":"publisher","first-page":"138","DOI":"10.1038\/nature03686","volume":"436","author":"T Schalch","year":"2005","unstructured":"Schalch T, Duda S, Sargent DF, Richmond TJ (2005) X-ray structure of a tetranucleosome and its implications for the chromatin fibre. Nature 436:138\u2013141","journal-title":"Nature"},{"key":"1863_CR39","doi-asserted-by":"crossref","unstructured":"Scheffer M.P., Eltsov M., Frangakis A.S. Evidence for short-range helical order in the 30-nm chromatin fibers of erythocyte nuclei (2011). Proc. Nat. Acad. Sci. (USA) 108:16992\u201316997","DOI":"10.1073\/pnas.1108268108"},{"key":"1863_CR40","doi-asserted-by":"publisher","first-page":"3872","DOI":"10.1021\/jz402068k","volume":"4","author":"R Tabor","year":"2013","unstructured":"Tabor R, Wu C, Grieser F, Dagastine RR, Chan DY (2013) Measurement of the hydrophobic force in a soft matter system. J Phys Chem Lett 4:3872\u20133877. https:\/\/doi.org\/10.1021\/jz402068k","journal-title":"J Phys Chem Lett"},{"key":"1863_CR41","doi-asserted-by":"crossref","unstructured":"Tachiwana H., Kagawa W., Osakabe A., Kawaguci K., Shiga T., Hayashi-Takanaka Y., Kimura H., Kurumizaka H. Structural basis of instability of the nucleosome containing a testis-specific histone variant, human H3T (2010). Proc. Natl. Acad. Sci. (USA), 107:10454\u201310459","DOI":"10.1073\/pnas.1003064107"},{"key":"1863_CR42","doi-asserted-by":"publisher","first-page":"403","DOI":"10.1083\/jcb.83.2.403","volume":"83","author":"F Thoma","year":"1979","unstructured":"Thoma F, Koller T, Klug A (1979) Involvement of histone H1 in the organization of the nucleosome and of the salt-dependent superstructures of chromatin. J Cell Biol 83:403\u2013427. https:\/\/doi.org\/10.1083\/jcb.83.2.403","journal-title":"J Cell Biol"},{"key":"1863_CR43","doi-asserted-by":"publisher","DOI":"10.3389\/fphy.2018.00087","volume":"6","author":"J Van der Gucht","year":"2018","unstructured":"Van der Gucht J (2018) Grand challenges in soft matter physics. Front Phys 6:87","journal-title":"Front Phys"},{"key":"1863_CR44","doi-asserted-by":"publisher","first-page":"1129","DOI":"10.1038\/sj.embor.7401115","volume":"8","author":"C Wu","year":"2007","unstructured":"Wu C, Bassett A, Travers A (2007) A variable topologyfor the 30-nm chromatin fibre. EMBO Rep 8:1129\u20131134. https:\/\/doi.org\/10.1038\/sj.embor.7401115","journal-title":"EMBO Rep"}],"container-title":["European Biophysics Journal"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00249-026-01863-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s00249-026-01863-2","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00249-026-01863-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,25]],"date-time":"2026-09-25T20:05:28Z","timestamp":1790366728000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s00249-026-01863-2"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,25]]},"references-count":44,"alternative-id":["1863"],"URL":"https:\/\/doi.org\/10.1007\/s00249-026-01863-2","relation":{},"ISSN":["0175-7571","1432-1017"],"issn-type":[{"value":"0175-7571","type":"print"},{"value":"1432-1017","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,25]]},"assertion":[{"value":"5 December 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"6 July 2026","order":2,"name":"revised","label":"Revised","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"25 August 2026","order":3,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"25 September 2026","order":4,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}}]}}