{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,11]],"date-time":"2026-09-11T02:14:26Z","timestamp":1789092866596,"version":"build-2803163510"},"reference-count":66,"publisher":"MDPI AG","issue":"21","license":[{"start":{"date-parts":[[2023,11,1]],"date-time":"2023-11-01T00:00:00Z","timestamp":1698796800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100012166","name":"National Key R&amp;D Program of China","doi-asserted-by":"publisher","award":["2020YFA0908100"],"award-info":[{"award-number":["2020YFA0908100"]}],"id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012166","name":"National Key R&amp;D Program of China","doi-asserted-by":"publisher","award":["31971151"],"award-info":[{"award-number":["31971151"]}],"id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012166","name":"National Key R&amp;D Program of China","doi-asserted-by":"publisher","award":["81627801"],"award-info":[{"award-number":["81627801"]}],"id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012166","name":"National Key R&amp;D Program of China","doi-asserted-by":"publisher","award":["81972909"],"award-info":[{"award-number":["81972909"]}],"id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012166","name":"National Key R&amp;D Program of China","doi-asserted-by":"publisher","award":["no. 2023M732238"],"award-info":[{"award-number":["no. 2023M732238"]}],"id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["2020YFA0908100"],"award-info":[{"award-number":["2020YFA0908100"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["31971151"],"award-info":[{"award-number":["31971151"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["81627801"],"award-info":[{"award-number":["81627801"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["81972909"],"award-info":[{"award-number":["81972909"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["no. 2023M732238"],"award-info":[{"award-number":["no. 2023M732238"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Education Foundation","award":["2020YFA0908100"],"award-info":[{"award-number":["2020YFA0908100"]}]},{"name":"Education Foundation","award":["31971151"],"award-info":[{"award-number":["31971151"]}]},{"name":"Education Foundation","award":["81627801"],"award-info":[{"award-number":["81627801"]}]},{"name":"Education Foundation","award":["81972909"],"award-info":[{"award-number":["81972909"]}]},{"name":"Education Foundation","award":["no. 2023M732238"],"award-info":[{"award-number":["no. 2023M732238"]}]},{"name":"China Postdoctoral Science Foundation","award":["2020YFA0908100"],"award-info":[{"award-number":["2020YFA0908100"]}]},{"name":"China Postdoctoral Science Foundation","award":["31971151"],"award-info":[{"award-number":["31971151"]}]},{"name":"China Postdoctoral Science Foundation","award":["81627801"],"award-info":[{"award-number":["81627801"]}]},{"name":"China Postdoctoral Science Foundation","award":["81972909"],"award-info":[{"award-number":["81972909"]}]},{"name":"China Postdoctoral Science Foundation","award":["no. 2023M732238"],"award-info":[{"award-number":["no. 2023M732238"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Centromeric chromatin is thought to play a critical role in ensuring the faithful segregation of chromosomes during mitosis. However, our understanding of this role is presently limited by our poor understanding of the structure and composition of this unique chromatin. The nucleosomal variant, CENP-A, localizes to narrow regions within the centromere, where it plays a major role in centromeric function, effectively serving as a platform on which the kinetochore is assembled. Previous work found that, within a given cell, the number of microtubules within kinetochores is essentially unchanged between CENP-A-localized regions of different physical sizes. However, it is unknown if the amount of CENP-A is also unchanged between these regions of different sizes, which would reflect a strict structural correspondence between these two key characteristics of the centromere\/kinetochore assembly. Here, we used super-resolution optical microscopy to image and quantify the amount of CENP-A and DNA within human centromere chromatin. We found that the amount of CENP-A within CENP-A domains of different physical sizes is indeed the same. Further, our measurements suggest that the ratio of CENP-A- to H3-containing nucleosomes within these domains is between 8:1 and 11:1. Thus, our results not only identify an unexpectedly strict relationship between CENP-A and microtubules stoichiometries but also that the CENP-A centromeric domain is almost exclusively composed of CENP-A nucleosomes.<\/jats:p>","DOI":"10.3390\/ijms242115871","type":"journal-article","created":{"date-parts":[[2023,11,1]],"date-time":"2023-11-01T07:24:04Z","timestamp":1698823444000},"page":"15871","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":3,"title":["Quantitative Super-Resolution Microscopy Reveals the Relationship between CENP-A Stoichiometry and Centromere Physical Size"],"prefix":"10.3390","volume":"24","author":[{"given":"Yaqian","family":"Li","sequence":"first","affiliation":[{"name":"State Key Laboratory of Systems Medicine for Cancer, School of Biomedical Engineering, Shanghai Jiao Tong University, Shanghai 200240, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1257-0727","authenticated-orcid":false,"given":"Jiabin","family":"Wang","sequence":"additional","affiliation":[{"name":"School of Chemistry and Chemical Engineering, Shanghai Jiao Tong University, Shanghai 200240, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xuecheng","family":"Chen","sequence":"additional","affiliation":[{"name":"Shanghai Center for Systems Biomedicine, Shanghai Jiao Tong University, Shanghai 200240, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2745-9546","authenticated-orcid":false,"given":"Daniel M.","family":"Czajkowsky","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Systems Medicine for Cancer, School of Biomedical Engineering, Shanghai Jiao Tong University, Shanghai 200240, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhifeng","family":"Shao","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Systems Medicine for Cancer, School of Biomedical Engineering, Shanghai Jiao Tong University, Shanghai 200240, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2023,11,1]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"1591","DOI":"10.1126\/science.270.5242.1591","article-title":"The centromere: Hub of chromosomal activities","volume":"270","author":"Pluta","year":"1995","journal-title":"Science"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"R562","DOI":"10.1016\/S0960-9822(99)80355-8","article-title":"Chromosome segregation: Dual control ensures fidelity","volume":"9","author":"Taylor","year":"1999","journal-title":"Curr. Biol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"7549","DOI":"10.1093\/nar\/15.18.7549","article-title":"Nucleotide sequence heterogeneity of alpha satellite repetitive DNA: A survey of alphoid sequences from different human chromosomes","volume":"15","author":"Waye","year":"1987","journal-title":"Nucleic Acids Res."},{"key":"ref_4","first-page":"524","article-title":"Chromosome-specific organization of human alpha satellite DNA","volume":"37","author":"Willard","year":"1985","journal-title":"Am. J. Hum. Genet."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"301","DOI":"10.1146\/annurev.genom.7.080505.115613","article-title":"Structural and functional dynamics of human centromeric chromatin","volume":"7","author":"Schueler","year":"2006","journal-title":"Annu. Rev. Genom. Hum. Genet."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1038\/nrm2310","article-title":"Molecular architecture of the kinetochore-microtubule interface","volume":"9","author":"Cheeseman","year":"2008","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"515","DOI":"10.1007\/978-3-319-58592-5_21","article-title":"Centromere Structure and Function","volume":"56","author":"Bloom","year":"2017","journal-title":"Prog. Mol. Subcell. Biol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1359","DOI":"10.1101\/gad.2055511","article-title":"Phosphorylation of H4 Ser 47 promotes HIRA-mediated nucleosome assembly","volume":"25","author":"Kang","year":"2011","journal-title":"Genes Dev."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"E28","DOI":"10.1038\/35050669","article-title":"The spindle: A dynamic assembly of microtubules and motors","volume":"3","author":"Wittmann","year":"2001","journal-title":"Nat. Cell Biol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1016\/S0074-7696(07)65003-7","article-title":"Mechanisms of mitotic spindle assembly and function","volume":"265","author":"Walczak","year":"2008","journal-title":"Int. Rev. Cytol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"eabl4178","DOI":"10.1126\/science.abl4178","article-title":"Complete genomic and epigenetic maps of human centromeres","volume":"376","author":"Altemose","year":"2022","journal-title":"Science"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"711","DOI":"10.1038\/s41592-022-01475-6","article-title":"DiMeLo-seq: A long-read, single-molecule method for mapping protein\u2013DNA interactions genome wide","volume":"19","author":"Altemose","year":"2022","journal-title":"Nat. Methods"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"687","DOI":"10.1038\/nsmb.2562","article-title":"The octamer is the major form of CENP-A nucleosomes at human centromeres","volume":"20","author":"Hasson","year":"2013","journal-title":"Nat. Struct. Mol. Biol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1038\/s41586-021-03420-7","article-title":"The structure, function and evolution of a complete human chromosome 8","volume":"593","author":"Logsdon","year":"2021","journal-title":"Nature"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1076","DOI":"10.1038\/nsmb845","article-title":"Centromeric chromatin exhibits a histone modification pattern that is distinct from both euchromatin and heterochromatin","volume":"11","author":"Sullivan","year":"2004","journal-title":"Nat. Struct. Mol. Biol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1016\/S0076-6879(03)75017-4","article-title":"Histone variant CENP-A purification, nucleosome reconstitution","volume":"375","author":"Yoda","year":"2004","journal-title":"Methods Enzym."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"789","DOI":"10.1083\/jcb.201503132","article-title":"A cell-free CENP-A assembly system defines the chromatin requirements for centromere maintenance","volume":"209","author":"Westhorpe","year":"2015","journal-title":"J. Cell Biol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"576","DOI":"10.1038\/s41467-019-08314-x","article-title":"The CENP-A centromere targeting domain facilitates H4K20 monomethylation in the nucleosome by structural polymorphism","volume":"10","author":"Arimura","year":"2019","journal-title":"Nat. Commun."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1023\/A:1009239013215","article-title":"Relevance of kinetochore size and microtubule-binding capacity for stable chromosome attachment during mitosis in PtK1 cells","volume":"6","author":"McEwen","year":"1998","journal-title":"Chromosome Res."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"e02137","DOI":"10.7554\/eLife.02137","article-title":"The quantitative architecture of centromeric chromatin","volume":"3","author":"Bodor","year":"2014","journal-title":"Elife"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"529","DOI":"10.1016\/j.tig.2004.08.008","article-title":"Analysis of the centromeric regions of the human genome assembly","volume":"20","author":"Rudd","year":"2004","journal-title":"Trends Genet."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"17981","DOI":"10.1039\/C9NR06364A","article-title":"Optical super-resolution microscopy unravels the molecular composition of functional protein complexes","volume":"11","author":"Dietz","year":"2019","journal-title":"Nanoscale"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"730","DOI":"10.1016\/j.tcb.2015.10.004","article-title":"Super-Resolution Microscopy: From Single Molecules to Supramolecular Assemblies","volume":"25","author":"Sydor","year":"2015","journal-title":"Trends Cell Biol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1016\/j.ccr.2018.08.006","article-title":"Organic fluorescent probes for stochastic optical reconstruction microscopy (STORM): Recent highlights and future possibilities","volume":"380","author":"Samanta","year":"2019","journal-title":"Coord. Chem. Rev."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"443001","DOI":"10.1088\/0022-3727\/48\/44\/443001","article-title":"The 2015 super-resolution microscopy roadmap","volume":"48","author":"Hell","year":"2015","journal-title":"J. Phys. D Appl. Phys."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1038\/nmeth.2843","article-title":"Precisely and accurately localizing single emitters in fluorescence microscopy","volume":"11","author":"Deschout","year":"2014","journal-title":"Nat. Methods"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"14072","DOI":"10.1038\/srep14072","article-title":"One, two or three? Probing the stoichiometry of membrane proteins by single-molecule localization microscopy","volume":"5","author":"Fricke","year":"2015","journal-title":"Sci. Rep."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1045","DOI":"10.1038\/s41592-019-0574-9","article-title":"Nuclear pores as versatile reference standards for quantitative superresolution microscopy","volume":"16","author":"Thevathasan","year":"2019","journal-title":"Nat. Methods"},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Lanzuolo, C., and Marasca, F. (2023). Polycomb Group Proteins: Methods and Protocols, Springer.","DOI":"10.1007\/978-1-0716-3143-0"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"4650","DOI":"10.1038\/ncomms5650","article-title":"Quantitative super-resolution imaging of Bruchpilot distinguishes active zone states","volume":"5","author":"Ehmann","year":"2014","journal-title":"Nat. Commun."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"301","DOI":"10.1146\/annurev-biophys-102521-112912","article-title":"Super-Resolution Microscopy for Structural Cell Biology","volume":"51","author":"Liu","year":"2022","journal-title":"Annu. Rev. Biophys."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"863610","DOI":"10.3389\/fchem.2022.863610","article-title":"3D Single Molecule Super-Resolution Microscopy of Whole Nuclear Lamina","volume":"10","author":"Rozario","year":"2022","journal-title":"Front. Chem."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"1523","DOI":"10.1091\/mbc.E20-11-0728","article-title":"Three-dimensional superresolution fluorescence microscopy maps the variable molecular architecture of the nuclear pore complex","volume":"32","author":"Sabinina","year":"2021","journal-title":"Mol. Biol. Cell"},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"Hoboth, P., \u0160ebesta, O., and Hoz\u00e1k, P. (2021). How Single-Molecule Localization Microscopy Expanded Our Mechanistic Understanding of RNA Polymerase II Transcription. Int. J. Mol. Sci., 22.","DOI":"10.20944\/preprints202106.0250.v1"},{"key":"ref_35","first-page":"253","article-title":"Immunofluorescence techniques applied to mitotic chromosome preparations","volume":"29","author":"Jeppesen","year":"1994","journal-title":"Methods Mol. Biol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1023\/A:1013168323754","article-title":"Immunolocalization of HP1 proteins in metaphasic mammalian chromosomes","volume":"23","author":"Minc","year":"2001","journal-title":"Methods Cell Sci."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"8030","DOI":"10.1021\/acsnano.2c01025","article-title":"Single-Molecule Micromanipulation and Super-Resolution Imaging Resolve Nanodomains Underlying Chromatin Folding in Mitotic Chromosomes","volume":"16","author":"Wang","year":"2022","journal-title":"ACS Nano"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"7276","DOI":"10.1021\/acs.chemrev.6b00729","article-title":"Unraveling the Thousand Word Picture: An Introduction to Super-Resolution Data Analysis","volume":"117","author":"Lee","year":"2017","journal-title":"Chem. Rev."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Annibale, P., Vanni, S., Scarselli, M., Rothlisberger, U., and Radenovic, A. (2011). Quantitative photo activated localization microscopy: Unraveling the effects of photoblinking. PLoS ONE, 6.","DOI":"10.1371\/journal.pone.0022678"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1027","DOI":"10.1038\/nmeth.1768","article-title":"Evaluation of fluorophores for optimal performance in localization-based super-resolution imaging","volume":"8","author":"Dempsey","year":"2011","journal-title":"Nat. Methods"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1007\/s00418-014-1195-0","article-title":"Quantitative single-molecule localization microscopy combined with rule-based modeling reveals ligand-induced TNF-R1 reorganization toward higher-order oligomers","volume":"142","author":"Fricke","year":"2014","journal-title":"Histochem. Cell Biol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"3595","DOI":"10.1016\/j.febslet.2014.06.014","article-title":"Challenges in quantitative single molecule localization microscopy","volume":"588","author":"Shivanandan","year":"2014","journal-title":"FEBS Lett."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"047001","DOI":"10.1088\/2050-6120\/acf546","article-title":"Quasi-Equilibrium State Based Quantification of Biological Macromolecules in Single-Molecule Localization Microscopy","volume":"11","author":"Xuecheng","year":"2023","journal-title":"Methods Appl. Fluoresc."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"e53807","DOI":"10.7554\/eLife.53807","article-title":"Microneedle manipulation of the mammalian spindle reveals specialized, short-lived reinforcement near chromosomes","volume":"9","author":"Suresh","year":"2020","journal-title":"Elife"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"4909","DOI":"10.1021\/acsnano.8b01963","article-title":"Super-resolution Imaging of Individual Human Subchromosomal Regions in Situ Reveals Nanoscopic Building Blocks of Higher-Order Structure","volume":"12","author":"Fang","year":"2018","journal-title":"ACS Nano"},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Hao, X., Parmar, J.J., Lelandais, B., Aristov, A., Ouyang, W., Weber, C., and Zimmer, C. (2021). Super-resolution visualization and modeling of human chromosomal regions reveals cohesin-dependent loop structures. Genome Biol., 22.","DOI":"10.1186\/s13059-021-02343-w"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1093\/nar\/gkab1215","article-title":"Super resolution microscopy reveals how elongating RNA polymerase II and nascent RNA interact with nucleosome clutches","volume":"50","author":"Vicario","year":"2022","journal-title":"Nucleic Acids Res."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"eaba8811","DOI":"10.1126\/sciadv.aba8811","article-title":"Chromatin arranges in chains of mesoscale domains with nanoscale functional topography independent of cohesin","volume":"6","author":"Miron","year":"2020","journal-title":"Sci. Adv."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"1259","DOI":"10.1074\/jbc.M109.067330","article-title":"Dynamic and selective nucleosome repositioning during endotoxin tolerance","volume":"285","author":"Liu","year":"2010","journal-title":"J. Biol. Chem."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"844","DOI":"10.1126\/science.abn3810","article-title":"Structure of the human inner kinetochore bound to a centromeric CENP-A nucleosome","volume":"376","author":"Yatskevich","year":"2022","journal-title":"Science"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1038\/38444","article-title":"Crystal structure of the nucleosome core particle at 2.8\u2009\u00c5 resolution","volume":"389","author":"Luger","year":"1997","journal-title":"Nature"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"8075","DOI":"10.1128\/MCB.01235-06","article-title":"Differential regulation of short- and long-tract gene conversion between sister chromatids by Rad51C","volume":"26","author":"Nagaraju","year":"2006","journal-title":"Mol. Cell Biol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"1018","DOI":"10.1016\/j.cub.2011.05.032","article-title":"Cohesion Fatigue Induces Chromatid Separation in Cells Delayed at Metaphase","volume":"21","author":"Daum","year":"2011","journal-title":"Curr. Biol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1083\/jcb.200210005","article-title":"Human centromere chromatin protein hMis12, essential for equal segregation, is independent of CENP-A loading pathway","volume":"160","author":"Goshima","year":"2003","journal-title":"J. Cell Biol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"10036","DOI":"10.1038\/ncomms10036","article-title":"Stable kinetochore\u2013microtubule attachment is sufficient to silence the spindle assembly checkpoint in human cells","volume":"6","author":"Tauchman","year":"2015","journal-title":"Nat. Commun."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"946","DOI":"10.1016\/j.semcdb.2011.09.016","article-title":"The CCAN complex: Linking centromere specification to control of kinetochore\u2013microtubule dynamics","volume":"22","author":"McAinsh","year":"2011","journal-title":"Semin. Cell Dev. Biol."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"278","DOI":"10.1038\/s41586-019-1609-1","article-title":"Structure of the inner kinetochore CCAN complex assembled onto a centromeric nucleosome","volume":"574","author":"Yan","year":"2019","journal-title":"Nature"},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Cheeseman, I.M. (2014). The kinetochore. Cold Spring Harb Perspect Biol., 6.","DOI":"10.1101\/cshperspect.a015826"},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Musacchio, A., and Desai, A. (2017). A Molecular View of Kinetochore Assembly and Function. Biology, 6.","DOI":"10.3390\/biology6010005"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"279","DOI":"10.1146\/annurev-genet-072820-034559","article-title":"The Four Causes: The Functional Architecture of Centromeres and Kinetochores","volume":"56","author":"McAinsh","year":"2022","journal-title":"Annu Rev Genet"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"344","DOI":"10.1038\/s41586-023-06457-y","article-title":"The complete sequence of a human Y chromosome","volume":"621","author":"Rhie","year":"2023","journal-title":"Nature"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1016\/S1534-5807(02)00135-1","article-title":"Conserved organization of centromeric chromatin in flies and humans","volume":"2","author":"Blower","year":"2002","journal-title":"Dev. Cell"},{"key":"ref_63","doi-asserted-by":"crossref","unstructured":"Chen, G., and Deng, X. (2018). Cell Synchronization by Double Thymidine Block. Bio. Protoc., 8.","DOI":"10.21769\/BioProtoc.2994"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"2389","DOI":"10.1093\/bioinformatics\/btu202","article-title":"ThunderSTORM: A comprehensive ImageJ plug-in for PALM and STORM data analysis and super-resolution imaging","volume":"30","author":"Borkovec","year":"2014","journal-title":"Bioinformatics"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"101372","DOI":"10.1016\/j.mex.2021.101372","article-title":"Dual-color dSTORM imaging and ThunderSTORM image reconstruction and analysis to study the spatial organization of the nuclear phosphatidylinositol phosphates","volume":"8","author":"Hoboth","year":"2021","journal-title":"MethodsX"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1016\/j.jsb.2014.10.001","article-title":"Separation of replication and transcription domains in nucleoli","volume":"188","author":"Smirnov","year":"2014","journal-title":"J. Struct. Biol."}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/24\/21\/15871\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T21:15:38Z","timestamp":1760130938000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/24\/21\/15871"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,11,1]]},"references-count":66,"journal-issue":{"issue":"21","published-online":{"date-parts":[[2023,11]]}},"alternative-id":["ijms242115871"],"URL":"https:\/\/doi.org\/10.3390\/ijms242115871","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,11,1]]}}}