Genetic and epigenetic regulation of centromeres: a look at HAC formation

Chromosome Research

Volumn 23, Issue 1, 2015, Pages 87-103

  Ohzeki, Jun ichirou ^a   Larionov, Vladimir ^b   Earnshaw, William C ^c   Masumoto, Hiroshi ^a  

^a KAZUSA DNA RESEARCH INSTITUTE   (Japan)

^b NATIONAL CANCER INSTITUTE   (United States)

^c UNIVERSITY OF EDINBURGH   (United Kingdom)

Author keywords

CENP A; CENP B; Centromere; HAC; HAT; Heterochromatin; Kinetochore

Indexed keywords

CENTROMERE PROTEIN A; CENTROMERE PROTEIN B; DNA FRAGMENT; GENOMIC DNA; PLASMID DNA; REPETITIVE DNA; SATELLITE DNA; AUTOANTIGEN; NONHISTONE PROTEIN;

ARTICLE; CENTROMERE; CHROMATIN; CHROMATIN ASSEMBLY AND DISASSEMBLY; DNA SEQUENCE; EPIGENETICS; GENE EXPRESSION REGULATION; HELA CELL LINE; HETEROCHROMATIN; HUMAN; HUMAN ARTIFICIAL CHROMOSOME; NONHUMAN; PRIORITY JOURNAL; SACCHAROMYCES CEREVISIAE; SCHIZOSACCHAROMYCES POMBE; TWO HYBRID SYSTEM; U2OS CELL LINE; ACETYLATION; CELL LINE; DNA MICROARRAY; GENETIC EPIGENESIS; GENETICS; METABOLISM; YEAST;

ACETYLATION; AUTOANTIGENS; CELL LINE; CENTROMERE; CHROMOSOMAL PROTEINS, NON-HISTONE; CHROMOSOMES, ARTIFICIAL, HUMAN; EPIGENESIS, GENETIC; HUMANS; OLIGONUCLEOTIDE ARRAY SEQUENCE ANALYSIS; YEASTS;

EID: 84925544981     PISSN: 09673849     EISSN: 15736849     Source Type: Journal    
DOI: 10.1007/s10577-015-9470-z     Document Type: Article

References (133)

1. Aldrup-Macdonald ME, Sullivan BA (2014) The past, present, and future of human centromere genomics. Genes 5(1):33–50
2. Allshire RC, Karpen GH (2008) Epigenetic regulation of centromeric chromatin: old dogs, new tricks? Nat Rev Genet 9(12):923–937
3. Alonso A, Fritz B, Hasson D, Abrusan G, Cheung F, Yoda K, Radlwimmer B, Ladurner AG, Warburton PE (2007) Co-localization of CENP-C and CENP-H to discontinuous domains of CENP-A chromatin at human neocentromeres. Genome Biol 8(7):R148
4. Ando S, Yang H, Nozaki N, Okazaki T, Yoda K (2002) CENP-A, -B, and -C chromatincomplex that contains the I-type alpha-satellite array constitutes the prekinetochore in HeLa cells. Mol Cell Biol 22(7):2229–2241
5. Barnhart MC, Kuich PH, Stellfox ME, Ward JA, Bassett EA, Black BE, Foltz DR (2011) HJURP is a CENP-A chromatin assembly factor sufficient to form a functional de novo kinetochore. J Cell Biol 194(2):229–243
6. Basu J, Stromberg G, Compitello G, Willard HF, Van Bokkelen G (2005) Rapid creation of BAC-based human artificial chromosome vectors by transposition with synthetic alpha-satellite arrays. Nucleic Acids Res 33(2):587–596
7. Bergmann JH, Rodríguez MG, Martins NM, Kimura H, Kelly DA, Masumoto H, Larionov V, Jansen LE, Earnshaw WC (2011) Epigenetic engineering shows H3K4me2 is required for HJURP targeting and CENP-A assembly on a synthetic human kinetochore. EMBO J 30(2):328–340
8. Bergmann JH, Jakubsche JN, Martins NM, Kagansky A, Nakano M, Kimura H, Kelly DA, Turner BM, Masumoto H, Larionov V, Earnshaw WC (2012a) Epigenetic engineering: histone H3K9 acetylation is compatible with kinetochore structure and function. J Cell Sci 125(Pt 2):411–421
9. Bergmann JH, Martins NM, Larionov V, Masumoto H, Earnshaw WC (2012b) HACking the centromere chromatin code: insights from human artificial chromosomes. Chromosome Res 20(5):505–519
10. Bernard P, Maure JF, Partridge JF, Genier S, Javerzat JP, Allshire (2001) Requirement of heterochromatin for cohesion at centromeres. Science 294(5551):2539–2542
11. Black BE, Cleveland DW (2011) Epigenetic centromere propagation and the nature of CENP-a nucleosomes. Cell 144(4):471–479
12. Brown KE, Barnett MA, Burgtorf C, Shaw P, Buckle VJ, Brown WR (1994) Dissecting the centromere of the human Y chromosome with cloned telomeric DNA. Hum Mol Genet 3(8):1227–1237
13. Cam HP, Noma K, Ebina H, Levin HL, Grewal SI (2008) Host genome surveillance for retrotransposons by transposon-derived proteins. Nature 451(7177):431–436
14. Camahort R, Shivaraju M, Mattingly M, Li B, Nakanishi S, Zhu D, Shilatifard A, Workman JL, Gerton JL (2009) Cse4 is part of an octameric nucleosome in budding yeast. Mol Cell 35(6):794–805
15. Cardinale S, Bergmann JH, Kelly D, Nakano M, Valdivia MM, Kimura H, Masumoto H, Larionov V, Earnshaw WC (2009) Hierarchical inactivation of a synthetic human kinetochore by a chromatin modifier. Mol Biol Cell 20(19):4194–4204
16. Carroll CW, Milks KJ, Straight AF (2010) Dual recognition of CENP-A nucleosomes is required for centromere assembly. J Cell Biol 189(7):1143–1155
17. Chan FL, Wong LH (2012) Transcription in the maintenance of centromere chromatin identity. Nucleic Acids Res 40(22):11178–11188
18. Cheeseman IM, Chappie JS, Wilson-Kubalek EM, Desai A (2006) The conserved KMN network constitutes the core microtubule-binding site of the kinetochore. Cell 127:983–997
19. Chikashige Y, Kinoshita N, Nakaseko Y, Matsumoto T, Murakami S, Niwa O, Yanagida M (1989) Composite motifs and repeat symmetry in S. pombe centromeres: direct analysis by integration of NotI restriction sites. Cell 57(5):739–751
20. Choi ES, Strålfors A, Catania S, Castillo AG, Svensson JP, Pidoux AL, Ekwall K, Allshire RC (2012) Factors that promote H3 chromatin integrity during transcription prevent promiscuous deposition of CENP-A (Cnp1) in fission yeast. PLoS Genet 8(9):e1002985
21. Choo KH, Vissel B, Nagy A, Earle E, Kalitsis P (1991) A survey of the genomic distribution of alpha satellite DNA on all the human chromosomes, and derivation of a new consensus sequence. Nucleic Acids Res 19(6):1179–1182
22. Clarke L, Carbon J (1980) Isolation of a yeast centromere and construction of functional small circular chromosomes. Nature 287:504–509
23. Cleveland DW, Mao Y, Sullivan KF (2003) Centromeres and kinetochores: from epigenetics to mitotic checkpoint signaling. Cell 112(4):407–421
24. Craig JM, Earle E, Canham P, Wong LH, Anderson M, Choo KH (2003) Analysis of mammalian proteins involved in chromatin modification reveals new metaphase centromeric proteins and distinct chromosomal distribution patterns. Hum Mol Genet 12(23):3109–3121
25. Dambacher S, Deng W, Hahn M, Sadic D, Fröhlich J, Nuber A, Hoischen C, Diekmann S, Leonhardt H, Schotta G (2012) CENP-C facilitates the recruitment of M18BP1 to centromeric chromatin. Nucleus 3(1):101–110
26. Deyter GM, Biggins S (2014) The FACT complex interacts with the E3 ubiquitin ligase Psh1 to prevent ectopic localization of CENP-A. Genes Dev 28(16):1815–1826
27. Distèche C, Hagemeijer A, Frederic J, Progneaux D (1972) An abnormal large human chromosome identified as an end-to-end fusion of two X’s by combined results of the new banding techniques and microdensitometry. Clin Genet 3(5):388–395
28. du Sart D, Cancilla MR, Earle E, Mao JI, Saffery R, Tainton KM, Kalitsis P, Martyn J, Barry AE, Choo KH (1997) A functional neo-centromere formed through activation of a latent human centromere and consisting of non-alpha-satellite DNA. Nat Genet 16(2):144–153
29. Dunleavy EM, Roche D, Tagami H, Lacoste N, Ray-Gallet D, Nakamura Y, Daigo Y, Nakatani Y, Almouzni-Pettinotti G (2009) HJURP is a cell-cycle-dependent maintenance and deposition factor of CENP-A at centromeres. Cell 137(3):485–497
30. Earnshaw WC, Migeon BR (1985) Three related centromere proteins are absent from the inactive centromere of a stable isodicentric chromosome. Chromosoma 92(4):290–296
31. Earnshaw WC, Rothfield N (1985) Identification of a family of human centromere proteins using autoimmune sera from patients with scleroderma. Chromosoma 91(3–4):313–321
32. Earnshaw WC, Sullivan KF, Machlin PS, Cooke CA, Kaiser DA, Pollard TD, Rothfield NF, Cleveland DW (1987) Molecular cloning of cDNA for CENP-B, the major human centromere autoantigen. J Cell Biol 104(4):817–829
33. Earnshaw WC, Ratrie H 3rd, Stetten G (1989) Visualization of centromere proteins CENP-B and CENP-C on a stable dicentric chromosome in cytological spreads. Chromosoma 98(1):1–12
34. Earnshaw WC, Bernat RL, Cooke CA, Rothfield NF (1991) Role of the centromere/kinetochore in cell cycle control. Cold Spring Harb Symp Quant Biol 56:675–685
35. Earnshaw WC, Allshire RC, Black BE, Bloom K, Brinkley BR, Brown W, Cheeseman IM, Choo KH, Copenhaver GP, Deluca JG, Desai A, Diekmann S, Erhardt S, Fitzgerald-Hayes M, Foltz D, Fukagawa T, Gassmann R, Gerlich DW, Glover DM, Gorbsky GJ, Harrison SC, Heun P, Hirota T, Jansen LE, Karpen G, Kops GJ, Lampson MA, Lens SM, Losada A, Luger K, Maiato H, Maddox PS, Margolis RL, Masumoto H, McAinsh AD, Mellone BG, Meraldi P, Musacchio A, Oegema K, O’Neill RJ, Salmon ED, Scott KC, Straight AF, Stukenberg PT, Sullivan BA, Sullivan KF, Sunkel CE, Swedlow JR, Walczak CE, Warburton PE, Westermann S, Willard HF, Wordeman L, Yanagida M, Yen TJ, Yoda K, Cleveland DW (2013) Esperanto for histones: CENP-A, not CenH3, is the centromeric histone H3 variant. Chromosome Res 21(2):101–106
36. Ebersole TA, Ross A, Clark E, McGill N, Schindelhauer D, Cooke H, Grimes B (2000) Mammalian artificial chromosome formation from circular alphoid input DNA does not require telomere repeats. Hum Mol Genet 9(11):1623–1631
37. Ebersole T, Kim JH, Samoshkin A, Kouprina N, Pavlicek A, White RJ, Larionov V (2011) tRNA genes protect a reporter gene from epigenetic silencing in mouse cells. Cell Cycle 10(16):2779–2791
38. Erliandri I, Fu H, Nakano M, Kim JH, Miga KH, Liskovykh M, Earnshaw WC, Masumoto H, Kouprina N, Aladjem MI, Larionov V (2014) Replication of alpha-satellite DNA arrays in endogenous human centromeric regions and in human artificial chromosome. Nucleic Acids Res
39. Fachinetti D, Folco HD, Nechemia-Arbely Y, Valente LP, Nguyen K, Wong AJ, Zhu Q, Holland AJ, Desai A, Jansen LE, Cleveland DW (2013) A two-step mechanism for epigenetic specification of centromere identity and function. Nat Cell Biol 15(9):1056–1066
40. Farr CJ, Bayne RA, Kipling D, Mills W, Critcher R, Cooke HJ (1995) Generation of a human X-derived minichromosome using telomere-associated chromosome fragmentation. EMBO J 14(21):5444–5454
41. Fitzgerald-Hayes M, Clarke L, Carbon J (1982) Nucleotide sequence comparisons and functional analysis of yeast centromere DNAs. Cell 29(1):235–244
42. Folco HD, Pidoux AL, Urano T, Allshire RC (2008) Heterochromatin and RNAi are required to establish CENP-A chromatin at centromeres. Science 319(5859):94–97
43. Foltz DR, Jansen LE, Black BE, Bailey AO, Yates JR 3rd, Cleveland DW (2006) The human CENP-A centromeric nucleosome-associated complex. Nat Cell Biol 8(5):458–469
44. Foltz DR, Jansen LE, Bailey AO, Yates JR 3rd, Bassett EA, Wood S, Black BE, Cleveland DW (2009) Centromere-specific assembly of CENP-a nucleosomes is mediated by HJURP. Cell 137(3):472–484
45. Fujita Y, Hayashi T, Kiyomitsu T, Toyoda Y, Kokubu A, Obuse C, Yanagida M (2007) Priming of centromere for CENP-A recruitment by human hMis18alpha, hMis18beta, and M18BP1. Dev Cell 12(1):17–30
46. Fukagawa T, Earnshaw WC (2014) The centromere: chromatin foundation for the kinetochore machinery. Dev Cell 30(5):496–508
47. Furuyama S, Biggins S (2007) Centromere identity is specified by a single centromeric nucleosome in budding yeast. Proc Natl Acad Sci U S A 104(37):14706–14711
48. Gascoigne KE, Takeuchi K, Suzuki A, Hori T, Fukagawa T, Cheeseman IM (2011) Induced ectopic kinetochore assembly bypasses the requirement for CENP-A nucleosomes. Cell 145(3):410–422
49. Grewal SI, Jia S (2007) Heterochromatin revisited. Nat Rev Genet 8(1):35–46
50. Grimes BR, Rhoades AA, Willard HF (2002) Alpha-satellite DNA and vector composition influence rates of human artificial chromosome formation. Mol Ther 5(6):798–805
51. Grimes BR, Babcock J, Rudd MK, Chadwick B, Willard HF (2004) Assembly and characterization of heterochromatin and euchromatin on human artificial chromosomes. Genome Biol 5(11):R89
52. Hahnenberger KM, Baum MP, Polizzi CM, Carbon J, Clarke L (1989) Construction of functional artificial minichromosomes in the fission yeast Schizosaccharomyces pombe. Proc Natl Acad Sci U S A 86(2):577–581
53. Harrington JJ, Van Bokkelen G, Mays RW, Gustashaw K, Willard HF (1997) Formation of de novo centromeres and construction of first-generation human artificial microchromosomes. Nat Genet 15(4):345–355
54. Hasegawa Y, Ishikura T, Hasegawa T, Watanabe T, Suzuki J, Nakayama M, Okamura Y, Okazaki T, Koseki H, Ohara O, Ikeno M, Masumoto H (2014) Generating a transgenic mouse line stably expressing human MHC surface antigen from a HAC carrying multiple genomic BACs. Chromosoma. doi:10.1007/s00412-014-0488-3
55. Hegemann JH, Shero JH, Cottarel G, Philippsen P, Hieter P (1988) Mutational analysis of centromere DNA from chromosome VI of Saccharomyces cerevisiae. Mol Cell Biol 8(6):2523–2535
56. Helfricht A, Wiegant WW, Thijssen PE, Vertegaal AC, Luijsterburg MS, van Attikum H (2013) Remodeling and spacing factor 1 (RSF1) deposits centromere proteins at DNA double-strand breaks to promote non-homologous end-joining. Cell Cycle 12(18):3070–3082
57. Hiratsuka M, Uno N, Ueda K, Kurosaki H, Imaoka N, Kazuki K, Ueno E, Akakura Y, Katoh M, Osaki M, Kazuki Y, Nakagawa M, Yamanaka S, Oshimura M (2011) Integration-free iPS cells engineered using human artificial chromosome vectors. PLoS One 6(10):e25961
58. Hori T, Shang WH, Takeuchi K, Fukagawa T (2013) The CCAN recruits CENP-A to the centromere and forms the structural core for kinetochore assembly. J Cell Biol 200(1):45–60
59. Hori T, Shang WH, Toyoda A, Misu S, Monma N, Ikeo K, Molina O, Vargiu G, Fujiyama A, Kimura H, Earnshaw WC, Fukagawa T (2014) Histone H4 Lys 20 monomethylation of the CENP-A nucleosome is essential for kinetochore assembly. Dev Cell 29(6):740–749
60. Howman EV, Fowler KJ, Newson AJ, Redward S, MacDonald AC, Kalitsis P, Choo KH (2000) Early disruption of centromeric chromatin organization in centromere protein A (Cenpa) null mice. Proc Natl Acad Sci U S A 97(3):1148–1153
61. Hudson DF, Fowler KJ, Earle E, Saffery R, Kalitsis P, Trowell H, Hill J, Wreford NG, de Kretser DM, Cancilla MR, Howman E, Hii L, Cutts SM, Irvine DV, Choo KH (1998) Centromere protein B null mice are mitotically and meiotically normal but have lower body and testis weights. J Cell Biol 141(2):309–319
62. Ikeno M, Masumoto H, Okazaki T (1994) Distribution of CENP-B boxes reflected in CREST centromere antigenic sites on long-range alpha-satellite DNA arrays of human chromosome 21. Hum Mol Genet 3(8):1245–1257
63. Ikeno M, Grimes B, Okazaki T, Nakano M, Saitoh K, Hoshino H, McGill NI, Cooke H, Masumoto H (1998) Construction of YAC-based mammalian artificial chromosomes. Nat Biotechnol 16(5):431–439
64. Ikeno M, Suzuki N, Hasegawa Y, Okazaki T (2009) Manipulating transgenes using a chromosome vector. Nucleic Acids Res 37:e44
65. Ishii K, Ogiyama Y, Chikashige Y, Soejima S, Masuda F, Kakuma T, Hiraoka Y, Takahashi K (2008) Heterochromatin integrity affects chromosome reorganization after centromere dysfunction. Science 321(5892):1088–1091
66. Izuta H, Ikeno M, Suzuki N, Tomonaga T, Nozaki N, Obuse C, Kisu Y, Goshima N, Nomura F, Nomura N, Yoda K (2006) Comprehensive analysis of the ICEN (Interphase Centromere Complex) components enriched in the CENP-A chromatin of human cells. Genes Cells 11(6):673–684
67. Jansen LE, Black BE, Foltz DR, Cleveland DW (2007) Propagation of centromeric chromatin requires exit from mitosis. J Cell Biol 176(6):795–805
68. Kagansky A, Folco HD, Almeida R, Pidoux AL, Boukaba A, Simmer F, Urano T, Hamilton GL, Allshire RC (2009) Synthetic heterochromatin bypasses RNAi and centromeric repeats to establish functional centromeres. Science 324(5935):1716–1719
69. Kapoor M, de Montes de Oca Luna R, Liu G, Lozano G, Cummings C, Mancini M, Ouspenski I, Brinkley BR, May GS (1998) The CENPB gene is not essential in mice. Chromosoma 107(8):570–576
70. Kim JH, Ebersole T, Kouprina N, Noskov VN, Ohzeki J, Masumoto H, Mravinac B, Sullivan BA, Pavlicek A, Dovat S, Pack SD, Kwon YW, Flanagan PT, Loukinov D, Lobanenkov V, Larionov V (2009) Human gamma-satellite DNA maintains open chromatin structure and protects a transgene from epigenetic silencing. Genome Res 19(4):533–544
71. Kim JH, Kononenko A, Erliandri I, Kim TA, Nakano M, Iida Y, Barrett JC, Oshimura M, Masumoto H, Earnshaw WC, Larionov V, Kouprina N (2011) Human artificial chromosome (HAC) vector with a conditional centromere for correction of genetic deficiencies in human cells. Proc Natl Acad Sci U S A 108(50):20048–20053
72. Kipling D, Mitchell AR, Masumoto H, Wilson HE, Nicol L, Cooke HJ (1995) CENP-B binds a novel centromeric sequence in the Asian mouse Mus caroli. Mol Cell Biol 15(8):4009–4020
73. Kitagawa K, Hieter P (2001) Evolutionary conservation between budding yeast and human kinetochores. Nat Rev Mol Cell Biol 2(9):678–687
74. Kitagawa K, Masumoto H, Ikeda M, Okazaki T (1995) Analysis of protein-DNA and protein-protein interactions of centromere protein B (CENP-B) and properties of the DNA-CENP-B complex in the cell cycle. Mol Cell Biol 15(3):1602–1612
75. Kononenko AV, Bansal R, Lee NC, Grimes BR, Masumoto H, Earnshaw WC, Larionov V, Kouprina N (2014) A portable BRCA1-HAC (human artificial chromosome) module for analysis of BRCA1 tumor suppressor function. Nucleic Acids Res
76. Maehara K, Takahashi K, Saitoh S (2010) CENP-A reduction induces a p53-dependent cellular senescence response to protect cells from executing defective mitoses. Mol Cell Biol 30(9):2090–2104
77. Masumoto H, Masukata H, Muro Y, Nozaki N, Okazaki T (1989) A human centromere antigen (CENP-B) interacts with a short specific sequence in alphoid DNA, a human centromeric satellite. J Cell Biol 109(5):1963–1973
78. Masumoto H, Ikeno M, Nakano M, Okazaki T, Grimes B, Cooke H, Suzuki N (1998) Assay of centromere function using a human artificial chromosome. Chromosoma 107(6–7):406–416
79. Masumoto H, Nakano M, Ohzeki J (2004) The role of CENP-B and alpha-satellite DNA: de novo assembly and epigenetic maintenance of human centromeres. Chromosome Res 12(6):543–556
80. McClintock B (1941) The stability of broken ends of chromosomes in Zea mays. Genetics 26(2):234–282
81. Mejía JE, Willmott A, Levy E, Earnshaw WC, Larin Z (2001) Functional complementation of a genetic deficiency with human artificial chromosomes. Am J Hum Genet 69(2):315–326
82. Merry DE, Pathak S, Hsu TC, Brinkley BR (1985) Anti-kinetochore antibodies: use as probes for inactive centromeres. Am J Hum Genet 37(2):425–430
83. Miga KH, Newton Y, Jain M, Altemose N, Willard HF, Kent WJ (2014) Centromere reference models for human chromosomes X and Y satellite arrays. Genome Res 24(4):697–707
84. Mills W, Critcher R, Lee C, Farr CJ (1999) Generation of an approximately 2.4 Mb human X centromere-based minichromosome by targeted telomere-associated chromosome fragmentation in DT40. Hum Mol Genet 8(5):751–761
85. Moree B, Meyer CB, Fuller CJ, Straight AF (2011) CENP-C recruits M18BP1 to centromeres to promote CENP-A chromatin assembly. J Cell Biol 194(6):855–871
86. Moroi Y, Peebles C, Fritzler MJ, Steigerwald J, Tan EM (1980) Autoantibody to centromere (kinetochore) in scleroderma sera. Proc Natl Acad Sci U S A 77(3):1627–1631
87. Muro Y, Masumoto H, Yoda K, Nozaki N, Ohashi M, Okazaki T (1992) Centromere protein B assembles human centromeric alpha-satellite DNA at the 17-bp sequence, CENP-B box. J Cell Biol 116(3):585–596
88. Murray AW, Szostak JW (1983) Construction of artificial chromosomes in yeast. Nature 305(5931):189–193
89. Musacchio A, Salmon ED (2007) The spindle-assembly checkpoint in space and time. Nat Rev Mol Cell Biol 8(5):379–393
90. Nakagawa H, Lee JK, Hurwitz J, Allshire RC, Nakayama J, Grewal SI, Tanaka K, Murakami Y (2002) Fission yeast CENP-B homologs nucleate centromeric heterochromatin by promoting heterochromatin-specific histone tail modifications. Genes Dev 16(14):1766–1778
91. Nakano M, Okamoto Y, Ohzeki J, Masumoto H (2003) Epigenetic assembly of centromeric chromatin at ectopic alpha-satellite sites on human chromosomes. J Cell Sci 116:4021–4034
92. Nakano M, Cardinale S, Noskov VN, Gassmann R, Vagnarelli P, Kandels-Lewis S, Larionov V, Earnshaw WC, Masumoto H (2008) Inactivation of a human kinetochore by specific targeting of chromatin modifiers. Dev Cell 14(4):507–522
93. Nakaseko Y, Adachi Y, Funahashi S, Niwa O, Yanagida M (1986) Chromosome walking shows a highly homologous repetitive sequence present in all the centromere regions of fission yeast. EMBO J 5(5):1011–1021
94. Nakashima H, Nakano M, Ohnishi R, Hiraoka Y, Kaneda Y, Sugino A, Masumoto H (2005) Assembly of additional heterochromatin distinct from centromere-kinetochore chromatin is required for de novo formation of human artificial chromosome. J Cell Sci 118(Pt 24):5885–5898
95. Nonaka N, Kitajima T, Yokobayashi S, Xiao G, Yamamoto M, Grewal SI, Watanabe Y (2002) Recruitment of cohesin to heterochromatic regions by Swi6/HP1 in fission yeast. Nat Cell Biol 4(1):89–93
96. Obuse C, Yang H, Nozaki N, Goto S, Okazaki T, Yoda K (2004) Proteomics analysis of the centromere complex from HeLa interphase cells: UV-damaged DNA binding protein 1 (DDB-1) is a component of the CEN-complex, while BMI-1 is transiently co-localized with the centromeric region in interphase. Genes Cells 9(2):105–120
97. Oegema K, Desai A, Rybina S, Kirkham M, Hyman AA (2001) Functional analysis of kinetochore assembly in Caenorhabditis elegans. J Cell Biol 153(6):1209–1226
98. Ogiyama Y, Ishii K (2012) The smooth and stable operation of centromeres. Genes Genet Syst 87(2):63–73
99. Ohta S, Bukowski-Wills JC, Sanchez-Pulido L, Alves Fde L, Wood L, Chen ZA, Platani M, Fischer L, Hudson DF, Ponting CP, Fukagawa T, Earnshaw WC, Rappsilber J (2010) The protein composition of mitotic chromosomes determined using multiclassifier combinatorial proteomics. Cell 142(5):810–821
100. Ohzeki J, Nakano M, Okada T, Masumoto H (2002) CENP-B box is required for de novo centromere chromatin assembly on human alphoid DNA. J Cell Biol 159(5):765–775
101. Ohzeki J, Bergmann JH, Kouprina N, Noskov VN, Nakano M, Kimura H, Earnshaw WC, Larionov V, Masumoto H (2012) Breaking the HAC Barrier: histone H3K9 acetyl/methyl balance regulates CENP-A assembly. EMBO J 31(10):2391–2402
102. Okada M, Cheeseman IM, Hori T, Okawa K, McLeod IX, Yates JR 3rd, Desai A, Fukagawa T (2006) The CENP-H-I complex is required for the efficient incorporation of newly synthesized CENP-A into centromeres. Nat Cell Biol 8(5):446–457
103. Okada T, Ohzeki J, Nakano M, Yoda K, Brinkley WR, Larionov V, Masumoto H (2007) CENP-B controls centromere formation depending on the chromatin context. Cell 131(7):1287–1300
104. Okamoto Y, Nakano M, Ohzeki J, Larionov V, Masumoto H (2007) A minimal CENP-A core is required for nucleation and maintenance of a functional human centromere. EMBO J 26(5):1279–1291
105. Palmer DK, O’Day K, Wener MH, Andrews BS, Margolis RL (1987) A 17-kD centromere protein (CENP-A) copurifies with nucleosome core particles and with histones. J Cell Biol 104(4):805–815
106. Perez-Castro AV, Shamanski FL, Meneses JJ, Lovato TL, Vogel KG, Moyzis RK, Pedersen R (1998) Centromeric protein B null mice are viable with no apparent abnormalities. Dev Biol 201(2):135–143
107. Perpelescu M, Nozaki N, Obuse C, Yang H, Yoda K (2009) Active establishment of centromeric CENP-A chromatin by RSF complex. J Cell Biol 185(3):397–407
108. Pertile MD, Graham AN, Choo KH, Kalitsis P (2009) Rapid evolution of mouse Y centromere repeat DNA belies recent sequence stability. Genome Res 19(12):2202–2213
109. Peters AH, O’Carroll D, Scherthan H, Mechtler K, Sauer S, Schöfer C, Weipoltshammer K, Pagani M, Lachner M, Kohlmaier A, Opravil S, Doyle M, Sibilia M, Jenuwein T (2001) Loss of the Suv39h histone methyltransferases impairs mammalian heterochromatin and genome stability. Cell 107(3):323–337
110. Pluta AF, Saitoh N, Goldberg I, Earnshaw WC (1992) Identification of a subdomain of CENP-B that is necessary and sufficient for localization to the human centromere. J Cell Biol 116(5):1081–1093
111. Przewloka MR, Venkei Z, Bolanos-Garcia VM, Debski J, Dadlez M, Glover DM (2011) CENP-C is a structural platform for kinetochore assembly. Curr Biol 21(5):399–405
112. Régnier V, Vagnarelli P, Fukagawa T, Zerjal T, Burns E, Trouche D, Earnshaw W, Brown W (2005) CENP-A is required for accurate chromosome segregation and sustained kinetochore association of BubR1. Mol Cell Biol 25(10):3967–3981
113. Saffery R, Wong LH, Irvine DV, Bateman MA, Griffiths B, Cutts SM, Cancilla MR, Cendron AC, Stafford AJ, Choo KH (2001) Construction of neocentromere-based human minichromosomes by telomere-associated chromosomal truncation. Proc Natl Acad Sci U S A 98(10):5705–5710
114. Saitoh H, Tomkiel J, Cooke CA, Ratrie H 3rd, Maurer M, Rothfield NF, Earnshaw WC (1992) CENP-C, an autoantigen in scleroderma, is a component of the human inner kinetochore plate. Cell 70(1):115–125
115. Schueler MG, Higgins AW, Rudd MK, Gustashaw K, Willard HF (2001) Genomic and genetic definition of a functional human centromere. Science 294(5540):109–115
116. Scott KC, Merrett SL, Willard HF (2006) A heterochromatin barrier partitions the fission yeast centromere into discrete chromatin domains. Curr Biol 16(2):119–129
117. Screpanti E, De Antoni A, Alushin GM, Petrovic A, Melis T, Nogales E, Musacchio A (2011) Direct binding of Cenp-C to the Mis12 complex joins the inner and outer kinetochore. Curr Biol 21(5):391–398
118. Shang WH, Hori T, Martins NM, Toyoda A, Misu S, Monma N, Hiratani I, Maeshima K, Ikeo K, Fujiyama A, Kimura H, Earnshaw WC, Fukagawa T (2013) Chromosome engineering allows the efficient isolation of vertebrate neocentromeres. Dev Cell 24(6):635–648
119. Stimpson KM, Matheny JE, Sullivan BA (2012) Dicentric chromosomes: unique models to study centromere function and inactivation. Chromosome Res 20(5):595–605
120. Stinchcomb DT, Struhl K, Davis RW (1979) Isolation and characterisation of a yeast chromosomal replicator. Nature 282(5734):39–43
121. Struhl K, Stinchcomb DT, Scherer S, Davis RW (1979) High-frequency transformation of yeast: autonomous replication of hybrid DNA molecules. Proc Natl Acad Sci U S A 76(3):1035–1039
122. Sullivan BA, Karpen GH (2004) Centromeric chromatin exhibits a histone modification pattern that is distinct from both euchromatin and heterochromatin. Nat Struct Mol Biol 11(11):1076–1083
123. Suzuki N, Nakano M, Nozaki N, Egashira S, Okazaki T, Masumoto H (2004) CENP-B interacts with CENP-C domains containing Mif2 regions responsible for centromere localization. J Biol Chem 279(7):5934–5946
124. Tachiwana H, Kagawa W, Shiga T, Osakabe A, Miya Y, Saito K, Hayashi-Takanaka Y, Oda T, Sato M, Park SY, Kimura H, Kurumizaka H (2011) Crystal structure of the human centromeric nucleosome containing CENP-A. Nature 476(7359):232–235
125. Tagami H, Ray-Gallet D, Almouzni G, Nakatani Y (2004) Histone H3.1 and H3.3 complexes mediate nucleosome assembly pathways dependent or independent of DNA synthesis. Cell 116(1):51–61
126. Takahashi K, Murakami S, Chikashige Y, Funabiki H, Niwa O, Yanagida M (1992) A low copy number central sequence with strict symmetry and unusual chromatin structure in fission yeast centromere. Mol Biol Cell 3(7):819–835
127. Takahashi K, Chen ES, Yanagida M (2000) Requirement of Mis6 centromere connector for localizing a CENP-A-like protein in fission yeast. Science 288(5474):2215–2219
128. Talbert PB, Ahmad K, Almouzni G, Ausió J, Berger F, Bhalla PL, Bonner WM, Cande WZ, Chadwick BP, Chan SW, Cross GA, Cui L, Dimitrov SI, Doenecke D, Eirin-López JM, Gorovsky MA, Hake SB, Hamkalo BA, Holec S, Jacobsen SE, Kamieniarz K, Khochbin S, Ladurner AG, Landsman D, Latham JA, Loppin B, Malik HS, Marzluff WF, Pehrson JR, Postberg J, Schneider R, Singh MB, Smith MM, Thompson E, Torres-Padilla ME, Tremethick DJ, Turner BM, Waterborg JH, Wollmann H, Yelagandula R, Zhu B, Henikoff S (2012) A unified phylogeny-based nomenclature for histone variants. Epigenetics Chromatin 5:7
129. Tsuduki T, Nakano M, Yasuoka N, Yamazaki S, Okada T, Okamoto Y, Masumoto H (2006) An artificially constructed de novo human chromosome behaves almost identically to its natural counterpart during metaphase and anaphase in living cells. Mol Cell Biol 26(20):7682–7695
130. Wakai M, Abe S, Kazuki Y, Oshimura M, Ishikawa F (2014) A human artificial chromosome recapitulates the metabolism of native telomeres in mammalian cells. PLoS One 9(2):e88530
131. Weuts A, Voet T, Verbeeck J, Lambrechts N, Wirix E, Schoonjans L, Danloy S, Marynen P, Froyen G (2012) Telomere length homeostasis and telomere position effect on a linear human artificial chromosome are dictated by the genetic background. Nucleic Acids Res 40(22):11477–11489
132. Willard HF, Waye JS (1987) Hierarchical order in chromosome-specific human alpha satellite DNA. Trends Genet 3:192–198
133. Yoda K, Kitagawa K, Masumoto H, Muro Y, Okazaki T (1992) A human centromere protein, CENP-B, has a DNA binding domain containing four potential alpha helices at the NH2 terminus, which is separable from dimerizing activity. J Cell Biol 119(6):1413–1427