The Centromere

  • Beth A. Sullivan


Centromeres are chromosomal loci that assemble the proteinaceous kinetochore, maintain sister chromatid cohesion, regulate chromosome attachment to the spindle, and direct chromosome movement during cell division. Although the function of centromeres and proteins that contribute to their complex structure are conserved in eukaryotes, centromeric DNAs are strikingly divergent. In this chapter, I review centromere organization in a range of organisms, including unicellular eukaryotes, fruit flies, plants, and mammals. Sequence features and epigenetic mechanisms of centromere identity and regulation, including DNA–protein interactions, post-translational modifications, RNA, and protein dosage that influence centromere-specific chromatin architecture are discussed. Understanding the assembly and organization of centromeres and the contributions of sequence and epigenetic features in centromere identity and diversity remain important areas of study in chromosome biology.


Fission Yeast Alpha Satellite Human Artificial Chromosome Centromeric Chromatin Kinetochore Assembly 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


  1. Abad JP et al. (2000) Pericentromeric regions containing 1.688 satellite DNA sequences show anti-kinetochore antibody staining in prometaphase chromosomes of Drosophila melanogaster. Mol Gen Genet 264: 371–377.PubMedGoogle Scholar
  2. Aker M, Huang HV (1996) Extreme heterogeneity of minor satellite repeat arrays in inbred strains of mice. Mamm Genome 7: 62–64.PubMedGoogle Scholar
  3. Alexandrov IA et al. (1993) Definition of a new alpha satellite suprachromosomal family characterized by monomeric organization. Nucleic Acids Res 21: 2209–2215.PubMedGoogle Scholar
  4. Allshire RC et al. (1994) Position effect variegation at fission yeast centromeres. Cell 76: 157–169.PubMedGoogle Scholar
  5. Allshire RC et al. (1995) Mutations derepressing silent centromeric domains in fission yeast disrupt chromosome segregation. Genes Dev 9: 218–233.PubMedGoogle Scholar
  6. Alonso A et al. (2007) Co-localization of CENP-C and CENP-H to discontinuous domains of CENP-A chromatin at human neocentromeres. Genome Biol 8: R148.PubMedGoogle Scholar
  7. Ando S et al. (2002) CENP-A, -B, and -C chromatin complex that contains the I-type alpha-satellite array constitutes the prekinetochore in HeLa cells. Mol Cell Biol 22: 2229–2241.PubMedGoogle Scholar
  8. Andreyeva EN et al. (2007) High-resolution analysis of Drosophila heterochromatin organization using SuUR Su(var)3-9 double mutants. Proc Natl Acad Sci USA 104: 12819–12824.PubMedGoogle Scholar
  9. Barbosa-Cisneros O et al. (1997) CENP-B autoantigen is a conserved protein from humans to higher plants: identification of the aminoterminal domain in Phaseolus vulgaris. Rev Rhum Engl Ed 64: 368–374.PubMedGoogle Scholar
  10. Baum M, Clarke L (2000) Fission yeast homologs of human CENP-B have redundant functions affecting cell growth and chromosome segregation. Mol Cell Biol 20: 2852–2864.PubMedGoogle Scholar
  11. Baum M et al. (1994) The centromeric K-type repeat and the central core are together sufficient to establish a functional Schizosaccharomyces pombe centromere. Mol Biol Cell 5: 747–761.PubMedGoogle Scholar
  12. Baum M et al. (2006) Formation of functional centromeric chromatin is specified epigenetically in Candida albicans. Proc Natl Acad Sci USA 103: 14877–14882.PubMedGoogle Scholar
  13. Bernard P et al. (2001) Requirement of Heterochromatin for Cohesion at Centromeres. Science 294: 2539–2542.PubMedGoogle Scholar
  14. Black BE et al. (2004) Structural determinants for generating centromeric chromatin. Nature 430: 578–582.PubMedGoogle Scholar
  15. Black BE et al. (2007) Centromere identity maintained by nucleosomes assembled with histone H3 containing the CENP-A targeting domain. Mol Cell 25: 309–322.PubMedGoogle Scholar
  16. Blower MD, Karpen GH (2001) The role of Drosophila CID in kinetochore formation, cell-cycle progression and heterochromatin interactions. Nat Cell Biol 3: 730–739.PubMedGoogle Scholar
  17. Blower MD et al. (2002) Conserved organization of centromeric chromatin in flies and humans. Dev Cell 2: 319–330.PubMedGoogle Scholar
  18. Bouzinba-Segard H et al. (2006) Accumulation of small murine minor satellite transcripts leads to impaired centromeric architecture and function. Proc Natl Acad Sci USA 103: 8709–8714.PubMedGoogle Scholar
  19. Brown KE et al. (1994) Dissecting the centromere of the human Y chromosome with cloned telomeric DNA. Hum Mol Genet 3: 1227–1237.PubMedGoogle Scholar
  20. Cam HP et al. (2005) Comprehensive analysis of heterochromatin- and RNAi-mediated epigenetic control of the fission yeast genome. Nat Genet 37: 809–819.PubMedGoogle Scholar
  21. Camahort R et al. (2007) Scm3 is essential to recruit the histone h3 variant cse4 to centromeres and to maintain a functional kinetochore. Mol Cell 26: 853–865.PubMedGoogle Scholar
  22. Castillo AG et al. (2007) Plasticity of fission yeast CENP-A chromatin driven by relative levels of histone H3 and H4. PLoS Genet 3: e121.PubMedGoogle Scholar
  23. Choo KH (2001) Domain organization at the centromere and neocentromere. Dev Cell 1: 165–177.PubMedGoogle Scholar
  24. Chueh AC et al. (2005) Variable and hierarchical size distribution of L1-retroelement-enriched CENP-A clusters within a functional human neocentromere. Hum Mol Genet 14: 85–93.PubMedGoogle Scholar
  25. Clarke L et al. (1993) Structure and function of Schizosaccharomyces pombe centromeres. Cold Spring Harb Symp Quant Biol 58: 687–695.PubMedGoogle Scholar
  26. Cleveland DW et al. (2003) Centromeres and kinetochores: from epigenetics to mitotic checkpoint signaling. Cell 112: 407–421.PubMedGoogle Scholar
  27. Collins KA et al. (2007) The overexpression of a Saccharomyces cerevisiae centromeric histone H3 variant mutant protein leads to a defect in kinetochore biorientation. Genetics 175: 513–525.PubMedGoogle Scholar
  28. Collins KA et al. (2004) Proteolysis contributes to the exclusive centromere localization of the yeast Cse4/CENP-A histone H3 variant. Curr Biol 14: 1968–1972.PubMedGoogle Scholar
  29. Conde e Silva N et al. (2007) CENP-A-containing nucleosomes: easier disassembly versus exclusive centromeric localization. J Mol Biol 370: 555–573.PubMedGoogle Scholar
  30. Copenhaver GP et al. (1999) Genetic definition and sequence analysis of Arabidopsis centromeres. Science 286: 2468–2474.PubMedGoogle Scholar
  31. Dalal Y et al. (2007a) Structure, dynamics, and evolution of centromeric nucleosomes. Proc Natl Acad Sci U S A. 104:15974–15981.Google Scholar
  32. Dalal Y et al. (2007b) Tetrameric structure of centromeric nucleosomes in interphase Drosophila cells. PLoS Biol 5: e218.Google Scholar
  33. Dawe RK et al. (1999) A maize homolog of mammalian CENPC is a constitutive component of the inner kinetochore. Plant Cell 11: 1227–1238.PubMedGoogle Scholar
  34. De Wulf P et al. (2003) Hierarchical assembly of the budding yeast kinetochore from multiple subcomplexes. Genes Dev 17: 2902–2921.PubMedGoogle Scholar
  35. Depinet TW et al. (1997) Characterization of neo-centromeres in marker chromosomes lacking detectable alpha-satellite DNA. Hum Mol Genet 6: 1195–1204.PubMedGoogle Scholar
  36. Di Stefano L et al. (2007) Mutation of Drosophila Lsd1 disrupts H3-K4 methylation, resulting in tissue-specific defects during development. Curr Biol 17: 808–812.PubMedGoogle Scholar
  37. du Sart D et al. (1997) A functional neo-centromere formed through activation of a latent human centromere and consisting of non-alpha-satellite DNA. Nat Genet 16: 144–153.PubMedGoogle Scholar
  38. Dykxhoorn DM et al. (2003) Killing the messenger: short RNAs that silence gene expression. Nat Rev Mol Cell Biol 4: 457–467.PubMedGoogle Scholar
  39. Earnshaw W et al. (1986) Three human chromosomal autoantigens are recognized by sera from patients with anti-centromere antibodies. J Clin Invest 77: 426–430.PubMedGoogle Scholar
  40. Earnshaw WC, Migeon BR (1985) Three related centromere proteins are absent from the inactive centromere of a stable isodicentric chromosome. Chromosoma 92: 290–296.PubMedGoogle Scholar
  41. Earnshaw WC, Rothfield N (1985) Identification of a family of human centromere proteins using autoimmune sera from patients with scleroderma. Chromosoma 91: 313–321.PubMedGoogle Scholar
  42. Ebert A et al. (2004) Su(var) genes regulate the balance between euchromatin and heterochromatin in Drosophila. Genes Dev 18: 2973–2983.PubMedGoogle Scholar
  43. Ekwall K et al. (1997) Transient inhibition of histone deacetylation alters the structural and functional imprint at fission yeast centromeres. Cell 91: 1021–1032.PubMedGoogle Scholar
  44. Farr CJ et al. (1995) Generation of a human X-derived minichromosome using telomere- associated chromosome fragmentation. EMBO J 14: 5444–5454.PubMedGoogle Scholar
  45. Fisher AM et al. (1997) Centromeric inactivation in a dicentric human Y;21 translocation chromosome. Chromosoma 106: 199–206.PubMedGoogle Scholar
  46. Foltz DR et al. (2006) The human CENP-A centromeric nucleosome-associated complex. Nat Cell Biol 8: 458–469.PubMedGoogle Scholar
  47. Fujita Y et al. (2007) Priming of centromere for CENP-A recruitment by human hMis18alpha, hMis18beta, and M18BP1. Dev Cell 12: 17–30.PubMedGoogle Scholar
  48. Fukagawa T et al. (2001a) CENP-H, a constitutive centromere component, is required for centromere targeting of CENP-C in vertebrate cells. Embo J 20: 4603–4617.Google Scholar
  49. Fukagawa T et al. (2004) Dicer is essential for formation of the heterochromatin structure in vertebrate cells. Nat Cell Biol 6: 784–791.PubMedGoogle Scholar
  50. Fukagawa T et al. (2001b) Creation and characterization of temperature-sensitive CENP-C mutants in vertebrate cells. Nucleic Acids Res 29: 3796–3803.Google Scholar
  51. Furuyama S, Biggins S (2007) Centromere identity is specified by a single centromeric nucleosome in budding yeast. Proc Natl Acad Sci USA 104: 14706–14711.PubMedGoogle Scholar
  52. Giet R, Glover DM (2001) Drosophila aurora B kinase is required for histone H3 phosphorylation and condensin recruitment during chromosome condensation and to organize the central spindle during cytokinesis. J Cell Biol 152: 669–682.PubMedGoogle Scholar
  53. Gilbert N, Allan J (2001) Distinctive higher-order chromatin structure at mammalian centromeres. Proc Natl Acad Sci USA 98: 11949–11954.PubMedGoogle Scholar
  54. Greaves IK et al. (2007) H2A.Z contributes to the unique 3D structure of the centromere. Proc Natl Acad Sci USA 104: 525–530.PubMedGoogle Scholar
  55. Grimes BR et al. (2002) alpha-Satellite DNA and Vector Composition Influence Rates of Human Artificial Chromosome Formation. Mol Ther 5: 798–805.PubMedGoogle Scholar
  56. Guenatri M et al. (2004) Mouse centric and pericentric satellite repeats form distinct functional heterochromatin. J Cell Biol 166: 493–505.PubMedGoogle Scholar
  57. Haaf T, Schmid M (1990) Y isochromosome associated with a mosaic karyotype and inactivation of the centromere. Hum Genet 85: 486–490.PubMedGoogle Scholar
  58. Haaf T, Ward DC (1994) Structural analysis of alpha-satellite DNA and centromere proteins using extended chromatin and chromosomes. Hum Mol Genet 3: 697–709.PubMedGoogle Scholar
  59. Hagstrom KA et al. (2002) C. elegans condensin promotes mitotic chromosome architecture, centromere organization, and sister chromatid segregation during mitosis and meiosis. Genes Dev 16: 729–742.PubMedGoogle Scholar
  60. Hahnenberger KM et al. (1991) Identification of DNA regions required for mitotic and meiotic functions within the centromere of Schizosaccharomyces pombe chromosome I. Mol Cell Biol 11: 2206–2215.PubMedGoogle Scholar
  61. Harrington JJ et al. (1997) Formation of de novo centromeres and construction of first-generation human artificial microchromosomes. Nat Genet 15: 345–355.PubMedGoogle Scholar
  62. Hayashi T et al. (2004) Mis16 and Mis18 are required for CENP-A loading and histone deacetylation at centromeres. Cell 118: 715–729.PubMedGoogle Scholar
  63. Hendzel MJ et al. (1997) Mitosis-specific phosphorylation of histone H3 initiates primarily within pericentromeric heterochromatin during G2 and spreads in an ordered fashion coincident with mitotic chromosome condensation. Chromosoma 106: 348–360.PubMedGoogle Scholar
  64. Heslop-Harrison JS et al. (2003) Tandemly repeated DNA sequences and centromeric chromosomal regions of Arabidopsis species. Chromosome Res 11: 241–253.PubMedGoogle Scholar
  65. Heun P et al. (2006) Mislocalization of the Drosophila centromere-specific histone CID promotes formation of functional ectopic kinetochores. Dev Cell 10: 303–315.PubMedGoogle Scholar
  66. Higgins AW et al. (2005) Engineered human dicentric chromosomes show centromere plasticity. Chromosome Res 13: 745–762.PubMedGoogle Scholar
  67. Hudson DF et al. (1998) Centromere protein B null mice are mitotically and meiotically normal but have lower body and testis weights. J Cell Biol 141: 309–319.PubMedGoogle Scholar
  68. Ikeno M et al. (1998) Construction of YAC-based mammalian artificial chromosomes. Nat Biotechnol 16: 431–439.PubMedGoogle Scholar
  69. Ikeno M et al. (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: 1245–1257.PubMedGoogle Scholar
  70. Ing PS, Smith SD (1983) Cytogenetic studies of a patient with mosaicism of isochromosome 13q and a dicentric (Y;13) translocation showing differential centromeric activity. Clin Genet 24: 194–199.PubMedGoogle Scholar
  71. Izuta H et al. (2006) Comprehensive analysis of the ICEN (Interphase Centromere Complex) components enriched in the CENP-A chromatin of human cells. Genes Cells 11: 673–684.PubMedGoogle Scholar
  72. Jager H et al. (2005) The Drosophila melanogaster condensin subunit Cap-G interacts with the centromere-specific histone H3 variant CID. Chromosoma 113: 350–361.PubMedGoogle Scholar
  73. Jansen LE et al. (2007) Propagation of centromeric chromatin requires exit from mitosis. J Cell Biol 176: 795–805.PubMedGoogle Scholar
  74. Jiang J et al. (2003) A molecular view of plant centromeres. Trends Plant Sci 8: 570–575.PubMedGoogle Scholar
  75. Jin W et al. (2004) Maize centromeres: organization and functional adaptation in the genetic background of oat. Plant Cell 16: 571–581.PubMedGoogle Scholar
  76. Kalitsis P et al. (2006) Mouse telocentric sequences reveal a high rate of homogenization and possible role in Robertsonian translocation. Proc Natl Acad Sci USA 103: 8786–8791.PubMedGoogle Scholar
  77. Kimura A, Horikoshi M (2004) Partition of distinct chromosomal regions: negotiable border and fixed border. Genes Cells 9: 499–508.PubMedGoogle Scholar
  78. Kipling D et al. (1991) Mouse minor satellite DNA genetically maps to the centromere and is physically linked to the proximal telomere. Genomics 11: 235–241.PubMedGoogle Scholar
  79. Kipling D et al. (1995) CENP-B binds a novel centromeric sequence in the Asian mouse Mus caroli. Mol Cell Biol 15: 4009–4020.PubMedGoogle Scholar
  80. Kitagawa K et al. (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: 1602–1612.PubMedGoogle Scholar
  81. Kuznetsova I et al. (2006) High-resolution organization of mouse centromeric and pericentromeric DNA. Cytogenet Genome Res 112: 248–255.PubMedGoogle Scholar
  82. Kuznetsova IS et al. (2005) New types of mouse centromeric satellite DNAs. Chromosome Res 13: 9–25.PubMedGoogle Scholar
  83. Lam AL et al. (2006) Human centromeric chromatin is a dynamic chromosomal domain that can spread over noncentromeric DNA. Proc Natl Acad Sci USA 103: 4186–4191.PubMedGoogle Scholar
  84. Lee C et al. (1997) Human centromeric DNAs. Hum Genet 100: 291–304.PubMedGoogle Scholar
  85. Lee HR et al. (2006) Transcription and evolutionary dynamics of the centromeric satellite repeat CentO in rice. Mol Biol Evol 23: 2505–2520.PubMedGoogle Scholar
  86. Lermontova I et al. (2006) Loading of Arabidopsis centromeric histone CENH3 occurs mainly during G2 and requires the presence of the histone fold domain. Plant Cell 18: 2443–2451.PubMedGoogle Scholar
  87. Locovei AM et al. (2006) The CENP-B homolog, Abp1, interacts with the initiation protein Cdc23 (MCM10) and is required for efficient DNA replication in fission yeast. Cell Div 1: 27.PubMedGoogle Scholar
  88. Maddox PS et al. (2007) Functional genomics identifies a Myb domain-containing protein family required for assembly of CENP-A chromatin. J Cell Biol 176: 757–763.PubMedGoogle Scholar
  89. Maddox PS et al. (2004) “Holo”er than thou: chromosome segregation and kinetochore function in C. elegans. Chromosome Res 12: 641–653.PubMedGoogle Scholar
  90. Maggert KA, Karpen GH (2001) The activation of a neocentromere in Drosophila requires proximity to an endogenous centromere. Genetics 158: 1615–1628.PubMedGoogle Scholar
  91. Martens JH et al. (2005) The profile of repeat-associated histone lysine methylation states in the mouse epigenome. Embo J 24: 800–812.PubMedGoogle Scholar
  92. May BP et al. (2005) Differential regulation of strand-specific transcripts from Arabidopsis centromeric satellite repeats. PLoS Genet 1: e79.PubMedGoogle Scholar
  93. McAinsh AD et al. (2003) Structure, function, and regulation of budding yeast kinetochores. Annu Rev Cell Dev Biol 19: 519–539.PubMedGoogle Scholar
  94. Mishra PK et al. (2007) Centromere size and position in Candida albicans are evolutionarily conserved independent of DNA sequence heterogeneity. Mol Genet Genomics 278:455–465.PubMedGoogle Scholar
  95. Mizuguchi G et al. (2007) Nonhistone Scm3 and histones CenH3-H4 assemble the core of centromere-specific nucleosomes. Cell 129: 1153–1164.PubMedGoogle Scholar
  96. Monen J et al. (2005) Differential role of CENP-A in the segregation of holocentric C. elegans chromosomes during meiosis and mitosis. Nat Cell Biol 7: 1248–1255.PubMedGoogle Scholar
  97. Moore LL, Roth MB (2001) HCP-4, a CENP-C-like protein in Caenorhabditis elegans, is required for resolution of sister centromeres. J Cell Biol 153: 1199–1208.PubMedGoogle Scholar
  98. Moreno-Moreno O et al. (2006) Proteolysis restricts localization of CID, the centromere-specific histone H3 variant of Drosophila, to centromeres. Nucleic Acids Res 34: 6247–6255.PubMedGoogle Scholar
  99. Motamedi MR et al. (2004) Two RNAi complexes, RITS and RDRC, physically interact and localize to noncoding centromeric RNAs. Cell 119: 789–802.PubMedGoogle Scholar
  100. Murphy TD, Karpen GH (1995) Interactions between the nod+ kinesin-like gene and extracentromeric sequences are required for transmission of a Drosophila minichromosome. Cell 81: 139–148.PubMedGoogle Scholar
  101. Nagaki K et al. (2004) Sequencing of a rice centromere uncovers active genes. Nat Genet 36: 138–145.PubMedGoogle Scholar
  102. Nagaki K et al. (2003) Chromatin immunoprecipitation reveals that the 180-bp satellite repeat is the key functional DNA element of Arabidopsis thaliana centromeres. Genetics 163: 1221–1225.PubMedGoogle Scholar
  103. Nakagawa H et al. (2002) Fission yeast CENP-B homologs nucleate centromeric heterochromatin by promoting heterochromatin-specific histone tail modifications. Genes Dev 16: 1766–1778.PubMedGoogle Scholar
  104. Nakashima H et al. (2005) Assembly of additional heterochromatin distinct from centromere-kinetochore chromatin is required for de novo formation of human artificial chromosome. J Cell Sci 118: 5885–5898.PubMedGoogle Scholar
  105. Nishihashi A et al. (2002) CENP-I is essential for centromere function in vertebrate cells. Dev Cell 2: 463–476.PubMedGoogle Scholar
  106. Noma K et al. (2006) A role for TFIIIC transcription factor complex in genome organization. Cell 125: 859–872.PubMedGoogle Scholar
  107. Nonaka N et al. (2002) Recruitment of cohesin to heterochromatic regions by Swi6/HP1 in fission yeast. Nat Cell Biol 4: 89–93.PubMedGoogle Scholar
  108. Nusbaum C et al. (2006) DNA sequence and analysis of human chromosome 8. Nature 439: 331–335.PubMedGoogle Scholar
  109. Oegema K et al. (2001) Functional analysis of kinetochore assembly in Caenorhabditis elegans. J Cell Biol 153: 1209–1226.PubMedGoogle Scholar
  110. Ogura Y et al. (2004) Characterization of a CENP-C homolog in Arabidopsis thaliana. Genes Genet Syst 79: 139–144.PubMedGoogle Scholar
  111. Ohzeki J et al. (2002) CENP-B box is required for de novo centromere chromatin assembly on human alphoid DNA. J Cell Biol 159: 765–775.PubMedGoogle Scholar
  112. Okada M et al. (2006) The CENP-H-I complex is required for the efficient incorporation of newly synthesized CENP-A into centromeres. Nat Cell Biol 8: 446–457.PubMedGoogle Scholar
  113. Okada T et al. (2007) CENP-B controls centromere formation depending on the chromatin context. Cell 131: 1287–1300.PubMedGoogle Scholar
  114. Okamoto Y et al. (2007) A minimal CENP-A core is required for nucleation and maintenance of a functional human centromere. Embo J 26: 1279–1291.PubMedGoogle Scholar
  115. Palmer DK et al. (1989) Biochemical analysis of CENP-A, a centromeric protein with histone-like properties. Prog Clin Biol Res 318: 61–72.PubMedGoogle Scholar
  116. Palmer DK et al. (1991) Purification of the centromere-specific protein CENP-A and demonstration that it is a distinctive histone. Proc Natl Acad Sci USA 88: 3734–3738.PubMedGoogle Scholar
  117. Palmer DK et al. (1987) A 17-kD centromere protein (CENP-A) copurifies with nucleosome core particles and with histones. J Cell Biol 104: 805–815.PubMedGoogle Scholar
  118. Partridge JF et al. (2002) cis-acting DNA from fission yeast centromeres mediates histone H3 methylation and recruitment of silencing factors and cohesin to an ectopic site. Curr Biol 12: 1652–1660.PubMedGoogle Scholar
  119. Peters AH et al. (2003) Partitioning and plasticity of repressive histone methylation states in mammalian chromatin. Mol Cell 12: 1577–1589.PubMedGoogle Scholar
  120. Pidoux AL, Allshire RC (2004) Kinetochore and heterochromatin domains of the fission yeast centromere. Chromosome Res 12: 521–534.PubMedGoogle Scholar
  121. Pidoux AL, Allshire RC (2005) The role of heterochromatin in centromere function. Philos Trans R Soc Lond B Biol Sci 360: 569–579.PubMedGoogle Scholar
  122. Politi V et al. (2002) CENP-C binds the alpha-satellite DNA in vivo at specific centromere domains. J Cell Sci 115: 2317–2327.PubMedGoogle Scholar
  123. Ross MT et al. (2005) The DNA sequence of the human X chromosome. Nature 434: 325–337.PubMedGoogle Scholar
  124. Rudd MK, Willard HF (2004) Analysis of the centromeric regions of the human genome assembly. Trends Genet 20: 529–533.PubMedGoogle Scholar
  125. Rudert F et al. (1995) Transcripts from opposite strands of gamma satellite DNA are differentially expressed during mouse development. Mamm Genome 6: 76–83.PubMedGoogle Scholar
  126. Saffery R et al. (2003) Transcription within a functional human centromere. Mol Cell 12: 509–516.PubMedGoogle Scholar
  127. Saitoh H et al. (1992) CENP-C, an autoantigen in scleroderma, is a component of the human inner kinetochore plate. Cell 70: 115–125.PubMedGoogle Scholar
  128. Saitoh S et al. (1997) Mis6, a fission yeast inner centromere protein, acts during G1/S and forms specialized chromatin required for equal segregation. Cell 90: 131–143.PubMedGoogle Scholar
  129. Sanyal K et al. (2004) Centromeric DNA sequences in the pathogenic yeast Candida albicans are all different and unique. Proc Natl Acad Sci USA 101: 11374–11379.PubMedGoogle Scholar
  130. Saunders M et al. (1988) Chromatin structure of altered yeast centromeres. Proc Natl Acad Sci USA 85: 175–179.PubMedGoogle Scholar
  131. Schuh M et al. (2007) Incorporation of Drosophila CID/CENP-A and CENP-C into centromeres during early embryonic anaphase. Curr Biol 17: 237–243.PubMedGoogle Scholar
  132. Scott KC et al. (2006) A heterochromatin barrier partitions the fission yeast centromere into discrete chromatin domains. Curr Biol 16: 119–129.PubMedGoogle Scholar
  133. Shelby RD et al. (2000) Chromatin assembly at kinetochores is uncoupled from DNA replication. J Cell Biol 151: 1113–1118.PubMedGoogle Scholar
  134. Shelby RD et al. (1997) Assembly of CENP-A into centromeric chromatin requires a cooperative array of nucleosomal DNA contact sites. J Cell Biol 136: 501–513.PubMedGoogle Scholar
  135. Shi J, Dawe RK (2006) Partitioning of the maize epigenome by the number of methyl groups on histone H3 lysines 9 and 27. Genetics 173: 1571–1583.PubMedGoogle Scholar
  136. Song K et al. (2002) Mutational analysis of the central centromere targeting domain of human centromere protein C, (CENP-C). Exp Cell Res 275: 81–91.PubMedGoogle Scholar
  137. Spence JM et al. (2002) Co-localization of centromere activity, proteins and topoisomerase II within a subdomain of the major human X alpha-satellite array. EMBO J 21: 5269–5280.PubMedGoogle Scholar
  138. Stoler S et al. (2007) Scm3, an essential Saccharomyces cerevisiae centromere protein required for G2/M progression and Cse4 localization. Proc Natl Acad Sci USA 104: 10571–10576.PubMedGoogle Scholar
  139. Sugata N et al. (2000) Human CENP-H multimers colocalize with CENP-A and CENP-C at active centromere--kinetochore complexes. Hum Mol Genet 9: 2919–2926.PubMedGoogle Scholar
  140. Sullivan BA (2002) Centromere round-up at the heterochromatin corral. Trends Biotechnol 20: 89–92.PubMedGoogle Scholar
  141. Sullivan BA et al. (2001) Determining centromere identity: cyclical stories and forking paths. Nat Rev Genet 2: 584–596.PubMedGoogle Scholar
  142. 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: 1076–1083.PubMedGoogle Scholar
  143. Sullivan BA, Schwartz S (1995) Identification of centromeric antigens in dicentric Robertsonian translocations: CENP-C and CENP-E are necessary components of functional centromeres. Hum Mol Genet 4: 2189–2197.PubMedGoogle Scholar
  144. Sun X et al. (2002) Sequence analysis of a functional Drosophila centromere. Genome Res 13: 182–188.Google Scholar
  145. Sun X et al. (1997) Molecular structure of a functional Drosophila centromere. Cell 91: 1007–1019.PubMedGoogle Scholar
  146. Takahashi K et al. (1992) A low copy number central sequence with strict symmetry and unusual chromatin structure in fission yeast centromere. Mol Biol Cell 3: 819–835.PubMedGoogle Scholar
  147. Talbert PB et al. (2004) Adaptive evolution of centromere proteins in plants and animals. J Biol 3: 18.PubMedGoogle Scholar
  148. Tanaka Y et al. (2005) Human centromere protein B induces translational positioning of nucleosomes on alpha-satellite sequences. J Biol Chem 280: 41609–41618.PubMedGoogle Scholar
  149. Tomkiel J et al. (1994) CENP-C is required for maintaining proper kinetochore size and for a timely transition to anaphase. J Cell Biol 125: 531–545.PubMedGoogle Scholar
  150. Topp CN et al. (2004) Centromere-encoded RNAs are integral components of the maize kinetochore. Proc Natl Acad Sci U S A 101: 15986–15991.PubMedGoogle Scholar
  151. Trazzi S et al. (2002) In vivo functional dissection of human inner kinetochore protein CENP-C. J Struct Biol 140: 39–48.PubMedGoogle Scholar
  152. Van Hooser AA et al. (2001) Specification of kinetochore-forming chromatin by the histone H3 variant CENP-A. J Cell Sci 114: 3529–3542.PubMedGoogle Scholar
  153. Verdel A et al. (2004) RNAi-mediated targeting of heterochromatin by the RITS complex. Science 303: 672–676.PubMedGoogle Scholar
  154. Verdel A, Moazed D (2005) RNAi-directed assembly of heterochromatin in fission yeast. FEBS Lett 579: 5872–5878.PubMedGoogle Scholar
  155. Vissel B, Choo KH (1989) Mouse major (gamma) satellite DNA is highly conserved and organized into extremely long tandem arrays: implications for recombination between nonhomologous chromosomes. Genomics 5: 407–414.PubMedGoogle Scholar
  156. Wevrick R et al. (1990) Partial deletion of alpha satellite DNA associated with reduced amounts of the centromere protein CENP-B in a mitotically stable human chromosome rearrangement. Mol Cell Biol 10: 6374–6380.PubMedGoogle Scholar
  157. Willard HF (1985) Chromosome-specific organization of human alpha satellite DNA. Am J Hum Genet 37: 524–532.PubMedGoogle Scholar
  158. Willard HF (1990) Centromeres of mammalian chromosomes. Trends Genet 6: 410–416.PubMedGoogle Scholar
  159. Willard HF, Waye JS (1987) Hierarchical order in chromosome-specific human alpha satellite DNA. Trends Genet 3: 192–198.Google Scholar
  160. Wong AK et al. (1990) The chromosomal distribution of the major and minor satellite is not conserved in the genus Mus. Chromosoma 99: 190–195.PubMedGoogle Scholar
  161. Wong AK, Rattner JB (1988) Sequence organization and cytological localization of the minor satellite of mouse. Nucleic Acids Res 16: 11645–11661.PubMedGoogle Scholar
  162. Wong LH et al. (2007) Centromere RNA is a key component for the assembly of nucleoproteins at the nucleolus and centromere. Genome Res 17: 1146–1160.PubMedGoogle Scholar
  163. Wong NC et al. (2006) Permissive transcriptional activity at the centromere through pockets of DNA hypomethylation. PLoS Genet 2: e17.PubMedGoogle Scholar
  164. Yan H et al. (2005) Transcription and histone modifications in the recombination-free region spanning a rice centromere. Plant Cell 17: 3227–3238.PubMedGoogle Scholar
  165. Yoda K et al. (2000) Human centromere protein A (CENP-A) can replace histone H3 in nucleosome reconstitution in vitro. Proc Natl Acad Sci U S A 97: 7266–7271.PubMedGoogle Scholar
  166. Zeng K et al. (2004) Localisation of centromeric proteins to a fraction of mouse minor satellite DNA on a mini-chromosome in human, mouse and chicken cells. Chromosoma 113: 84–91.PubMedGoogle Scholar
  167. Zinkowski RP et al. (1991) The centromere-kinetochore complex: a repeat subunit model. J Cell Biol 113: 1091–1110.PubMedGoogle Scholar

Copyright information

© Springer Science+Business Media, LLC 2009

Authors and Affiliations

  1. 1.Institute for Genome Sciences & Policy and Department of Molecular Genetics and MicrobiologyDuke UniversityDurhamU.S.A

Personalised recommendations