Centromeres: Sequences, Structure, and Biology



Although technological advances have continued to change the speed, cost, and number of plant genomes sequenced (see Flagel and Blackman 2012, this volume), parts of genomes remain to be sequenced and explored. Even the best-sequenced plant genomes, including Arabidopsis thaliana and rice, are missing 7–8% of their total genomic information (Kaul et al. 2000; Goff et al. 2002; Yu et al. 2002). One chromosomal region not often sequenced in genome projects is the centromere. Centromeres of almost all higher eukaryotes contain large stretches (up to several megabases) of tandemly repeated arrays of satellite DNA and retrotransposons. Such long arrays of highly homogenized repetitive DNA sequences cannot readily be cloned, sequenced, and assembled using the currently available cloning and sequencing technologies.


  1. Alfenito MR, Birchler JA (1993) Molecular characterization of a maize B chromosome centric sequence. Genetics 135:589–597PubMedGoogle Scholar
  2. Ananiev EV, Phillips RL, Rines HW (1998) Chromosome-specific molecular organization of maize (Zea mays L.) centromeric regions. Proc Natl Acad Sci USA 95:13073–13078PubMedCrossRefGoogle Scholar
  3. Bao W, Zhang W, Yang Q, Zhang Y, Han B, Gu M, Xue Y, Cheng Z (2006) Diversity of centromeric repeats in two closely related wild rice species, Oryza officinalis and Oryza rhizomatis. Mol Genet Genomics 275:421–430PubMedCrossRefGoogle Scholar
  4. Black BE, Foltz DR, Chakravarthy S, Luger K, Woods VL, Cleveland DW (2004) Structural determinants for generating centromeric chromatin. Nature 430:578–582PubMedCrossRefGoogle Scholar
  5. Bouzinba-Segard H, Guais A, Francastel C (2006) Accumulation of small murine minor satellite transcripts leads to impaired centromeric architecture and function. Proc Natl Acad Sci USA 103:8709–8714PubMedCrossRefGoogle Scholar
  6. Brown JD, O’Neill RJ (2010) Chromosomes, conflict, and epigenetics: chromosomal speciation revisited. Annu Rev Genomics Hum Genet 11:291–316PubMedCrossRefGoogle Scholar
  7. Buchwitz BJ, Ahmad K, Moore LL, Roth MB, Henikoff S (1999) Cell division: a histone-H3-like protein in C. elegans. Nature 401:547–548PubMedCrossRefGoogle Scholar
  8. Carone D, Longo M, Ferreri G, Hall L, Harris M, Shook N, Bulazel K, Carone B, Obergfell C, O’Neill M, O’Neill R (2009) A new class of retroviral and satellite encoded small RNAs emanates from mammalian centromeres. Chromosoma 118:113–125PubMedCrossRefGoogle Scholar
  9. Cheng Z, Dong F, Langdon T, Ouyang S, Buell CR, Gu M, Blattner FR, Jiang J (2002) Functional rice centromeres are marked by a satellite repeat and a centromere-specific retrotransposon. Plant Cell 14:1691–1704PubMedCrossRefGoogle Scholar
  10. Cooper JL, Henikoff S (2004) Adaptive evolution of the histone fold domain in centromeric histones. Mol Biol Evol 21:1712–1718PubMedCrossRefGoogle Scholar
  11. Dong F, Miller JT, Jackson SA, Wang G-L, Ronald PC, Jiang J (1998) Rice (Oryza sativa) centromeric regions consist of complex DNA. Proc Natl Acad Sci USA 95:8135–8140PubMedCrossRefGoogle Scholar
  12. Du J, Tian Z, Hans CS, Laten HM, Cannon SB, Jackson SA, Shoemaker RC, Ma J (2010a) Evolutionary conservation, diversity and specificity of LTR-retrotransposons in flowering plants: insights from genome-wide analysis and multi-specific comparison. Plant J 63:584–598PubMedCrossRefGoogle Scholar
  13. Du Y, Topp CN, Dawe RK (2010b) DNA binding of centromere protein C (CENPC) is stabilized by single-stranded RNA. PLoS Genet 6:e1000835PubMedCrossRefGoogle Scholar
  14. 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 137:485–497PubMedCrossRefGoogle Scholar
  15. Earnshaw WC, Rothfield N (1985) Identification of a family of human centromere proteins using autoimmune sera from patients with scleroderma. Chromosoma 91:313–321PubMedCrossRefGoogle Scholar
  16. Flagel L, Blackman B (2012) The first ten years of plant genome sequencing and prospects for the next decade. In: Wendel JF (ed) Plant genome diversity, vol 1, Plant genomes, their residents, and their evolutionary dynamics. Springer, Wien, New YorkGoogle Scholar
  17. Foltz DR, Jansen LET, Black BE, Bailey AO, Yates JR, Cleveland DW (2006) The human CENP-A centromeric nucleosome-associated complex. Nat Cell Biol 8:458–469PubMedCrossRefGoogle Scholar
  18. Foltz DR, Jansen LET, Bailey AO, Yates Iii JR, Bassett EA, Wood S, Black BE, Cleveland DW (2009) Centromere-specific assembly of CENP-A nucleosomes is mediated by HJURP. Cell 137:472–484PubMedCrossRefGoogle Scholar
  19. Fujita Y, Hayashi T, Kiyomitsu T, Toyoda Y, Kokubu A, Obuse C, Yanagida M (2007) Priming of centromere for CENP-A recruitment by human hMis18α, hMis18β, and M18BP1. Dev Cell 12:17–30PubMedCrossRefGoogle Scholar
  20. Fukagawa T, Nogami M, Yoshikawa M, Ikeno M, Okazaki T, Takami Y, Nakayama T, Oshimura M (2004) Dicer is essential for formation of the heterochromatin structure in vertebrate cells. Nat Cell Biol 6:784–791PubMedCrossRefGoogle Scholar
  21. Gao D, Gill N, Kim HR, Walling JG, Zhang W, Fan C, Yu Y, Ma J, SanMiguel P, Jiang N, Cheng Z, Wing RA, Jiang J, Jackson SA (2009) A lineage-specific centromere retrotransposon in Oryza brachyantha. Plant J 60:820–831PubMedCrossRefGoogle Scholar
  22. Goff SA, Ricke D, Lan TH, Presting G, Wang RL, Dunn M, Glazebrook J, Sessions A, Oeller P, Varma H, Hadley D, Hutchinson D, Martin C, Katagiri F, Lange BM, Moughamer T, Xia Y, Budworth P, Zhong JP, Miguel T, Paszkowski U, Zhang SP, Colbert M, Sun WL, Chen LL, Cooper B, Park S, Wood TC, Mao L, Quail P, Wing R, Dean R, Yu YS, Zharkikh A, Shen R, Sahasrabudhe S, Thomas A, Cannings R, Gutin A, Pruss D, Reid J, Tavtigian S, Mitchell J, Eldredge G, Scholl T, Miller RM, Bhatnagar S, Adey N, Rubano T, Tusneem N, Robinson R, Feldhaus J, Macalma T, Oliphant A, Briggs S (2002) A draft sequence of the rice genome (Oryza sativa L. ssp japonica). Science 296:92–100PubMedCrossRefGoogle Scholar
  23. Gong Z, Yu H, Huang J, Yi C, Gu M (2009) Unstable transmission of rice chromosomes without functional centromeric repeats in asexual propagation. Chromosome Res 17:863–872PubMedCrossRefGoogle Scholar
  24. Guo Y-L, Ge S (2005) Molecular phylogeny of Oryzeae (Poaceae) based on DNA sequences from chloroplast, mitochondrial, and nuclear genomes. Am J Bot 92:1548–1558PubMedCrossRefGoogle Scholar
  25. Hayashi T, Fujita Y, Iwasaki O, Adachi Y, Takahashi K, Yanagida M (2004) Mis16 and Mis18 are required for CENP-A loading and histone deacetylation at centromeres. Cell 118:715–729PubMedCrossRefGoogle Scholar
  26. Hemmerich P, Weidtkamp-Peters S, Hoischen C, Schmiedeberg L, Erliandri I, Diekmann S (2008) Dynamics of inner kinetochore assembly and maintenance in living cells. J Cell Biol 180:1101–1114PubMedCrossRefGoogle Scholar
  27. Henikoff S, Ahmad K, Platero JS, van Steensel B (2000) Heterochromatic deposition of centromeric histone H3-like proteins. Proc Natl Acad Sci USA 97:716–721PubMedCrossRefGoogle Scholar
  28. Henikoff S, Ahmad K, Malik HS (2001) The centromere paradox: stable inheritance with rapidly evolving DNA. Science 293:1098–1102PubMedCrossRefGoogle Scholar
  29. Heslop-Harrison JS, Murata M, Ogura Y, Schwarzacher T, Motoyoshi F (1999) Polymorphisms and genomic organization of repetitive DNA from centromeric regions of Arabidopsis chromosomes. Plant Cell 11:31–42PubMedCrossRefGoogle Scholar
  30. Heslop-Harrison JS, Brandes A, Schwarzacher T (2003) Tandemly repeated DNA sequences and centromeric chromosomal regions of Arabidopsis species. Chromosome Res 11:241–253PubMedCrossRefGoogle Scholar
  31. Hirsch CD, Wu YF, Yan HH, Jiang JM (2009) Lineage-specific adaptive evolution of the centromeric protein CENH3 in diploid and allotetraploid Oryza species. Mol Biol Evol 26:2877–2885PubMedCrossRefGoogle Scholar
  32. Houben A, Schroeder-Reiter E, Nagaki K, Nasuda S, Wanner G, Murata M, Endo TR (2007) CENH3 interacts with the centromeric retrotransposon cereba and GC-rich satellites and locates to centromeric substructures in barley. Chromosoma 116:275–283PubMedCrossRefGoogle Scholar
  33. Hudakova S, Michalek W, Presting GG, Rt H, Kd S, Jasencakova Z, Schubert I (2001) Sequence organization of barley centromeres. Nucleic Acids Res 29:5029–5035PubMedCrossRefGoogle Scholar
  34. Jansen LET, Black BE, Foltz DR, Cleveland DW (2007) Propagation of centromeric chromatin requires exit from mitosis. J Cell Biol 176:795–805PubMedCrossRefGoogle Scholar
  35. Jiang JM, Birchler JA, Parrott WA, Dawe RK (2003) A molecular view of plant centromeres. Trends Plant Sci 8:570–575PubMedCrossRefGoogle Scholar
  36. Jin WW, Melo JR, Nagaki K, Talbert PB, Henikoff S, Dawe RK, Jiang JM (2004) Maize centromeres: organization and functional adaptation in the genetic background of oat. Plant Cell 16:571–581PubMedCrossRefGoogle Scholar
  37. Jin W, Lamb JC, Vega JM, Dawe RK, Birchler JA, Jiang J (2005) Molecular and functional dissection of the maize B chromosome centromere. Plant Cell 17:1412–1423PubMedCrossRefGoogle Scholar
  38. Jin W, Lamb J, Zhang W, Kolano B, Birchler J, Jiang J (2008) Histone modifications associated with both A and B chromosomes of maize. Chromosome Res 16:1203–1214PubMedCrossRefGoogle Scholar
  39. Kamm A, Galasso I, Schmidt T, Heslop-Harrison JS (1995) Analysis of a repetitive DNA family from Arabidopsis arenosa and relationships between Arabidopsis species. Plant Mol Biol 27:853–862PubMedCrossRefGoogle Scholar
  40. Kaul S, Koo HL, Jenkins J, Rizzo M, Rooney T, Tallon LJ, Feldblyum T, Nierman W, Benito MI, Lin XY, Town CD, Venter JC, Fraser CM, Tabata S, Nakamura Y, Kaneko T, Sato S, Asamizu E, Kato T, Kotani H, Sasamoto S, Ecker JR, Theologis A, Federspiel NA, Palm CJ, Osborne BI, Shinn P, Conway AB, Vysotskaia VS, Dewar K, Conn L, Lenz CA, Kim CJ, Hansen NF, Liu SX, Buehler E, Altafi H, Sakano H, Dunn P, Lam B, Pham PK, Chao Q, Nguyen M, Yu GX, Chen HM, Southwick A, Lee JM, Miranda M, Toriumi MJ, Davis RW, Wambutt R, Murphy G, Dusterhoft A, Stiekema W, Pohl T, Entian KD, Terryn N, Volckaert G, Salanoubat M, Choisne N, Rieger M, Ansorge W, Unseld M, Fartmann B, Valle G, Artiguenave F, Weissenbach J, Quetier F, Wilson RK, de la Bastide M, Sekhon M, Huang E, Spiegel L, Gnoj L, Pepin K, Murray J, Johnson D, Habermann K, Dedhia N, Parnell L, Preston R, Hillier L, Chen E, Marra M, Martienssen R, McCombie WR, Mayer K, White O, Bevan M, Lemcke K, Creasy TH, Bielke C, Haas B, Haase D, Maiti R, Rudd S, Peterson J, Schoof H, Frishman D, Morgenstern B, Zaccaria P, Ermolaeva M, Pertea M, Quackenbush J, Volfovsky N, Wu DY, Lowe TM, Salzberg SL, Mewes HW, Rounsley S, Bush D, Subramaniam S, Levin I, Norris S, Schmidt R, Acarkan A, Bancroft I, Brennicke A, Eisen JA, Bureau T, Legault BA, Le QH, Agrawal N, Yu Z, Copenhaver GP, Luo S, Pikaard CS, Preuss D, Paulsen IT, Sussman M, Britt AB, Selinger DA, Pandey R, Mount DW, Chandler VL, Jorgensen RA, Pikaard C, Juergens G, Meyerowitz EM, Dangl J, Jones JDG, Chen M, Chory J, Somerville MC, Ar Gen I (2000) Analysis of the genome sequence of the flowering plant Arabidopsis thaliana. Nature 408:796–815CrossRefGoogle Scholar
  41. Kawabe A, Nasuda S (2005) Structure and genomic organization of centromeric repeats in Arabidopsis species. Mol Genet Genomics 272:593–602PubMedCrossRefGoogle Scholar
  42. Kejnovsky E, Hawkins J, Feschotte C (2012) Plant transposable elements: biology and evolution. In: Wendel JF (ed) Plant genome diversity, vol 1, Plant genomes, their residents, and their evolutionary dynamics. Springer, Wien, New YorkGoogle Scholar
  43. Kikuchi S, Kishii M, Shimizu M, Tsujimoto H (2005) Centromere-specific repetitive sequences from Torenia, a model plant for interspecific fertilization, and whole-mount FISH of its interspecific hybrid embryos. Cytogenet Genome Res 109:228–235PubMedCrossRefGoogle Scholar
  44. Koo D-H, Jiang J (2009) Super-stretched pachytene chromosomes for fluorescence in situ hybridization mapping and immunodetection of DNA methylation. Plant J 59:509–516PubMedCrossRefGoogle Scholar
  45. Kordis D (2005) A genomic perspective on the chromodomain-containing retrotransposons: chromoviruses. Gene 347:161–173PubMedCrossRefGoogle Scholar
  46. Kumekawa N, Hosouchi T, Tsuruoka H, Kotani H (2000) The size and sequence organization of the centromeric region of Arabidopsis thaliana chromosome 5. DNA Res 7:315–321PubMedCrossRefGoogle Scholar
  47. Kumekawa N, Hosouchi T, Tsuruoka H, Kotani H (2001) The size and sequence organization of the centromeric region of Arabidopsis thaliana chromosome 4. DNA Res 8:285–290PubMedCrossRefGoogle Scholar
  48. Langdon T, Seago C, Mende M, Leggett M, Thomas H, Forster JW, Thomas H, Jones RN, Jenkins G (2000) Retrotransposon evolution in diverse plant genomes. Genetics 156:313–325PubMedGoogle Scholar
  49. Lee HR, Zhang WL, Langdon T, Jin WW, Yan HH, Cheng ZK, Jiang JM (2005) Chromatin immunoprecipitation cloning reveals rapid evolutionary patterns of centromeric DNA in Oryza species. Proc Natl Acad Sci USA 102:11793–11798PubMedCrossRefGoogle Scholar
  50. Lee HR, Neumann P, Macas J, Jiang JM (2006) Transcription and evolutionary dynamics of the centromeric satellite repeat CentO in rice. Mol Biol Evol 23:2505–2520PubMedCrossRefGoogle Scholar
  51. Lermontova I, Schubert V, Fuchs J, Klatte S, Macas J, Schubert I (2006) Loading of Arabidopsis centromeric histone CENH3 occurs mainly during G2 and requires the presence of the histone fold domain. Plant Cell 18:2443–2451PubMedCrossRefGoogle Scholar
  52. Lermontova I, Fuchs Jr, Schubert V, Schubert I (2007) Loading time of the centromeric histone H3 variant differs between plants and animals. Chromosoma 116:507–510PubMedCrossRefGoogle Scholar
  53. .01w?>Liu Z, Yue W, Li D, Wang R, Kong X, Lu K, Wang G, Dong Y, Jin W, Zhang X (2008) Structure and dynamics of retrotransposons at wheat centromeres and pericentromeres. Chromosoma 117:445–456PubMedCrossRefGoogle Scholar
  54. Malik HS, Henikoff S (2001) Adaptive evolution of cid, a centromere-specific histone in drosophila. Genetics 157:1293–1298PubMedGoogle Scholar
  55. Malik HS, Henikoff S (2002) Conflict begets complexity: the evolution of centromeres. Curr Opin Genet Dev 12:711–718PubMedCrossRefGoogle Scholar
  56. Maluszynska J, Heslop-Harrison J (1991) Localization of tandemly repeated DMA sequences in Arabidopsis thaliana. Plant J 1:159–166CrossRefGoogle Scholar
  57. Marshall OJ, Chueh AC, Wong LH, Choo KHA (2008) Neocentromeres: new insights into centromere structure, disease development, and karyotype evolution. Am J Hum Genet 82:261–282PubMedCrossRefGoogle Scholar
  58. Martinez-Zapater JM, Estelle MA, Somerville CR (1986) A highly repeated DNA sequence in Arabidopsis thaliana. Mol Gen Genet 204:417–423CrossRefGoogle Scholar
  59. Meluh PB, Yang PR, Glowczewski L, Koshland D, Smith MM (1998) Cse4p is a component of the core centromere of Saccharomyces cerevisiae. Cell 94:607–613PubMedCrossRefGoogle Scholar
  60. Miller JT, Dong F, Jackson SA, Song J, Jiang J (1998) Retrotransposon-related DNA sequences in the centromeres of grass chromosomes. Genetics 150:1615–1623PubMedGoogle Scholar
  61. Murata M, Ogura Y, Motoyoshi F (1994) Centromeric repetitive sequences in Arabidopsis thaliana. Jap J Genet 69:361–370CrossRefGoogle Scholar
  62. Nagaki K, Murata M (2005) Characterization of CENH3 and centromere-associated DNA sequences in sugarcane. Chromosome Res 13:195–203PubMedCrossRefGoogle Scholar
  63. Nagaki K, Talbert PB, Zhong CX, Dawe RK, Henikoff S, Jiang JM (2003) Chromatin immunoprecipitation reveals that the 180-bp satellite repeat is the key functional DNA element of Arabidopsis thaliana centromeres. Genetics 163:1221–1225PubMedGoogle Scholar
  64. Nagaki K, Cheng ZK, Ouyang S, Talbert PB, Kim M, Jones KM, Henikoff S, Buell CR, Jiang JM (2004) Sequencing of a rice centromere uncovers active genes. Nat Genet 36:138–145PubMedCrossRefGoogle Scholar
  65. Nagaki K, Neumann P, Zhang D, Ouyang S, Buell CR, Cheng Z, Jiang J (2005) Structure, divergence, and distribution of the CRR centromeric retrotransposon family in rice. Mol Biol Evol 22:845–855PubMedCrossRefGoogle Scholar
  66. Nagaki K, Kashihara K, Murata M (2009) A centromeric DNA sequence colocalized with a centromere-specific histone H3 in tobacco. Chromosoma 118:249–257PubMedCrossRefGoogle Scholar
  67. Nasuda S, Hudakova S, Schubert I, Houben A, Endo TR (2005) Stable barley chromosomes without centromeric repeats. Proc Natl Acad Sci USA 102:9842–9847PubMedCrossRefGoogle Scholar
  68. Neumann P, Yan HH, Jiang JM (2007) The centromeric retrotransposons of rice are transcribed and differentially processed by RNA interference. Genetics 176:749–761PubMedCrossRefGoogle Scholar
  69. Nonomura KI, Kurata N (1999) Organization of the 1.9-kb repeat unit RCE1 in the centromeric region of rice chromosomes. Mol Gen Genet MGG 261:1–10CrossRefGoogle Scholar
  70. Okada M, Cheeseman IM, Hori T, Okawa K, McLeod IX, Yates JR, 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 8:446–457PubMedCrossRefGoogle Scholar
  71. Pidoux AL, Choi ES, Abbott JKR, Liu X, Kagansky A, Castillo AG, Hamilton GL, Richardson W, Rappsilber J, He X, Allshire RC (2009) Fission yeast Scm3: a CENP-A receptor required for integrity of subkinetochore chromatin. Mol Cell 33:299–311PubMedCrossRefGoogle Scholar
  72. Presting GG, Malysheva L, Fuchs J, Schubert I (1998) ATY3/GYPSYretrotransposon-like sequence localizes to the centromeric regions of cereal chromosomes. Plant J 16:721–728PubMedCrossRefGoogle Scholar
  73. Ravi M, Kwong PN, Menorca RMG, Valencia JT, Ramahi JS, Stewart JL, Tran RK, Sundaresan V, Comai L, Chan SW-L (2010) The rapidly evolving centromere-specific histone has stringent functional requirements in Arabidopsis thaliana. Genetics: genetics.110.120337Google Scholar
  74. Shi J, Dawe RK (2006) Partitioning of the maize epigenome by the number of methyl groups on histone H3 lysines 9 and 27. Genetics 173:1571–1583PubMedCrossRefGoogle Scholar
  75. Shibata F, Murata M (2004) Differential localization of the centromere-specific proteins in the major centromeric satellite of Arabidopsis thaliana. J Cell Sci 117:2963–2970PubMedCrossRefGoogle Scholar
  76. Shuaib M, Ouararhni K, Dimitrov S, Hamiche A (2010) HJURP binds CENP-A via a highly conserved N-terminal domain and mediates its deposition at centromeres. Proc Natl Acad Sci USA 107:1349–1354PubMedCrossRefGoogle Scholar
  77. Slotkin R, Nuthikattu S, Jiang N (2012) The impact of transposable elements on gene and genome evolution. In: Wendel JF (ed) Plant genome diversity, vol 1, Plant genomes, their residents, and their evolutionary dynamics. Springer, Wien, New YorkGoogle Scholar
  78. Stoler S, Keith KC, Curnick KE, Fitzgerald-Hayes M (1995) A mutation in CSE4, an essential gene encoding a novel chromatin-associated protein in yeast, causes chromosome nondisjunction and cell cycle arrest at mitosis. Genes Dev 9:573–586PubMedCrossRefGoogle Scholar
  79. Sullivan BA, Karpen GH (2004) Centromeric chromatin exhibits a histone modification pattern that is distinct from both euchromatin and heterochromatin. Nat Struct Mol Biol 11:1076–1083PubMedCrossRefGoogle Scholar
  80. Sullivan KF, Hechenberger M, Masri K (1994) Human CENP-A contains a histone H3 related histone fold domain that is required for targeting to the centromere. J Cell Biol 127:581–592PubMedCrossRefGoogle Scholar
  81. Talbert PB, Masuelli R, Tyagi AP, Comai L, Henikoff S (2002) Centromeric localization and adaptive evolution of an Arabidopsis histone H3 variant. Plant Cell 14:1053–1066PubMedCrossRefGoogle Scholar
  82. Talbert P, Bryson T, Henikoff S (2004) Adaptive evolution of centromere proteins in plants and animals. J Biol 3:18PubMedCrossRefGoogle Scholar
  83. Tek AL, Jiang J (2004) The centromeric regions of potato chromosomes contain megabase-sized tandem arrays of telomere-similar sequence. Chromosoma 113:77–83PubMedCrossRefGoogle Scholar
  84. Tek A, Kashihara K, Murata M, Nagaki K (2010) Functional centromeres in soybean include two distinct tandem repeats and a retrotransposon. Chromosome Res 18:337–347PubMedCrossRefGoogle Scholar
  85. Topp CN, Zhong CX, Dawe RK (2004) Centromere-encoded RNAs are integral components of the maize kinetochore. Proc Natl Acad Sci USA 101:15986–15991PubMedCrossRefGoogle Scholar
  86. Topp CN, Okagaki RJ, Melo JR, Kynast RG, Phillips RL, Dawe RK (2009) Identification of a maize neocentromere in an oat-maize addition line. Cytogenet Genome Res 124:228–238PubMedCrossRefGoogle Scholar
  87. Torras-Llort M, Moreno-Moreno O, Azorin F (2009) Focus on the centre: the role of chromatin on the regulation of centromere identity and function. EMBO J 28:2337–2348PubMedCrossRefGoogle Scholar
  88. Williams JS, Hayashi T, Yanagida M, Russell P (2009) Fission yeast Scm3 mediates stable assembly of Cnp1/CENP-A into centromeric chromatin. Mol Cell 33:287–298PubMedCrossRefGoogle Scholar
  89. Wong LH, Brettingham-Moore KH, Chan L, Quach JM, Anderson MA, Northrop EL, Hannan R, Saffery R, Shaw ML, Williams E, Choo KHA (2007) Centromere RNA is a key component for the assembly of nucleoproteins at the nucleolus and centromere. Genome Res 17:1146–1160PubMedCrossRefGoogle Scholar
  90. Yan HH, Ito H, Nobuta K, Ouyang S, Jin WW, Tian SL, Lu C, Venu RC, Wang GL, Green PJ, Wing RA, Buell CR, Meyers BC, Jiang JM (2006) Genomic and genetic characterization of rice Cen3 reveals extensive transcription and evolutionary implications of a complex centromere. Plant Cell 18:2123–2133PubMedCrossRefGoogle Scholar
  91. Yan H, Talbert PB, Lee H-R, Jett J, Henikoff S, Chen F, Jiang J (2008) Intergenic locations of rice centromeric chromatin. PLoS Biol 6:e286PubMedCrossRefGoogle Scholar
  92. Yang C, Tomkiel J, Saitoh H, Johnson D, Earnshaw W (1996) Identification of overlapping DNA-binding and centromere-targeting domains in the human kinetochore protein CENP-C. Mol Cell Biol 16:3576–3586PubMedGoogle Scholar
  93. Yu J, Hu SN, Wang J, Wong GKS, Li SG, Liu B, Deng YJ, Dai L, Zhou Y, Zhang XQ, Cao ML, Liu J, Sun JD, Tang JB, Chen YJ, Huang XB, Lin W, Ye C, Tong W, Cong LJ, Geng JN, Han YJ, Li L, Li W, Hu GQ, Huang XG, Li WJ, Li J, Liu ZW, Liu JP, Qi QH, Liu JS, Li T, Wang XG, Lu H, Wu TT, Zhu M, Ni PX, Han H, Dong W, Ren XY, Feng XL, Cui P, Li XR, Wang H, Xu X, Zhai WX, Xu Z, Zhang JS, He SJ, Zhang JG, Xu JC, Zhang KL, Zheng XW, Dong JH, Zeng WY, Tao L, Ye J, Tan J, Ren XD, Chen XW, He J, Liu DF, Tian W, Tian CG, Xia HG, Bao QY, Li G, Gao H, Cao T, Zhao WM, Li P, Chen W, Wang XD, Zhang Y, Hu JF, Liu S, Yang J, Zhang GY, Xiong YQ, Li ZJ, Mao L, Zhou CS, Zhu Z, Chen RS, Hao BL, Zheng WM, Chen SY, Guo W, Li GJ, Liu SQ, Tao M, Zhu LH, Yuan LP, Yang HM (2002) A draft sequence of the rice genome (Oryza sativa L. ssp. indica). Science 296:79–92PubMedCrossRefGoogle Scholar
  94. Zappulla DC, Cech TR (2004) Yeast telomerase RNA: a flexible scaffold for protein subunits. Proc Natl Acad Sci USA 101:10024–10029PubMedCrossRefGoogle Scholar
  95. Zhang L-B, Ge S (2007) Multilocus analysis of nucleotide variation and speciation in Oryza officinalis and its close relatives. Mol Biol Evol 24:769–783PubMedCrossRefGoogle Scholar
  96. Zhang W, Lee H-R, Koo D-H, Jiang J (2008) Epigenetic modification of centromeric chromatin: hypomethylation of DNA sequences in the CENH3-associated chromatin in Arabidopsis thaliana and maize. Plant Cell 20:25–34PubMedCrossRefGoogle Scholar
  97. Zhong CX, Marshall JB, Topp C, Mroczek R, Kato A, Nagaki K, Birchler JA, Jiang JM, Dawe RK (2002) Centromeric retroelements and satellites interact with maize kinetochore protein CENH3. Plant Cell 14:2825–2836PubMedCrossRefGoogle Scholar

Copyright information

© Springer Verlag Wien 2012

Authors and Affiliations

  1. 1.Department of HorticultureUniversity of Wisconsin-MadisonMadisonUSA
  2. 2.Department of HorticultureUniversity of Wisconsin-MadisonMadisonUSA

Personalised recommendations