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CENP-A and centromere evolution in equids.

Abstract: While the centromeric function is conserved and epigenetically specified by CENP-A, centromeric DNA, typically composed of satellite repeats, is highly divergent and rapidly evolving. In the species of the genus Equus (horses, asses and zebras), also known as equids, the numerous centromeres devoid of satellite repeats enabled us to carry out molecular analysis of centromeric chromatin establishing a unique model system for mammalian centromere biology. In this review, after a brief description of the rapid evolution of equids, we outline one of our most relevant initial discoveries: the position of CENP-A binding domains is variable among individuals giving rise to epialleles which are inherited as Mendelian traits. This positional variability was recently confirmed in human centromeres whose repetitive DNA organization could be analyzed thanks to telomere-to-telomere (T2T) genome assemblies. Another unexpected observation was that, in equids, CENP-B does not bind the centromeric core and is uncoupled from CENP-A and CENP-C. CENP-B is absent from the majority of chromosomes while the CENP-B binding DNA sequence (CENP-B box) is comprised within a satellite that is localized at pericentromeric or terminal positions. Finally, comparative molecular and cytogenetic analyses of satellite-free centromeres revealed that the birth of neocentromeres during the evolution of this genus occurred through two alternative mechanisms: centromere repositioning and Robertsonian fusion. These events played a key role in karyotype reshuffling and speciation. Investigating centromere organization in equids provided new insights into the complexity of centromere organization across the vast biodiversity of the mammalian world, where the majority of species remain understudied.
Publication Date: 2025-06-30 PubMed ID: 40586953PubMed Central: PMC12208984DOI: 10.1007/s10577-025-09773-3Google Scholar: Lookup
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Summary

This research summary has been generated with artificial intelligence and may contain errors and omissions. Refer to the original study to confirm details provided. Submit correction.

The research article is a molecular review focused on centromeres, specifically their diversity and rapid evolution in different species of the Equus genus (horses, asses, and zebras). The study highlights key discoveries about the positional variability of CENP-A binding domains and their inheritance as Mendelian traits, and CENP-B’s uncoupling from CENP-A and CENP-C.

Centromeres and the Equus Genus

  • The study focuses on the varied and rapidly evolving centromeric DNA in the species from the Equus genus. These species have numerous centromeres devoid of satellite repeats which allowed for an in-depth molecular analysis.
  • On top of studying the rapid evolution of equids, the researchers also established these animals as a unique model system for the study of mammalian centromere biology.

The Positional Variability of CENP-A Binding Domains

  • One significant finding from the research is the discovery that the position of CENP-A binding domains can vary among individuals.
  • This variation results in different epialleles being created, and these are inherited as Mendelian traits (traits that follow specific patterns of inheritance).
  • Recent research in human centromeres confirmed this variability, thanks to ‘telomere-to-telomere’ genome assemblies that allowed the analysis of repetitive DNA organization.

CENP-B’s Uncoupling from CENP-A and CENP-C

  • The research also uncovered a surprise in equid centromeres: CENP-B doesn’t bind to the centromeric core and is instead uncoupled from CENP-A and CENP-C.
  • Most chromosomes are missing CENP-B, while the DNA sequence it can bind to (the CENP-B box) is found inside a satellite at pericentromeric or terminal positions.

Neocentromeres and Karyotype Reshuffling

  • Finally, when conducting a comparative molecular and cytogenetic analysis of satellite-free centromeres, they found two alternative mechanisms that created neocentromeres (new centromeres): centromere repositioning and Robertsonian fusion.
  • These events played a crucial role in the reshuffling of karyotypes (the number and visual appearance of chromosomes in the cell nuclei) and speciation.

Implications and Conclusion

  • This research adds valuable information to our understanding of centromere organization in equids, which may have wider implications for the study of diverse mammals, as many species remain understudied.
  • This research gains importance given that centromeres play a crucial role in cell division, therefore understanding their organization and functioning could have significant implications for understanding species evolution and potentially tackling diseases related to cell division.

Cite This Article

APA
(2025). CENP-A and centromere evolution in equids. Chromosome Res, 33(1), 13. https://doi.org/10.1007/s10577-025-09773-3

Publication

ISSN: 1573-6849
NlmUniqueID: 9313452
Country: Netherlands
Language: English
Volume: 33
Issue: 1
Pages: 13
PII: 13

Researcher Affiliations

MeSH Terms

  • Animals
  • Centromere / genetics
  • Centromere / metabolism
  • Centromere Protein A / genetics
  • Equidae / genetics
  • Evolution, Molecular
  • DNA, Satellite / genetics
  • Humans
  • Horses / genetics
  • Chromosomal Proteins, Non-Histone / genetics
  • Chromosomal Proteins, Non-Histone / metabolism
  • Centromere Protein B / genetics

Grant Funding

  • 2019-67015-29340/Project Accession 1018854 / National Institute of Food and Agriculture
  • 2019-67015-29340/Project Accession 1018854 / National Institute of Food and Agriculture
  • 2019-67015-29340/Project Accession 1018854 / National Institute of Food and Agriculture
  • 2019-67015-29340/Project Accession 1018854 / National Institute of Food and Agriculture
  • 2019-67015-29340/Project Accession 1018854 / National Institute of Food and Agriculture
  • 2019-67015-29340/Project Accession 1018854 / National Institute of Food and Agriculture
  • 2019-67015-29340/Project Accession 1018854 / National Institute of Food and Agriculture
  • 2019-67015-29340/Project Accession 1018854 / National Institute of Food and Agriculture
  • 2019-67015-29340/Project Accession 1018854 / National Institute of Food and Agriculture

Conflict of Interest Statement

Declarations. Ethics approval: Not applicable. Consent to publish: Not applicable. Competing interests: The authors declare no competing interests.

References

This article includes 105 references
  1. Altemose N, Logsdon GA, Bzikadze AV, Sidhwani P, Langley SA, Caldas GV, Hoyt SJ, Uralsky L, Ryabov FD, Shew CJ. Complete genomic and epigenetic maps of human centromeres. Science 376:eabl4178.
    pmc: PMC9233505pubmed: 35357911
  2. Amor DJ, Choo KH. Neocentromeres: role in human disease, evolution, and centromere study. Am J Hum Genet 71(4):695–714.
    pmc: PMC378529pubmed: 12196915
  3. Anglana M, Bertoni L, Giulotto E. Cloning of a polymorphic sequence from the nontranscribed spacer of horse rDNA. Mamm Genome 7:539–541.
    pubmed: 8672135
  4. Arora UP, Sullivan BA, Dumont BL. Variation in the CENP-A sequence association landscape across diverse inbred mouse strains. Cell Rep 42:113178.
    pmc: PMC10873113pubmed: 37742188
  5. Barbosa AC, Xu Z, Karari K, Williams W, Hauf S, Brown WRA. Mutation and selection explain why many eukaryotic centromeric DNA sequences are often A + T rich. Nucleic Acids Res 50:579–596.
    pmc: PMC8754631pubmed: 34928384
  6. Brannan EO, Hartley GA, O’Neill RJ. Mechanisms of rapid karyotype evolution in mammals. Genes (Basel) .
    pmc: PMC10815390pubmed: 38254952doi: 10.3390/genes15010062google scholar: lookup
  7. Cappelletti E, Piras FM, Badiale C, Bambi M, Santagostino M, Vara C, Masterson TA, Sullivan KF, Nergadze SG, Ruiz-Herrera A. CENP-A binding domains and recombination patterns in horse spermatocytes. Sci Rep 9:15800.
    pmc: PMC6825197pubmed: 31676881
  8. Cappelletti E, Piras FM, Sola L, Santagostino M, Abdelgadir WA, Raimondi E, Lescai F, Nergadze SG, Giulotto E. Robertsonian fusion and centromere repositioning contributed to the formation of satellite-free centromeres during the evolution of zebras. Mol Biol Evol .
    pmc: PMC9356731pubmed: 35881460doi: 10.1093/molbev/msac162google scholar: lookup
  9. Cappelletti E, Piras FM, Sola L, Santagostino M, Petersen JL, Bellone RR, Finno CJ, Peng S, Kalbfleisch TS, Bailey E. The localization of centromere protein A is conserved among tissues. Commun Biol 6:963.
    pmc: PMC10514049pubmed: 37735603
  10. Cappelletti E, Piras FM, Biundo M, Raimondi E, Nergadze SG, Giulotto E. CENP-A/CENP-B uncoupling in the evolutionary reshuffling of centromeres in equids. Genome Biol .
    pmc: PMC11804003pubmed: 39915813doi: 10.1186/s13059-025-03490-0google scholar: lookup
  11. Carbone L, Nergadze SG, Magnani E, Misceo D, Francesca Cardone M, Roberto R, Bertoni L, Attolini C, Francesca Piras M, de Jong P. Evolutionary movement of centromeres in horse, donkey, and zebra. Genomics 87:777–782.
    pubmed: 16413164
  12. Cerutti F, Gamba R, Mazzagatti A, Piras FM, Cappelletti E, Belloni E, Nergadze SG, Raimondi E, Giulotto E. The major horse satellite DNA family is associated with centromere competence. Mol Cytogenet 9:35.
    pmc: PMC4847189pubmed: 27123044
  13. Chabot BJ, Sun R, Amjad A, Hoyt SJ, Ouyang L, Courret C, Drennan R, Leo L, Larracuente AM, Core LJ. Transcription of a centromere-enriched retroelement and local retention of its RNA are significant features of the CENP-A chromatin landscape. Genome Biol 25:295.
    pmc: PMC11575011pubmed: 39558354
  14. Chang CH, Chavan A, Palladino J, Wei X, Martins NMC, Santinello B, Chen CC, Erceg J, Beliveau BJ, Wu CT. Islands of retroelements are major components of Drosophila centromeres. PLoS Biol 17:e3000241.
    pmc: PMC6516634pubmed: 31086362
  15. Chmátal L, Gabriel SI, Mitsainas GP, Martínez-Vargas J, Ventura J, Searle JB, Schultz RM, Lampson MA. Centromere strength provides the cell biological basis for meiotic drive and karyotype evolution in mice. Curr Biol 24:2295–2300.
    pmc: PMC4189972pubmed: 25242031
  16. Choo KH. Why is the centromere so cold?. Genome Res 8:81–82.
    pubmed: 9477334
  17. Choo KH. Centromerization. Trends Cell Biol 10:182–188.
    pubmed: 10754560
  18. Clarke L, Carbon J. The structure and function of yeast centromeres. Annu Rev Genet 19:29–55.
    pubmed: 3909945
  19. Dudka D, Nguyen AL, Boese KG, Marescal O, Akins RB, Black BE, Cheeseman IM, Lampson MA. Adaptive evolution of CENP-T modulates centromere binding. Curr Biol 35:1012-1022.e1015.
    pmc: PMC11903153pubmed: 39947176
  20. Dumont M, Gamba R, Gestraud P, Klaasen S, Worrall JT, De Vries SG, Boudreau V, Salinas-Luypaert C, Maddox PS, Lens SM. Human chromosome-specific aneuploidy is influenced by DNA-dependent centromeric features. EMBO J 39:e102924.
    pmc: PMC6960447pubmed: 31750958
  21. Fachinetti D, Han JS, McMahon MA, Ly P, Abdullah A, Wong AJ, Cleveland DW. DNA sequence-specific binding of CENP-B enhances the fidelity of human centromere function. Dev Cell 33:314–327.
    pmc: PMC4421092pubmed: 25942623
  22. Fishman L, Saunders A. Centromere-associated female meiotic drive entails male fitness costs in monkeyflowers. Science 322:1559–1562.
    pubmed: 19056989
  23. Fowler KJ, Hudson DF, Salamonsen LA, Edmondson SR, Earle E, Sibson MC, Choo KH. Uterine dysfunction and genetic modifiers in centromere protein B-deficient mice. Genome Res 10:30–41.
    pmc: PMC310504pubmed: 10645947
  24. Froehlich DJ. Quo vadis eohippus? The systematics and taxonomy of the early Eocene equids (Perissodactyla). Zool J Linnean Soc 134:141–256.
  25. Gamba R, Fachinetti D. From evolution to function: two sides of the same CENP-B coin?. Exp Cell Res 390:111959.
    pubmed: 32173469
  26. Garagna S, Marziliano N, Zuccotti M, Searle JB, Capanna E, Redi CA. Pericentromeric organization at the fusion point of mouse Robertsonian translocation chromosomes. Proc Natl Acad Sci U S A 98:171–175.
    pmc: PMC14563pubmed: 11136254
  27. Garagna S, Page J, Fernandez-Donoso R, Zuccotti M, Searle JB. The Robertsonian phenomenon in the house mouse: mutation, meiosis and speciation. Chromosoma 123:529–544.
    pubmed: 25053180
  28. Giulotto E, Raimondi E, Sullivan KF. The Unique DNA Sequences Underlying Equine Centromeres. Prog Mol Subcell Biol 56:337–354.
    pubmed: 28840244
  29. Hartley GA, Okhovat M, Hoyt SJ, Fuller E, Pauloski N, Alexandre N, Alexandrov I, Drennan R, Dubocanin D, Gilbert DM. Centromeric transposable elements and epigenetic status drive karyotypic variation in the eastern hoolock gibbon. Cell Genom .
    pmc: PMC12008813pubmed: 40088887doi: 10.1016/j.xgen.2025.100808google scholar: lookup
  30. Hasson D, Panchenko T, Salimian KJ, Salman MU, Sekulic N, Alonso A, Warburton PE, Black BE. The octamer is the major form of CENP-A nucleosomes at human centromeres. Nat Struct Mol Biol 20:687–695.
    pmc: PMC3760417pubmed: 23644596
  31. Henikoff S, Ahmad K, Malik HS. The centromere paradox: stable inheritance with rapidly evolving DNA. Science 293:1098–1102.
    pubmed: 11498581
  32. Heslop-Harrison JS, Schwarzacher T. Nucleosomes and centromeric DNA packaging. Proc Natl Acad Sci U S A 110:19974–19975.
    pmc: PMC3864337pubmed: 24282300
  33. Hoffmann S, Dumont M, Barra V, Ly P, Nechemia-Arbely Y, McMahon MA, Hervé S, Cleveland DW, Fachinetti D. CENP-A is dispensable for mitotic centromere function after initial centromere/kinetochore assembly. Cell Rep 17:2394–2404.
    pmc: PMC5134894pubmed: 27880912
  34. Hori T, Kagawa N, Toyoda A, Fujiyama A, Misu S, Monma N, Makino F, Ikeo K, Fukagawa T. Constitutive centromere-associated network controls centromere drift in vertebrate cells. J Cell Biol 216:101–113.
    pmc: PMC5223601pubmed: 27940888
  35. Houck ML, Kingswood SC, Kumamoto AT. Comparative cytogenetics of tapirs, genus Tapirus (Perissodactyla, Tapiridae). Cytogenet Cell Genet 89:110–115.
    pubmed: 10894950
  36. Hudson DF, Fowler KJ, Earle E, Saffery R, Kalitsis P, Trowell H, Hill J, Wreford NG, de Kretser DM, Cancilla MR. Centromere protein B null mice are mitotically and meiotically normal but have lower body and testis weights. J Cell Biol 141:309–319.
    pmc: PMC2148459pubmed: 9548711
  37. Iwata-Otsubo A, Dawicki-McKenna JM, Akera T, Falk SJ, Chmátal L, Yang K, Sullivan BA, Schultz RM, Lampson MA, Black BE. Expanded satellite repeats amplify a discrete CENP-A nucleosome assembly site on chromosomes that drive in female meiosis. Curr Biol 27:2365-2373.e2368.
    pmc: PMC5567862pubmed: 28756949
  38. Jónsson H, Schubert M, Seguin-Orlando A, Ginolhac A, Petersen L, Fumagalli M, Albrechtsen A, Petersen B, Korneliussen TS, Vilstrup JT. Speciation with gene flow in equids despite extensive chromosomal plasticity. Proc Natl Acad Sci U S A 111:18655–18660.
    pmc: PMC4284605pubmed: 25453089
  39. Kalitsis P, Griffiths B, Choo KH. Mouse telocentric sequences reveal a high rate of homogenization and possible role in Robertsonian translocation. Proc Natl Acad Sci U S A 103:8786–8791.
    pmc: PMC1482656pubmed: 16731628
  40. Kapoor M, de Oca M, Luna R, Liu G, Lozano G, Cummings C, Mancini M, Ouspenski I, Brinkley BR, May GS. The cenpB gene is not essential in mice. Chromosoma 107:570–576.
    pubmed: 9933410
  41. Kasinathan S, Henikoff S. Non-B-form DNA is enriched at centromeres. Mol Biol Evol 35:949–962.
    pmc: PMC5889037pubmed: 29365169
  42. Kipling D, Warburton PE. Centromeres, CENP-B and Tigger too. Trends Genet 13:141–145.
    pubmed: 9097724
  43. Klein SJ, O’Neill RJ. Transposable elements: genome innovation, chromosome diversity, and centromere conflict. Chromosome Res 26:5–23.
    pmc: PMC5857280pubmed: 29332159
  44. Kumon T, Ma J, Akins RB, Stefanik D, Nordgren CE, Kim J, Levine MT, Lampson MA. Parallel pathways for recruiting effector proteins determine centromere drive and suppression. Cell 184:4904-4918.e4911.
    pmc: PMC8448984pubmed: 34433012
  45. Kursel LE, Malik HS. The cellular mechanisms and consequences of centromere drive. Curr Opin Cell Biol 52:58–65.
    pmc: PMC5988936pubmed: 29454259
  46. Li H, Durbin R. Genome assembly in the telomere-to-telomere era. Nat Rev Genet 25:658–670.
    pubmed: 38649458
  47. Librado P, Orlando L. Genomics and the evolutionary history of equids. Annu Rev Anim Biosci 9(1):81–101.
    pubmed: 33197207
  48. Lindsay EH, Opdyke ND, Johnson NM. Pliocene dispersal of the horse Equus and late Cenozoic mammalian dispersal events. Nature 287:135–138.
  49. Locke DP, Hillier LW, Warren WC, Worley KC, Nazareth LV, Muzny DM, Yang SP, Wang Z, Chinwalla AT, Minx P. Comparative and demographic analysis of orang-utan genomes. Nature 469:529–533.
    pmc: PMC3060778pubmed: 21270892
  50. Logsdon GA, Gambogi CW, Liskovykh MA, Barrey EJ, Larionov V, Miga KH, Heun P, Black BE. Human Artificial Chromosomes that Bypass Centromeric DNA. Cell 178:624-639.e619.
    pmc: PMC6657561pubmed: 31348889
  51. Logsdon GA, Rozanski AN, Ryabov F, Potapova T, Shepelev VA, Catacchio CR, Porubsky D, Mao Y, Yoo D, Rautiainen M. The variation and evolution of complete human centromeres. Nature 629:136–145.
    pmc: PMC11062924pubmed: 38570684
  52. Longo MS, Carone DM, Green ED, O’Neill MJ, O’Neill RJ, Program NCS. Distinct retroelement classes define evolutionary breakpoints demarcating sites of evolutionary novelty. BMC Genomics 10:334.
    pmc: PMC2736999pubmed: 19630942
  53. Macfadden BJ. Evolution. Fossil horses–evidence for evolution. Science 307:1728–1730.
    pubmed: 15774746
  54. MacFadden BJ, Hulbert RC. Explosive speciation at the base of the adaptive radiation of Miocene grazing horses. Nature 336:466–468.
  55. Mahlke MA, Lumerman L, Ly P, Nechemia-Arbely Y. Epigenetic centromere identity is precisely maintained through DNA replication but is uniquely specified among human cells. Life Sci Alliance .
    pmc: PMC9811134pubmed: 36596606doi: 10.26508/lsa.202201807google scholar: lookup
  56. Maloney KA, Sullivan LL, Matheny JE, Strome ED, Merrett SL, Ferris A, Sullivan BA. Functional epialleles at an endogenous human centromere. Proc Natl Acad Sci U S A 109:13704–13709.
    pmc: PMC3427087pubmed: 22847449
  57. Marshall OJ, Chueh AC, Wong LH, Choo KH. Neocentromeres: new insights into centromere structure, disease development, and karyotype evolution. Am J Hum Genet 82:261–282.
    pmc: PMC2427194pubmed: 18252209
  58. Masumoto H, Masukata H, Muro Y, Nozaki N, Okazaki T. A human centromere antigen (CENP-B) interacts with a short specific sequence in alphoid DNA, a human centromeric satellite. J Cell Biol 109:1963–1973.
    pmc: PMC2115871pubmed: 2808515
  59. Miga KH, Eichler EE. Envisioning a new era: Complete genetic information from routine, telomere-to-telomere genomes. Am J Hum Genet 110:1832–1840.
    pmc: PMC10645551pubmed: 37922882
  60. Montefalcone G, Tempesta S, Rocchi M, Archidiacono N. Centromere repositioning. Genome Res 9:1184–1188.
    pmc: PMC311001pubmed: 10613840
  61. Murillo-Pineda M, Valente LP, Dumont M, Mata JF, Fachinetti D, Jansen LET. Induction of spontaneous human neocentromere formation and long-term maturation. J Cell Biol .
    pmc: PMC7812830pubmed: 33443568doi: 10.1083/jcb.202007210google scholar: lookup
  62. Murphy WJ, Pringle TH, Crider TA, Springer MS, Miller W. Using genomic data to unravel the root of the placental mammal phylogeny. Genome Res 17:413–421.
    pmc: PMC1832088pubmed: 17322288
  63. Musilova P, Kubickova S, Vahala J, Rubes J. Subchromosomal karyotype evolution in Equidae. Chromosome Res 21:175–187.
    pubmed: 23532666
  64. Nagpal H, Ali-Ahmad A, Hirano Y, Cai W, Halic M, Fukagawa T, Sekulić N, Fierz B. CENP-A and CENP-B collaborate to create an open centromeric chromatin state. Nat Commun 14:8227.
    pmc: PMC10716449pubmed: 38086807
  65. Nergadze SG, Lupotto M, Pellanda P, Santagostino M, Vitelli V, Giulotto E. Mitochondrial DNA insertions in the nuclear horse genome. Anim Genet 41(Suppl 2):176–185.
    pubmed: 21070293
  66. Nergadze SG, Belloni E, Piras FM, Khoriauli L, Mazzagatti A, Vella F, Bensi M, Vitelli V, Giulotto E, Raimondi E. Discovery and comparative analysis of a novel satellite, EC137, in horses and other equids. Cytogenet Genome Res 144:114–123.
    pubmed: 25342230
  67. Nergadze SG, Piras FM, Gamba R, Corbo M, Cerutti F, McCarter JGW, Cappelletti E, Gozzo F, Harman RM, Antczak DF. Birth, evolution, and transmission of satellite-free mammalian centromeric domains. Genome Res 28:789–799.
    pmc: PMC5991519pubmed: 29712753
  68. Nurk S, Koren S, Rhie A, Rautiainen M, Bzikadze AV, Mikheenko A, Vollger MR, Altemose N, Uralsky L, Gershman A. The complete sequence of a human genome. Science 376:44–53.
    pmc: PMC9186530pubmed: 35357919
  69. Oakenfull EA, Lim HN, Ryder OA. A survey of equid mitochondrial DNA: Implications for the evolution, genetic diversity and conservation of Equus. Conserv Genet 1:341–355.
  70. Orlando L. Equids. Curr Biol 25:R973-978.
    pubmed: 26485367
  71. Orlando L, Ginolhac A, Zhang G, Froese D, Albrechtsen A, Stiller M, Schubert M, Cappellini E, Petersen B, Moltke I. Recalibrating Equus evolution using the genome sequence of an early Middle Pleistocene horse. Nature 499:74–78.
    pubmed: 23803765
  72. Pardo-Manuel de Villena F, Sapienza C. Female meiosis drives karyotypic evolution in mammals. Genetics 159:1179–1189.
    pmc: PMC1461872pubmed: 11729161
  73. Peng S, Petersen JL, Bellone RR, Kalbfleisch T, Kingsley NB, Barber AM, Cappelletti E, Giulotto E, Finno CJ. Decoding the equine genome: lessons from ENCODE. Genes .
    pmc: PMC8625040pubmed: 34828313doi: 10.3390/genes12111707google scholar: lookup
  74. Piras FM, Nergadze SG, Poletto V, Cerutti F, Ryder OA, Leeb T, Raimondi E, Giulotto E. Phylogeny of horse chromosome 5q in the genus Equus and centromere repositioning. Cytogenet Genome Res 126:165–172.
    pubmed: 20016166
  75. Piras FM, Nergadze SG, Magnani E, Bertoni L, Attolini C, Khoriauli L, Raimondi E, Giulotto E. Uncoupling of satellite DNA and centromeric function in the genus Equus. PLoS Genet 6:e1000845.
    pmc: PMC2820525pubmed: 20169180
  76. Piras FM, Cappelletti E, Santagostino M, Nergadze SG, Giulotto E, Raimondi E. Molecular dynamics and evolution of centromeres in the genus Equus. Int J Mol Sci .
    pmc: PMC9024551pubmed: 35457002doi: 10.3390/ijms23084183google scholar: lookup
  77. Piras FM, Cappelletti E, Abdelgadir WA, Salamon G, Vignati S, Santagostino M, Sola L, Nergadze SG, Giulotto E. A satellite-free centromere in Equus przewalskii chromosome 10. Int J Mol Sci .
    pmc: PMC9961726pubmed: 36835543doi: 10.3390/ijms24044134google scholar: lookup
  78. Plohl M, Luchetti A, Mestrović N, Mantovani B. Satellite DNAs between selfishness and functionality: structure, genomics and evolution of tandem repeats in centromeric (hetero)chromatin. Gene 409:72–82.
    pubmed: 18182173
  79. Plohl M, Meštrović N, Mravinac B. Centromere identity from the DNA point of view. Chromosoma 123:313–325.
    pmc: PMC4107277pubmed: 24763964
  80. Prothero DR, Schoch RM. The Evolution of Perissodactyls. Oxford University Press .
  81. Purgato S, Belloni E, Piras FM, Zoli M, Badiale C, Cerutti F, Mazzagatti A, Perini G, Della Valle G, Nergadze SG. Centromere sliding on a mammalian chromosome. Chromosoma 124:277–287.
    pmc: PMC4446527pubmed: 25413176
  82. Roberti A, Bensi M, Mazzagatti A, Piras FM, Nergadze SG, Giulotto E, Raimondi E. Satellite DNA at the centromere is dispensable for segregation fidelity. Genes .
    pmc: PMC6627300pubmed: 31226862doi: 10.3390/genes10060469google scholar: lookup
  83. Rocchi M, Archidiacono N, Schempp W, Capozzi O, Stanyon R. Centromere repositioning in mammals. Hered (Edinb) 108:59–67.
    pmc: PMC3238114pubmed: 22045381
  84. Ryder OA, Epel NC, Benirschke K. Chromosome banding studies of the Equidae. Cytogenet Cell Genet 20:332–350.
    pubmed: 648186
  85. Santagostino M, Khoriauli L, Gamba R, Bonuglia M, Klipstein O, Piras FM, Vella F, Russo A, Badiale C, Mazzagatti A. Genome-wide evolutionary and functional analysis of the Equine Repetitive Element 1: an insertion in the myostatin promoter affects gene expression. BMC Genet 16:126.
    pmc: PMC4623272pubmed: 26503543
  86. Santagostino M, Piras FM, Cappelletti E, Del Giudice S, Semino O, Nergadze SG, Giulotto E. Insertion of telomeric repeats in the human and horse genomes: an evolutionary perspective. Int J Mol Sci .
    pmc: PMC7215372pubmed: 32325780doi: 10.3390/ijms21082838google scholar: lookup
  87. Schubert I. What is behind “centromere repositioning”?. Chromosoma 127:229–234.
    pubmed: 29705818
  88. Shang WH, Hori T, Toyoda A, Kato J, Popendorf K, Sakakibara Y, Fujiyama A, Fukagawa T. Chickens possess centromeres with both extended tandem repeats and short non-tandem-repetitive sequences. Genome Res 20:1219–1228.
    pmc: PMC2928500pubmed: 20534883
  89. Sola L, Nergadze SG, Cappelletti E, Piras FM, Giulotto E, Santagostino M. Telomeric-like repeats flanked by sequences retrotranscribed from the telomerase RNA inserted at DNA double-strand break sites during vertebrate genome evolution. Int J Mol Sci .
    pmc: PMC8537989pubmed: 34681704doi: 10.3390/ijms222011048google scholar: lookup
  90. Sultana T, van Essen D, Siol O, Bailly-Bechet M, Philippe C, Zine El Aabidine A, Pioger L, Nigumann P, Saccani S, Andrau JC. The Landscape of L1 Retrotransposons in the Human Genome Is Shaped by Pre-insertion Sequence Biases and Post-insertion Selection. Mol Cell 74:555-570.e557.
    pubmed: 30956044
  91. Talbert PB, Henikoff S. Centromeres convert but don’t cross. PLoS Biol 8:e1000326.
    pmc: PMC2834710pubmed: 20231873
  92. Talbert PB, Henikoff S. What makes a centromere?. Exp Cell Res 389:111895.
    pubmed: 32035948
  93. Talbert PB, Henikoff S. The genetics and epigenetics of satellite centromeres. Genome Res 32:608–615.
    pmc: PMC8997354pubmed: 35361623
  94. Talbert PB, Bryson TD, Henikoff S. Adaptive evolution of centromere proteins in plants and animals. J Biol 3:18.
    pmc: PMC549713pubmed: 15345035
  95. Tolomeo D, Capozzi O, Stanyon RR, Archidiacono N, D’Addabbo P, Catacchio CR, Purgato S, Perini G, Schempp W, Huddleston J. Epigenetic origin of evolutionary novel centromeres. Sci Rep 7:41980.
    pmc: PMC5290474pubmed: 28155877
  96. Tougard C, Delefosse T, Hänni C, Montgelard C. Phylogenetic relationships of the five extant Rhinoceros species (Rhinocerotidae, Perissodactyla) based on mitochondrial cytochrome b and 12S rRNA genes. Mol Phylogenet Evol 19:34–44.
    pubmed: 11286489
  97. Trifonov VA, Stanyon R, Nesterenko AI, Fu B, Perelman PL, O’Brien PC, Stone G, Rubtsova NV, Houck ML, Robinson TJ. Multidirectional cross-species painting illuminates the history of karyotypic evolution in Perissodactyla. Chromosome Res 16:89–107.
    pubmed: 18293107
  98. Trifonov VA, Musilova P, Kulemsina AI. Chromosome evolution in Perissodactyla. Cytogenet Genome Res 137:208–217.
    pubmed: 22813844
  99. van de Werken HJG, Haan JC, Feodorova Y, Bijos D, Weuts A, Theunis K, Holwerda SJB, Meuleman W, Pagie L, Thanisch K. Small chromosomal regions position themselves autonomously according to their chromatin class. Genome Res 27:922–933.
    pmc: PMC5453326pubmed: 28341771
  100. . The complete diploid reference genome of RPE-1 identifies human phased epigenetic landscapes. bioRxiv .
  101. Wade CM, Giulotto E, Sigurdsson S, Zoli M, Gnerre S, Imsland F, Lear TL, Adelson DL, Bailey E, Bellone RR. Genome sequence, comparative analysis, and population genetics of the domestic horse. Science 326:865–867.
    pmc: PMC3785132pubmed: 19892987
  102. Wei KH, Reddy HM, Rathnam C, Lee J, Lin D, Ji S, Mason JM, Clark AG, Barbash DA. A Pooled Sequencing Approach Identifies a Candidate Meiotic Driver. Genetics 206:451–465.
    pmc: PMC5419488pubmed: 28258181
  103. Xu X, Janke A, Arnason U. The complete mitochondrial DNA sequence of the greater Indian rhinoceros, Rhinoceros unicornis, and the phylogenetic relationship among Carnivora, Perissodactyla, and Artiodactyla (+ Cetacea). Mol Biol Evol 13:1167–1173.
    pubmed: 8896369
  104. Yadav V, Sun S, Billmyre RB, Thimmappa BC, Shea T, Lintner R, Bakkeren G, Cuomo CA, Heitman J, Sanyal K. RNAi is a critical determinant of centromere evolution in closely related fungi. Proc Natl Acad Sci U S A 115:3108–3113.
    pmc: PMC5866544pubmed: 29507212
  105. Yang F, Fu B, O’Brien PC, Robinson TJ, Ryder OA, Ferguson-Smith MA. Karyotypic relationships of horses and zebras: results of cross-species chromosome painting. Cytogenet Genome Res 102:235–243.
    pubmed: 14970709

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