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Centromere and kinetochore | chromosome structure and function
Centromere and kinetochore | chromosome structure and function
Published: 2015/02/25
Channel: Shomu's Biology
What is CENTROMERE? What does CENTROMERE mean? CENTROMERE meaning, definition & explanation
What is CENTROMERE? What does CENTROMERE mean? CENTROMERE meaning, definition & explanation
Published: 2016/10/18
Channel: The Audiopedia
MCAT Centromeres
MCAT Centromeres
Published: 2016/01/15
Channel: MCAT Prep by PremedHQ
Centromere 3-D
Centromere 3-D
Published: 2015/03/19
Channel: National Human Genome Research Institute
Centromeres and Telomeres tutorial
Centromeres and Telomeres tutorial
Published: 2014/03/21
Channel: Streaming Tutors
Chromosome and Kinetochore
Chromosome and Kinetochore
Published: 2013/09/30
Channel: WEHImovies
Centromere
Centromere
Published: 2014/09/28
Channel: Audiopedia
GENETICS: KINETOCHORE & MITOSIS
GENETICS: KINETOCHORE & MITOSIS
Published: 2012/08/12
Channel: Walter Jahn
Tandem Repeats of the Centromere
Tandem Repeats of the Centromere
Published: 2017/05/01
Channel: Craig Paardekooper
Telomeres, Chromatids, Chromatin, Centromeres, Genes
Telomeres, Chromatids, Chromatin, Centromeres, Genes
Published: 2017/01/29
Channel: Nikolay's Genetics Lessons
Difference Between Centriole and Centrosome
Difference Between Centriole and Centrosome
Published: 2016/01/05
Channel: Difference Between
[read] The Centromere
[read] The Centromere
Published: 2017/06/12
Channel: kjhfufu
Centromere Clustering
Centromere Clustering
Published: 2014/03/14
Channel: Stowers Institute for Medical Research
Mitosis, Meiosis, Chromosomes, Chromatids, Chromatin and Centromeres . Dr,, Dawe Video.
Mitosis, Meiosis, Chromosomes, Chromatids, Chromatin and Centromeres . Dr,, Dawe Video.
Published: 2015/02/21
Channel: DrChrisDawe VideoTutor
Centromere Meaning
Centromere Meaning
Published: 2015/04/22
Channel: SDictionary
What Is A Telomere?
What Is A Telomere?
Published: 2013/03/13
Channel: Kerry McCarthy
What different types of chromosomes exist?
What different types of chromosomes exist?
Published: 2013/10/12
Channel: Nikolay's Genetics Lessons
Another Century
Another Century's Episode 3 - Centromere
Published: 2010/09/24
Channel: Kasidej Kamalanavin
Evolution Of Centromere Models
Evolution Of Centromere Models
Published: 2017/04/12
Channel: Alyona Fulp
"Genetics", Centromeres
"Genetics", Centromeres
Published: 2015/06/18
Channel: MyCyberCollege
CHROMOSOME TERMINOLOGY BASED ON CENTROMERE POSITION
CHROMOSOME TERMINOLOGY BASED ON CENTROMERE POSITION
Published: 2016/07/20
Channel: Walter Jahn
Evolution Of Centromere Models v2
Evolution Of Centromere Models v2
Published: 2017/04/07
Channel: Josh Lawrimore
JCB : Telomeres and centromeres have interchangeable roles in promoting meiotic spindle formation
JCB : Telomeres and centromeres have interchangeable roles in promoting meiotic spindle formation
Published: 2015/02/28
Channel: ScienceVio
Genetics: KNOW: The 4 Types of Chromosomes
Genetics: KNOW: The 4 Types of Chromosomes
Published: 2014/12/28
Channel: Neurogenesis Learning
centromere!
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Published: 2007/03/24
Channel: kharps
1717: Anti-SSA or Anti-Centromere
1717: Anti-SSA or Anti-Centromere
Published: 2016/09/28
Channel: latisom
mappingorderedtetrad
mappingorderedtetrad
Published: 2015/02/24
Channel: daajjj
Centromere
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Published: 2011/06/17
Channel: aslstemforum
Centromere - edited by Heesung
Centromere - edited by Heesung
Published: 2014/09/19
Channel: 김희성
Centromere Cluster
Centromere Cluster
Published: 2011/11/02
Channel: LabSpaces Video Feed
What Is The Definition Of Centromere Medical Dictionary Free Online
What Is The Definition Of Centromere Medical Dictionary Free Online
Published: 2014/09/21
Channel: Medical Dictionary Online
170207 Centromere Cover I.O.I - whatta Man @DebsirinRomklao school
170207 Centromere Cover I.O.I - whatta Man @DebsirinRomklao school
Published: 2017/02/07
Channel: Nutdanai Churee
centromere
centromere
Published: 2015/01/20
Channel: SMARTSign Dictionary
Medical vocabulary: What does Centromere mean
Medical vocabulary: What does Centromere mean
Published: 2016/01/17
Channel: botcaster inc. bot
Karen Miga | Linear Assembly of a Human Y Centromere using MinION Nanopore Long Read Sequences
Karen Miga | Linear Assembly of a Human Y Centromere using MinION Nanopore Long Read Sequences
Published: 2017/06/19
Channel: Oxford Nanopore Technologies
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centromere: kurious oranj
Published: 2008/12/08
Channel: xenopusRTRT
Chromosome
Chromosome
Published: 2016/02/08
Channel: SliderBase
Centromere Structure and Evolution Progress in Molecular and Subcellular Biology
Centromere Structure and Evolution Progress in Molecular and Subcellular Biology
Published: 2017/02/05
Channel: herman
CENTROMERE 항체 양성이면 피부경화증(Scleroderma) 국소형인가 전신형인가
CENTROMERE 항체 양성이면 피부경화증(Scleroderma) 국소형인가 전신형인가
Published: 2017/01/22
Channel: 유창길한의원
Download Centromere Structure and Evolution Progress in Molecular and Subcellular Biology PDF
Download Centromere Structure and Evolution Progress in Molecular and Subcellular Biology PDF
Published: 2017/03/20
Channel: M Brodsky
Centromere Structure and Evolution Progress in Molecular and Subcellular Biology
Centromere Structure and Evolution Progress in Molecular and Subcellular Biology
Published: 2017/02/04
Channel: ben
Plant Centromere Biology
Plant Centromere Biology
Published: 2017/02/07
Channel: Amy P
Centromere Top # 12 Facts
Centromere Top # 12 Facts
Published: 2015/11/04
Channel: Parash Surayavan
How to say "centromere"! (High Quality Voices)
How to say "centromere"! (High Quality Voices)
Published: 2017/01/31
Channel: WordBox
Plant Centromere Biology
Plant Centromere Biology
Published: 2017/02/14
Channel: maya
Plant Centromere Biology
Plant Centromere Biology
Published: 2017/03/09
Channel: saddha
Centromere - ATP (Music Only)
Centromere - ATP (Music Only)
Published: 2014/05/12
Channel: Mark Perez
Sylvia Erhardt - A role for RNA in centromere biology and genome integrity
Sylvia Erhardt - A role for RNA in centromere biology and genome integrity
Published: 2017/07/10
Channel: CellNetworks
Centromere Structure and Evolution Progress in Molecular and Subcellular Biology
Centromere Structure and Evolution Progress in Molecular and Subcellular Biology
Published: 2017/02/09
Channel: Evie Maruff
Final  Animal Biology - Centromere Repositioning
Final Animal Biology - Centromere Repositioning
Published: 2013/04/25
Channel: Rhianna Daniels
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WIKIPEDIA ARTICLE

From Wikipedia, the free encyclopedia
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In this diagram of a duplicated chromosome, (2) identifies the centromere—the region that joins the two sister chromatids, or each half of the chromosome. In prophase of mitosis, specialized regions on centromeres called kinetochores attach chromosomes to spindle fibers.

The centromere is the part of a chromosome that links sister chromatids or a dyad. During mitosis, spindle fibers attach to the centromere via the kinetochore.[1] Centromeres were first thought to be genetic loci that direct the behavior of chromosomes.

The physical role of the centromere is to act as the site of assembly of the kinetochore – a highly complex multiprotein structure that is responsible for the actual events of chromosome segregation – i.e. binding microtubules and signalling to the cell cycle machinery when all chromosomes have adopted correct attachments to the spindle, so that it is safe for cell division to proceed to completion and for cells to enter anaphase.[2]

There are, broadly speaking, two types of centromeres. "Point centromeres" bind to specific proteins that recognize particular DNA sequences with high efficiency.[3] Any piece of DNA with the point centromere DNA sequence on it will typically form a centromere if present in the appropriate species. The best characterised point centromeres are those of the budding yeast, Saccharomyces cerevisiae. "Regional centromeres" is the term coined to describe most centromeres, which typically form on regions of preferred DNA sequence, but which can form on other DNA (Deoxyribonucleic acid ) sequences as well.[3] The signal for formation of a regional centromere appears to be epigenetic. Most organisms, ranging from the fission yeast Schizosaccharomyces pombe to humans, have regional centromeres.

Regarding mitotic chromosome structure, centromeres represent a constricted region of the chromosome (often referred to as the primary constriction) where two identical sister chromatids are most closely in contact. When cells enter mitosis, the sister chromatids (the two copies of each chromosomal DNA molecule resulting from DNA replication in chromatin form) are linked along their length by the action of the cohesin complex. It is now believed that this complex is mostly released from chromosome arms during prophase, so that by the time the chromosomes line up at the mid-plane of the mitotic spindle (also known as the metaphase plate), the last place where they are linked with one another is in the chromatin in and around the centromere.[4]

Positions[edit]

Classifications of Chromosomes
I: Telocentric - centromere placement very close to the top, p arms barely visible if visible at all
II: Acrocentric - q arms are still much longer than the p arms, but the p arms are longer than those in telocentric
III: Submetacentric - p and q arms are very close in length but not equal
IV: Metacentric - p q arms are equal in length
A: Short arm (p arm)
B: Centromere
C: Long arm (q arm)
D: Sister Chromatid

Each chromosome has two arms, labeled p (the shorter of the two) and q (the longer). Many remember that the short arm 'p' is named for the French word "petit" meaning 'small', although this explanation was shown to be apocryphal.[5] They can be connected in either metacentric, submetacentric, acrocentric or telocentric manner.[6][7]

Categorization of chromosomes according to the relative arms length[8]
Centromere position Arms length ratio Sign Description
Medial sensu stricto 1.0 – 1.6 M Metacentric
Medial region 1.7 m Metacentric
Submedial 3.0 sm Submetacentric
Subterminal 3.1 – 6.9 st Subtelocentric
Terminal region 7.0 t Acrocentric
Terminal sensu stricto T Telocentric
Notes Metacentric: M+m Atelocentric: M+m+sm+st+t

Metacentric[edit]

These are X-shaped chromosomes, with the centromere in the middle so that the two arms of the chromosomes are almost equal.

A chromosome is metacentric if its two arms are roughly equal in length. In a normal human karyotype, five chromosomes are considered metacentric: chromosomes 1, 3, 16, 19, and 20. In some cases, a metacentric chromosome is formed by balanced translocation: the fusion of two acrocentric chromosomes to form one metacentric chromosome.[9][10]

Submetacentric[edit]

If arms' lengths are unequal, the chromosome is said to be submetacentric. Their shape is j shape during anaphase[11]

Acrocentric[edit]

If the p (short) arm is so short that it is hard to observe, but still present, then the chromosome is acrocentric (the "acro-" in acrocentric refers to the Greek word for "peak"). The human genome includes six acrocentric chromosomes: 13, 14, 15, 21, 22 and the Y chromosome.[11]

In an acrocentric chromosome the p arm contains genetic material including repeated sequences such as nucleolar organizing regions, and can be translocated without significant harm, as in a balanced Robertsonian translocation. The domestic horse genome includes one metacentric chromosome that is homologous to two acrocentric chromosomes in the conspecific but undomesticated Przewalski's horse.[12] This may reflect either fixation of a balanced Robertsonian translocation in domestic horses or, conversely, fixation of the fission of one metacentric chromosome into two acrocentric chromosomes in Przewalski's horses. A similar situation exists between the human and great ape genomes; in this case, because more species are extant, it is apparent that the evolutionary sequence is a reduction of two acrocentric chromosomes in the great apes to one metacentric chromosome in humans (see Karyotype#Aneuploidy).[11]

Telocentric[edit]

A telocentric chromosome's centromere is located at the terminal end of the chromosome. Telomeres may extend from both ends of the chromosome.their shape is like i during anaphase. For example, the standard house mouse karyotype has only telocentric chromosomes.[13][14] Humans do not possess telocentric chromosomes.

Subtelocentric[edit]

If the chromosome's centromere is located closer to its end than to its center, it may be described as subtelocentric.[citation needed][15]

Holocentric[edit]

With holocentric chromosomes, the entire length of the chromosome acts as the centromere. Examples of this type of centromere can be found scattered throughout the plant and animal kingdoms,[16] with the most well-known example being the nematode Caenorhabditis elegans.

Human chromosomes[edit]

Chromosome Centromere position (Mbp) Chromosome Size (Mbp) Centromere size (Mbp)
1 125.0 metacentric 247.2 7.4
2 93.3 submetacentric 242.8 6.3
3 91.0 metacentric 199.4 6.0
4 50.4 submetacentric 191.3
5 48.4 submetacentric 180.8
6 61.0 submetacentric 170.9
7 59.9 submetacentric 158.8
8 45.6 submetacentric 146.3
9 49.0 submetacentric 140.4
10 40.2 submetacentric 135.4
11 53.7 submetacentric 134.5
12 35.8 submetacentric 132.3
13 17.9 acrocentric 114.1
14 17.6 acrocentric 106.3
15 19.0 acrocentric 100.3
16 36.6 metacentric 88.8
17 24.0 submetacentric 78.7
18 17.2 submetacentric 76.1
19 26.5 metacentric 63.8
20 27.5 metacentric 62.4
21 13.2 acrocentric 46.9
22 14.7 acrocentric 49.5
X 60.6 submetacentric 154.9
Y 12.5 acrocentric 57.7

Sequence[edit]

There are two types of centromeres.[17] In regional centromeres, DNA sequences contribute to but do not define function. Regional centromeres contain large amounts of DNA and are often packaged into heterochromatin. In most eukaryotes, the centromere's DNA sequence consists of large arrays of repetitive DNA (e.g. satellite DNA) where the sequence within individual repeat elements is similar but not identical. In humans, the primary centromeric repeat unit is called α-satellite (or alphoid), although a number of other sequence types are found in this region.[18]

Point centromeres are smaller and more compact. DNA sequences are both necessary and sufficient to specify centromere identity and function in organisms with point centromeres. In budding yeasts, the centromere region is relatively small (about 125 bp DNA) and contains two highly conserved DNA sequences that serve as binding sites for essential kinetochore proteins.[18]

Inheritance[edit]

Since centromeric DNA sequence is not the key determinant of centromeric identity in metazoans, it is thought that epigenetic inheritance plays a major role in specifying the centromere.[19] The daughter chromosomes will assemble centromeres in the same place as the parent chromosome, independent of sequence. It has been proposed that histone H3 variant CENP-A (Centromere Protein A) is the epigenetic mark of the centromere.[20] The question arises whether there must be still some original way in which the centromere is specified, even if it is subsequently propagated epigenetically. If the centromere is inherited epigenetically from one generation to the next, the problem is pushed back to the origin of the first metazoans.

Structure[edit]

The centromeric DNA is normally in a heterochromatin state, which is essential for the recruitment of the cohesin complex that mediates sister chromatid cohesion after DNA replication as well as coordinating sister chromatid separation during anaphase. In this chromatin, the normal histone H3 is replaced with a centromere-specific variant, CENP-A in humans.[21] The presence of CENP-A is believed to be important for the assembly of the kinetochore on the centromere. CENP-C has been shown to localise almost exclusively to these regions of CENP-A associated chromatin. In human cells, the histones are found to be most enriched for H4K20me3 and H3K9me3[22] which are known heterochromatic modifications.

In the yeast Schizosaccharomyces pombe (and probably in other eukaryotes), the formation of centromeric heterochromatin is connected to RNAi.[23] In nematodes such as Caenorhabditis elegans, some plants, and the insect orders Lepidoptera and Hemiptera, chromosomes are "holocentric", indicating that there is not a primary site of microtubule attachments or a primary constriction, and a "diffuse" kinetochore assembles along the entire length of the chromosome.

Centromeric aberrations[edit]

In rare cases in humans, neocentromeres can form at new sites on the chromosome. There are currently over 90 known human neocentromeres identified on 20 different chromosomes.[24][25] The formation of a neocentromere must be coupled with the inactivation of the previous centromere, since chromosomes with two functional centromeres (Dicentric chromosome) will result in chromosome breakage during mitosis. In some unusual cases human neocentromeres have been observed to form spontaneously on fragmented chromosomes. Some of these new positions were originally euchromatic and lack alpha satellite DNA altogether.

Centromere proteins are also the autoantigenic target for some anti-nuclear antibodies, such as anti-centromere antibodies.

Dysfunction and disease[edit]

It has been known that centromere misregulation contributes to mis-segregation of chromosomes, which is strongly related to cancer and abortion. Notably, overexpression of many centromere genes have been linked to cancer malignant phenotypes. Overexpression of these centromere genes is thought to increase genomic instability in cancers. Elevated genomic instability on one hand relates to malignant phenotypes; on the other hand, it makes the tumor cells more vulnerable to specific adjuvant therapies such as certain chemotherapies and radiotherapy.[26]. Instability of centromere repetitive DNA was recently shown in cancer and aging [27].

Etymology and pronunciation[edit]

The word centromere (/ˈsɛntrəˌmɪər/[28][29]) uses combining forms of centro- and -mere, yielding "central part", describing the centromere's location at the center of the chromosome.

See also[edit]

References[edit]

  1. ^ Pollard, T.D. (2007). Cell Biology. Philadelphia: Saunders. pp. 200–203. ISBN 978-1-4160-2255-8. 
  2. ^ Pollard, TD (2007). Cell Biology. Philadelphia: Saunders. pp. 227–230. ISBN 978-1-4160-2255-8. 
  3. ^ a b Pluta, A.; A.M. Mackay; A.M. Ainsztein; I.G. Goldberg; W.C. Earnshaw (1995). "The centromere: Hub of chromosomal activities.". Science. 270 (5242): 1591–1594. PMID 7502067. doi:10.1126/science.270.5242.1591. 
  4. ^ "Sister chromatid cohesion". Genetics Home Reference. United States National Library of Medicine. May 15, 2011. 
  5. ^ "p + q = Solved, Being the True Story of How the Chromosome Got Its Name". 
  6. ^ Nikolay's Genetics Lessons (2013-10-12), What different types of chromosomes exist?, retrieved 2017-05-28 
  7. ^ Levan A., Fredga K., Sandberg A. A. (1964): Nomenclature for centromeric position on chromosomes. Hereditas, Lund, 52: 201.
  8. ^ Levan A., Fredga K., Sandberg A. A. (1964): Nomenclature for centromeric position on chromosomes. Hereditas, Lund, 52: 201.
  9. ^ "Chromosomes, Chromosome Anomalies". 
  10. ^ *Gilbert F (1999). "Disease genes and chromosomes: disease maps of the human genome. Chromosome 16". Genet Test. 3 (2): 243–54. PMID 10464676. 
  11. ^ a b c https://www.amazon.com/Thompson-Genetics-Medicine-Sixth-Edition/dp/0721669026
  12. ^ Myka, J.L.; Lear, T.L.; Houck, M.L.; Ryder, O.A.; Bailey, E. (2003). "FISH analysis comparing genome organization in the domestic horse (Equus caballus) to that of the Mongolian wild horse (E. przewalskii)". Cytogenetic and Genome Research. 102 (1–4): 222–5. PMID 14970707. doi:10.1159/000075753. 
  13. ^ Silver, Lee M. (1995). "Karyotypes, Chromosomes, and Translocations". Mouse Genetics: Concepts and Applications. Oxford: Oxford University Press. pp. 83–92. ISBN 978-0-19-507554-0. 
  14. ^ Chinwalla, Asif T.; Cook, Lisa L.; Delehaunty, Kimberly D.; Fewell, Ginger A.; Fulton, Lucinda A.; Fulton, Robert S.; Graves, Tina A.; Hillier, Ladeana W.; et al. (2002). "Initial sequencing and comparative analysis of the mouse genome". Nature. 420 (6915): 520–62. PMID 12466850. doi:10.1038/nature01262. 
  15. ^ Margulis, Lynn; Matthews, Clifford; Haselton, Aaron (2000-01-01). Environmental Evolution: Effects of the Origin and Evolution of Life on Planet Earth. MIT Press. ISBN 9780262631976. 
  16. ^ Dernburg, A. F. (2001). "Here, There, and Everywhere: Kinetochore Function on Holocentric Chromosomes". The Journal of Cell Biology. 153 (6): F33–8. PMC 2192025Freely accessible. PMID 11402076. doi:10.1083/jcb.153.6.F33. 
  17. ^ Pluta, A. F.; MacKay, A. M.; Ainsztein, A. M.; Goldberg, I. G.; Earnshaw, W. C. (1995). "The Centromere: Hub of Chromosomal Activities". Science. 270 (5242): 1591–4. PMID 7502067. doi:10.1126/science.270.5242.1591. 
  18. ^ a b Mehta, G. D.; Agarwal, M.; Ghosh, S. K. (2010). "Centromere Identity: a challenge to be faced". Mol. Genet. Genomics. 284 (2): 75–94. PMID 20585957. doi:10.1007/s00438-010-0553-4. 
  19. ^ Dalal, Yamini (2009). "Epigenetic specification of centromeres". Biochemistry and Cell Biology. 87 (1): 273–82. PMID 19234541. doi:10.1139/O08-135. 
  20. ^ Bernad, Rafael; Sánchez, Patricia; Losada, Ana (2009). "Epigenetic specification of centromeres by CENP-A". Experimental Cell Research. 315 (19): 3233–41. PMID 19660450. doi:10.1016/j.yexcr.2009.07.023. 
  21. ^ Chueh, A. C.; Wong, LH; Wong, N; Choo, KH (2004). "Variable and hierarchical size distribution of L1-retroelement-enriched CENP-A clusters within a functional human neocentromere". Human Molecular Genetics. 14 (1): 85–93. PMID 15537667. doi:10.1093/hmg/ddi008. 
  22. ^ Rosenfeld, Jeffrey A; Wang, Zhibin; Schones, Dustin E; Zhao, Keji; Desalle, Rob; Zhang, Michael Q (2009). "Determination of enriched histone modifications in non-genic portions of the human genome". BMC Genomics. 10: 143. PMC 2667539Freely accessible. PMID 19335899. doi:10.1186/1471-2164-10-143. 
  23. ^ Volpe, T. A.; Kidner, C; Hall, IM; Teng, G; Grewal, SI; Martienssen, RA (2002). "Regulation of Heterochromatic Silencing and Histone H3 Lysine-9 Methylation by RNAi". Science. 297 (5588): 1833–7. PMID 12193640. doi:10.1126/science.1074973. 
  24. ^ Marshall, Owen J.; Chueh, Anderly C.; Wong, Lee H.; Choo, K.H. Andy (2008). "Neocentromeres: New Insights into Centromere Structure, Disease Development, and Karyotype Evolution". The American Journal of Human Genetics. 82 (2): 261–82. PMC 2427194Freely accessible. PMID 18252209. doi:10.1016/j.ajhg.2007.11.009. 
  25. ^ Warburton, Peter E. (2004). "Chromosomal dynamics of human neocentromere formation". Chromosome Research. 12 (6): 617–26. PMID 15289667. doi:10.1023/B:CHRO.0000036585.44138.4b. 
  26. ^ Zhang, W.; Mao, J-H.; Zhu, W.; Jain, A.K.; Liu, L.; Brown, J.B.; Karpen, G.H. (2016). "Centromere and kinetochore gene misexpression predicts cancer patient survival and response to radiotherapy and chemotherapy". Nature Communications. 7: 12619. PMC 5013662Freely accessible. PMID 27577169. doi:10.1038/ncomms12619. 
  27. ^ Giunta, S; Funabiki, H (21 February 2017). "Integrity of the human centromere DNA repeats is protected by CENP-A, CENP-C, and CENP-T.". Proceedings of the National Academy of Sciences of the United States of America. 114 (8): 1928–1933. PMID 28167779. doi:10.1073/pnas.1615133114. 
  28. ^ "Centromere". Merriam-Webster Dictionary. 
  29. ^ "Centromere". Dictionary.com Unabridged. Random House. 

Further reading[edit]

  • Mehta, G. D.; Agarwal, M.; Ghosh, S. K. (2010). "Centromere Identity: a challenge to be faced". Mol. Genet. Genomics. 284 (2): 75–94. PMID 20585957. doi:10.1007/s00438-010-0553-4. 
  • Lodish, Harvey; Berk, Arnold; Kaiser, Chris A.; Krieger, Monty; Scott, Matthew P.; Bretscher, Anthony; Ploegh, Hiddle; Matsudaira, Paul (2008). Molecular Cell Biology (6th ed.). New York: W.H. Freeman. ISBN 978-0-7167-7601-7. 
  • Nagaki, Kiyotaka; Cheng, Zhukuan; Ouyang, Shu; Talbert, Paul B; Kim, Mary; Jones, Kristine M; Henikoff, Steven; Buell, C Robin; Jiang, Jiming (2004). "Sequencing of a rice centromere uncovers active genes". Nature Genetics. 36 (2): 138–45. PMID 14716315. doi:10.1038/ng1289. Lay summaryScience Daily (January 13, 2004). 

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