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Kinetochores

Genetics and Genomics · NLM MeSH · updated 2026-10-29 · version 1

Citations verified against PubMedNot yet clinician-reviewed10 referencesApproximately 3 min read

Large multiprotein complexes that bind the centromeres of the chromosomes to the microtubules of the mitotic spindle during metaphase in the cell cycle.

Table of Contents

1. Definition2. Research landscape3. Reading the evidence4. Open-access studies5. Key reviews6. References7. About this article

Definition and research guide

This page combines a MeSH definition, publication statistics and selected research abstracts. It is a starting point for reading the literature, not a medical textbook or a systematic evidence review.

Content available: 4 complete abstracts (641 words), 10 references. Full article text is not stored or reproduced here; follow the PMID, DOI or PMC links to read the original sources and their access terms.

Definition

Large multiprotein complexes that bind the centromeres of the chromosomes to the microtubules of the mitotic spindle during metaphase in the cell cycle.

Definition: NLM Medical Subject Headings (MeSH) scope note (Public domain (U.S. National Library of Medicine); attribution requested). View the MeSH record.

Also known as: Kinetochore.

Where it fits in the MeSH classification

Broader topics
Centromere

Research landscape

PubMed indexes 3,843 records tagged with the MeSH heading “Kinetochores” (counts retrieved 2026-10-29). 723 of them (18.8%) were published from 2020 onward, and the busiest period so far is 2010-2019 with 1,614 records.

Reviews account for 592 records (15.4%) and meta-analyses for 0. Randomized controlled trials number 1 (0.0%) as a share of all records on the topic. Randomized trials make up only a small share of this literature, so much of what is known comes from laboratory, observational or descriptive work.

PubMed records by publication periodthrough 1999: 277 records; 2000-2009: 1,229 records; 2010-2019: 1,614 records; 2020 onward: 723 records277through 19991,2292000-20091,6142010-20197232020 onward
Records in PubMed indexed with this heading, by publication period. Real counts from NCBI E-utilities; the final period covers only part of the current year. Volume reflects how much has been published, not how strong the evidence is.
Show the data
PeriodRecords
through 1999277
2000-20091,229
2010-20191,614
2020 onward723
Share of records by publication typeReviews: 15.4 percent; Meta-analyses: 0.0 percent; Randomized controlled trials: 0.0 percentReviews15.4% (592)Meta-analyses0.0% (0)Randomized controlled trials0.0% (1)
Share of all PubMed records on this heading that carry each publication type. Types overlap with other records, so the shares do not sum to 100%.
Publication-type chart data
Publication types among 3,843 PubMed records
TypeRecordsShare
Reviews59215.4%
Meta-analyses00.0%
Randomized controlled trials10.0%

Method: PubMed query "Kinetochores"[MeSH Terms] with publication-type and date filters, retrieved 2026-10-29.

Reading the evidence in genetics and genomics

Genetic findings vary widely in strength. A variant linked to a condition in one study may not replicate in other ancestries, and most common variants have small effects that are meaningful only in aggregate. Studies of rare, high-impact variants can support strong causal claims about a gene, while association studies describe statistical links that need functional and independent replication. Check the sample size, the ancestry of participants, whether results were replicated, and whether a laboratory or clinical outcome was actually measured.

Evidence from open-access studies

These open-access studies come from PubMed’s best-match results for this heading (papers whose main topic it is), ranked here by citation count, with one recent addition. Each abstract is reproduced in full under its recorded open licence so that you can read the authors’ own words. They illustrate the literature; they are not a systematic summary of it. Citation counts measure attention, not reliability or clinical benefit.

Study comparison — selected abstracts, not a systematic review
Study / authorsYearSource label / designEurope PMC citationsLicence / source
The Kinetochore Receptor for the Cohesin Loading Complex
Hinshaw SM, Makrantoni V, Harrison SC, Marston AL
2017Research article
Design not recorded in structured metadata
104cc by
PMID 28938124 · DOI · PMC5610175
Three-dimensional structure of kinetochore-fibers in human mitotic spindles
Kiewisz R, Fabig G, Conway W, Baum D, Needleman D, Müller-Reichert T
2022Research article
Design not recorded in structured metadata
87cc by
PMID 35894209 · DOI · PMC9365394
The COMA complex interacts with Cse4 and positions Sli15/Ipl1 at the budding yeast inner kinetochore
Fischböck-Halwachs J, Singh S, Potocnjak M, Hagemann G, Solis-Mezarino V, Woike S, et al.
2019Research article
Design not recorded in structured metadata
83cc by
PMID 31112132 · DOI · PMC6546395
Progressive coevolution of the yeast centromere and kinetochore
Helsen J, Ramachandran K, Sherlock G, Dey G
2026Research article
Design not recorded in structured metadata
11cc by
PMID 41299172 · DOI · PMC12925627

The Kinetochore Receptor for the Cohesin Loading Complex [1]

Research article Hinshaw SM, Makrantoni V, Harrison SC, Marston AL · · 2017 · cited 104 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
The ring-shaped cohesin complex brings together distant DNA domains to maintain, express, and segregate the genome.
Closing excerpt
These findings represent a complete molecular description of targeted cohesin loading, a phenomenon with wide-ranging importance in chromosome segregation and, in multicellular organisms, transcription regulation.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (143 words)

The ring-shaped cohesin complex brings together distant DNA domains to maintain, express, and segregate the genome. Establishing specific chromosomal linkages depends on cohesin recruitment to defined loci. One such locus is the budding yeast centromere, which is a paradigm for targeted cohesin loading. The kinetochore, a multiprotein complex that connects centromeres to microtubules, drives the recruitment of high levels of cohesin to link sister chromatids together. We have exploited this system to determine the mechanism of specific cohesin recruitment. We show that phosphorylation of the Ctf19 kinetochore protein by a conserved kinase, DDK, provides a binding site for the Scc2/4 cohesin loading complex, thereby directing cohesin loading to centromeres. A similar mechanism targets cohesin to chromosomes in vertebrates. These findings represent a complete molecular description of targeted cohesin loading, a phenomenon with wide-ranging importance in chromosome segregation and, in multicellular organisms, transcription regulation.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 28938124 · DOI · PMC5610175

Three-dimensional structure of kinetochore-fibers in human mitotic spindles [2]

Research article Kiewisz R, Fabig G, Conway W, Baum D, Needleman D, Müller-Reichert T · · 2022 · cited 87 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
During cell division, kinetochore microtubules (KMTs) provide a physical linkage between the chromosomes and the rest of the spindle.
Closing excerpt
Finally, we also introduce a new visualization tool allowing an interactive display of our 3D spindle data that will serve as a resource for further structural studies on mitosis in human cells.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (202 words)

During cell division, kinetochore microtubules (KMTs) provide a physical linkage between the chromosomes and the rest of the spindle. KMTs in mammalian cells are organized into bundles, so-called kinetochore-fibers (k-fibers), but the ultrastructure of these fibers is currently not well characterized. Here, we show by large-scale electron tomography that each k-fiber in HeLa cells in metaphase is composed of approximately nine KMTs, only half of which reach the spindle pole. Our comprehensive reconstructions allowed us to analyze the three-dimensional (3D) morphology of k-fibers and their surrounding MTs in detail. We found that k-fibers exhibit remarkable variation in circumference and KMT density along their length, with the pole-proximal side showing a broadening. Extending our structural analysis then to other MTs in the spindle, we further observed that the association of KMTs with non-KMTs predominantly occurs in the spindle pole regions. Our 3D reconstructions have implications for KMT growth and k-fiber self-organization models as covered in a parallel publication applying complementary live-cell imaging in combination with biophysical modeling (Conway et al., 2022). Finally, we also introduce a new visualization tool allowing an interactive display of our 3D spindle data that will serve as a resource for further structural studies on mitosis in human cells.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 35894209 · DOI · PMC9365394

The COMA complex interacts with Cse4 and positions Sli15/Ipl1 at the budding yeast inner kinetochore [3]

Research article Fischböck-Halwachs J, Singh S, Potocnjak M, Hagemann G, Solis-Mezarino V, Woike S, et al. · · 2019 · cited 83 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
Kinetochores are macromolecular protein complexes at centromeres that ensure accurate chromosome segregation by attaching chromosomes to spindle microtubules and integrating safeguard mechanisms.
Closing excerpt
This study shows molecular characteristics of the point-centromere kinetochore architecture and suggests a role for the Ctf19 C-terminus in mediating CPC-binding and accurate chromosome segregation.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (150 words)

Kinetochores are macromolecular protein complexes at centromeres that ensure accurate chromosome segregation by attaching chromosomes to spindle microtubules and integrating safeguard mechanisms. The inner kinetochore is assembled on CENP-A nucleosomes and has been implicated in establishing a kinetochore-associated pool of Aurora B kinase, a chromosomal passenger complex (CPC) subunit, which is essential for chromosome biorientation. By performing crosslink-guided in vitro reconstitution of budding yeast kinetochore complexes we showed that the Ame1/Okp1CENP-U/Q heterodimer, which forms the COMA complex with Ctf19/Mcm21CENP-P/O, selectively bound Cse4CENP-A nucleosomes through the Cse4 N-terminus. The Sli15/Ipl1INCENP/Aurora-B core-CPC interacted with COMA in vitro through the Ctf19 C-terminus whose deletion affected chromosome segregation fidelity in Sli15 wild-type cells. Tethering Sli15 to Ame1/Okp1 rescued synthetic lethality upon Ctf19 depletion in a Sli15 centromere-targeting deficient mutant. This study shows molecular characteristics of the point-centromere kinetochore architecture and suggests a role for the Ctf19 C-terminus in mediating CPC-binding and accurate chromosome segregation.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 31112132 · DOI · PMC6546395

Progressive coevolution of the yeast centromere and kinetochore [4]

Research article Helsen J, Ramachandran K, Sherlock G, Dey G · · 2026 · cited 11 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
During mitosis, stable but dynamic interactions between centromere DNA and the kinetochore complex enable accurate and efficient chromosome segregation.
Closing excerpt
Together, our findings provide insight into the evolutionary constraints and trajectories shaping centromere evolution.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (146 words)

During mitosis, stable but dynamic interactions between centromere DNA and the kinetochore complex enable accurate and efficient chromosome segregation. Even though many proteins of the kinetochore are highly conserved1,2, centromeres are among the fastest evolving regions in a genome3,4, showing extensive variation even on short evolutionary timescales. Here we sought to understand how organisms evolve completely new sets of centromeres that still effectively engage with the kinetochore machinery by identifying and tracking thousands of centromeres across two major fungal clades, including more than 2,500 natural strain isolates and representing over 1,000 million years of evolution. We show that new centromeres spread progressively via drift and subsequent selection and that the kinetochore, which is evolving slowly in relative terms, appears to act as a filter to determine which new centromere variants are tolerated. Together, our findings provide insight into the evolutionary constraints and trajectories shaping centromere evolution.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 41299172 · DOI · PMC12925627

Key reviews and meta-analyses

The most cited review-type papers among PubMed’s best-match results for this heading, listed with bibliographic details only. Follow the links for the full text and access terms.

  • Research article Molecular architecture of the kinetochore-microtubule interface [5]
    Cheeseman IM, Desai A · Nat Rev Mol Cell Biol · 2008 · cited 769 times (Europe PMC)
  • Research article Un ménage à quatre: the molecular biology of chromosome segregation in meiosis [6]
    Petronczki M, Siomos MF, Nasmyth K · Cell · 2003 · cited 591 times (Europe PMC)
  • Research article Microtubule attachment and spindle assembly checkpoint signalling at the kinetochore [7]
    Foley EA, Kapoor TM · Nat Rev Mol Cell Biol · 2013 · cited 545 times (Europe PMC)
  • Research article How cells get the right chromosomes [8]
    Nicklas RB · Science · 1997 · cited 445 times (Europe PMC)
  • Research article The spindle checkpoint: structural insights into dynamic signalling [9]
    Musacchio A, Hardwick KG · Nat Rev Mol Cell Biol · 2002 · cited 404 times (Europe PMC)
  • Research article The life and miracles of kinetochores [10]
    Santaguida S, Musacchio A · EMBO J · 2009 · cited 404 times (Europe PMC)

References

  1. Hinshaw SM, Makrantoni V, Harrison SC, Marston AL. The Kinetochore Receptor for the Cohesin Loading Complex. . 2017. PMID 28938124 · DOI · PMC5610175
  2. Kiewisz R, Fabig G, Conway W, Baum D, Needleman D, Müller-Reichert T. Three-dimensional structure of kinetochore-fibers in human mitotic spindles. . 2022. PMID 35894209 · DOI · PMC9365394
  3. Fischböck-Halwachs J, Singh S, Potocnjak M, Hagemann G, Solis-Mezarino V, Woike S, et al.. The COMA complex interacts with Cse4 and positions Sli15/Ipl1 at the budding yeast inner kinetochore. . 2019. PMID 31112132 · DOI · PMC6546395
  4. Helsen J, Ramachandran K, Sherlock G, Dey G. Progressive coevolution of the yeast centromere and kinetochore. . 2026. PMID 41299172 · DOI · PMC12925627
  5. Cheeseman IM, Desai A. Molecular architecture of the kinetochore-microtubule interface. Nat Rev Mol Cell Biol. 2008. PMID 18097444 · DOI
  6. Petronczki M, Siomos MF, Nasmyth K. Un ménage à quatre: the molecular biology of chromosome segregation in meiosis. Cell. 2003. PMID 12600308 · DOI
  7. Foley EA, Kapoor TM. Microtubule attachment and spindle assembly checkpoint signalling at the kinetochore. Nat Rev Mol Cell Biol. 2013. PMID 23258294 · DOI · PMC3762224
  8. Nicklas RB. How cells get the right chromosomes. Science. 1997. PMID 9005842 · DOI
  9. Musacchio A, Hardwick KG. The spindle checkpoint: structural insights into dynamic signalling. Nat Rev Mol Cell Biol. 2002. PMID 12360190 · DOI
  10. Santaguida S, Musacchio A. The life and miracles of kinetochores. EMBO J. 2009. PMID 19629042 · DOI · PMC2722247

About this article

Sources
NLM Medical Subject Headings (definition, synonyms); NCBI PubMed (counts, identifiers); Europe PMC (open-access abstracts and citation counts).
Verification
Every reference was resolved against PubMed on 2026-10-31; title, year and DOI matched.
Review state
automated source verified; no human expert review has been recorded yet.
Version
1, last updated 2026-10-29.

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At a glance

3,843
PubMed records on this heading
19%
published since 2020
592
review articles
1
randomized trials
10
verified references below

Counts come from PubMed on 2026-10-29; they show how much has been written, not how strong the evidence is.

Related topics See all →

  • CentromereGenetics and Genomics
  • ChromatidsGenetics and Genomics
  • ChromatinGenetics and Genomics
  • EuchromatinGenetics and Genomics
  • NucleosomesGenetics and Genomics

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