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{{short description|Stage of cell division}}
{{short description|Stage of cell division}}
{{for|the Product Lifecycle Management software|SDRC#Products}}
he spindle]]
[[File:Spindle chromosomes-en.png|thumb|right|250px|The mitotic spindle checkpoint verifies that all the chromosomes are aligned properly on the metaphase plate and prevents premature entry into anaphase.]]
[[File:Metaphase chromosomes.jpg|thumb|Chromosomes lined up on the metaphase plate. Two views with the metaphase plate rotated 60°.]]
[[File:Metaphase chromosomes.jpg|thumb|Chromosomes lined up on the metaphase plate. Two views with the metaphase plate rotated 60°.]]
[[File:Stages of early mitosis in a vertebrate cell with micrographs of chromatids.svg|thumb|Stages of early mitosis in a vertebrate cell with micrographs of chromatids]]
[[File:Stages of early mitosis in a vertebrate cell with micrographs of chromatids.svg|thumb|Stages of early mitosis in a vertebrate cell with micrographs of chromatids]]


'''Metaphase''' (from the [[Ancient Greek|Greek]] μετά, "adjacent" and φάσις, "stage") is a stage of [[mitosis]] in the [[eukaryote|eukaryotic]] [[cell cycle]] in which chromosomes are at their second-most condensed and coiled stage (they are at their most condensed in [[anaphase]]).<ref>{{cite web|title=Chromosome condensation through mitosis|url=https://www.sciencedaily.com/releases/2007/06/070611122252.htm|publisher=Science Daily|access-date=12 June 2007}}</ref> These [[chromosome]]s, carrying [[DNA sequence|genetic information]], align in the equator of the [[cell (biology)|cell]] before being separated into each of the two daughter cells.ges/C/CellCycle.html|publisher=Kimball's Biology Pages|access-date=9 December 2012|archive-url=https://web.archive.org/web/20121119134405/http://users.rcn.com/jkimball.ma.ultranet/BiologyPages/C/CellCycle.html|archive-date=19 November 2012|url-status=dead}}</ref> In certain types of cells, chromosomes do not line up at the metaphase plate and instead move back and forth between the poles randomly, only roughly lining up along the middleline.{{citation needed|date=December 2012}} Early events of metaphase can coincide with the later events of prometaphase, as chromosomes with connected kinetochores will start the events of metaphase individually before other chromosomes with unconnected kinetochores that are still lingering in the events of prometaphase.{{citation needed|date=December 2012}}
'''Metaphase''' ({{etymology|grc|''[[wikt:μετα-|μετα-]]'' ([[Meta (prefix)|meta-]]) beyond, above, transcending}} and {{etymology|grc|''[[wikt:φάσις|φάσις]]'' (phásis)|appearance}}) is a stage of [[mitosis]] in the [[eukaryote|eukaryotic]] [[cell cycle]] in which chromosomes are at their second-most condensed and coiled stage (they are at their most condensed in [[anaphase]]).<ref>{{cite web |title=Chromosome condensation through mitosis |url=https://www.sciencedaily.com/releases/2007/06/070611122252.htm |website=[[ScienceDaily]] |access-date=12 June 2007}}</ref> These [[chromosome]]s, carrying [[DNA sequence|genetic information]], align in the equator of the [[cell (biology)|cell]] between the spindle poles at the '''metaphase plate''', before being separated into each of the two daughter nuclei. This alignment marks the beginning of metaphase.<ref name="Alberts">{{cite book |last1=Alberts |first1=Bruce |last2=Hopkin |first2=Karen |last3=Johnson |first3=Alexander |last4=Morgan |first4=David |last5=Raff |first5=Martin |last6=Roberts |first6=Keith |last7=Walter |first7=Peter |title=Essential cell biology |date=2019 |publisher=W. W. Norton & Company |location=New York London |isbn=9780393680393 |page=632–633 |edition=Fifth}}</ref> Metaphase accounts for approximately 4% of the [[cell cycle]]'s duration.{{citation needed|date=December 2012}}


In metaphase, microtubules from both duplicated [[centrosome]]s on opposite poles of the cell have completed attachment to [[kinetochore]]s on condensed chromosomes. The [[centromere]]s of the chromosomes convene themselves on the metaphase plate, an imaginary line that is equidistant from the two spindle poles.<ref>{{cite web|title=Metaphase plate|url=http://www.biology-online.org/dictionary/Metaphase_plate|work=Biology Dictionary|publisher=Biology Online|access-date=9 December 2012}}</ref> This even alignment is due to the counterbalance of the pulling powers generated by the opposing kinetochore microtubules,<ref>{{cite web|title=Metaphase|url=http://www.nature.com/scitable/definition/metaphase-249|publisher=Nature Education|access-date=9 December 2012}}</ref> analogous to a tug-of-war between two people of equal strength, ending with the destruction of B [[cyclin]].<ref name="Kimball's Biology Pages">{{cite web|title=The Cell Cycle|url=http://users.rcn.com/jkimball.ma.ultranet/BiologyPages/C/CellCycle.html|publisher=Kimball's Biology Pages|access-date=9 December 2012|archive-url=https://web.archive.org/web/20121119134405/http://users.rcn.com/jkimball.ma.ultranet/BiologyPages/C/CellCycle.html|archive-date=19 November 2012|url-status=dead}}</ref>
One of the [[cell cycle checkpoint]]s occurs during prometaphase and metaphase. Only after all chromosomes have become aligned at the ''metaphase plate'', when every kinetochore is properly attached to a bundle of microtubules, does the cell enter anaphase. It is thought that unattached or improperly attached kinetochores generate a signal to prevent premature progression to anaphase, even if most of kinetochores have been attached and most of the chromosomes have been aligned. Such a signal creates the mitotic [[spindle checkpoint]]. This would be accomplished by regulation of the [[anaphase-promoting complex]], [[securin]], and [[separase]].

In order to prevent deleterious [[nondisjunction]] events, a key [[cell cycle checkpoint]], the [[spindle checkpoint]], verifies this evenly balanced alignment and ensures that every kinetochore is properly attached to a bundle of microtubules and is under balanced bipolar tension. Sister chromatids require active [[separase]] to hydrolyze the [[cohesin]] that bind them together prior to progression to [[anaphase]]. Any unattached or improperly attached kinetochores generate signals that prevent the activation of the [[anaphase promoting complex]] (cyclosome or APC/C), a [[ubiquitin ligase]] which targets [[securin]] and [[cyclin B]] for degradation via the [[proteosome]]. As long as securin and cyclin B remain active, separase remains inactive, preventing premature progression to anaphase.


==Metaphase in cytogenetics and cancer studies==
==Metaphase in cytogenetics and cancer studies==
[[Image:NHGRI human male karyotype.png|thumb|left|Human metaphase chromosomes (normal male [[karyotype]])]]
[[Image:NHGRI human male karyotype.png|thumb|right|Human metaphase chromosomes (normal male [[karyotype]])]]


The analysis of metaphase [[chromosome]]s is one of the main tools of classical [[cytogenetics]] and [[cancer]] studies. Chromosomes are condensed (thickened) and highly coiled in metaphase, which makes them most suitable for visual analysis. Metaphase chromosomes make the classical picture of chromosomes ([[karyotype]]). For classical cytogenetic analyses, cells are grown in short term culture and arrested in metaphase using [[mitotic inhibitor]]. Further they are used for slide preparation and banding ([[staining]]) of chromosomes to be visualised under microscope to study structure and number of chromosomes (karyotype). [[Staining]] of the slides, often with [[Giemsa]] ([[G banding]]) or [[Quinacrine]], produces a pattern of in total up to several hundred bands. Normal metaphase spreads are used in methods like [[Fluorescence in situ hybridization|FISH]] and as a hybridization matrix for [[comparative genomic hybridization]] (CGH) experiments.
The analysis of metaphase [[chromosome]]s is one of the main tools of classical [[cytogenetics]] and [[cancer]] studies. Chromosomes are condensed (thickened) and highly coiled in metaphase, which makes them most suitable for visual analysis. Metaphase chromosomes make the classical picture of chromosomes ([[karyotype]]). For classical cytogenetic analyses, cells are grown in short term culture and arrested in metaphase using [[mitotic inhibitor]]. Further they are used for slide preparation and banding ([[staining]]) of chromosomes to be visualised under microscope to study structure and number of chromosomes (karyotype). [[Staining]] of the slides, often with [[Giemsa]] ([[G banding]]) or [[Quinacrine]], produces a pattern of in total up to several hundred bands. Normal metaphase spreads are used in methods like [[Fluorescence in situ hybridization|FISH]] and as a hybridization matrix for [[comparative genomic hybridization]] (CGH) experiments.


[[Malignant cell]]s from solid [[tumor]]s or [[leukemia]] samples can also be used for cytogenetic analysis to generate metaphase preparations. Inspection of the stained metaphase chromosomes allows the determination of numerical and structural changes in the tumor cell genome, for example, losses of chromosomal segments or [[chromosomal translocation|translocations]], which may lead to chimeric [[oncogene]]s, such as [[bcr-abl]] in [[chronic myelogenous leukemia]].
[[Malignant cell]]s from solid [[tumor]]s or [[leukemia]] samples can also be used for cytogenetic analysis to generate metaphase preparations. Inspection of the stained metaphase chromosomes allows the determination of numerical and structural changes in the tumor cell genome, for example, losses of chromosomal segments or [[chromosomal translocation|translocations]], which may lead to chimeric [[oncogene]]s, such as [[bcr-abl]] in [[chronic myelogenous leukemia]].

==See also==
*[[Interphase]]
*[[Prophase]]
*[[Prometaphase]]
*[[Anaphase]]
*[[Telophase]]
*[[Cytoskeleton]]


==References==
==References==

Latest revision as of 17:31, 16 September 2024

The mitotic spindle checkpoint verifies that all the chromosomes are aligned properly on the metaphase plate and prevents premature entry into anaphase.
Chromosomes lined up on the metaphase plate. Two views with the metaphase plate rotated 60°.
Stages of early mitosis in a vertebrate cell with micrographs of chromatids

Metaphase (from Ancient Greek μετα- (meta-) beyond, above, transcending and from Ancient Greek φάσις (phásis) 'appearance') is a stage of mitosis in the eukaryotic cell cycle in which chromosomes are at their second-most condensed and coiled stage (they are at their most condensed in anaphase).[1] These chromosomes, carrying genetic information, align in the equator of the cell between the spindle poles at the metaphase plate, before being separated into each of the two daughter nuclei. This alignment marks the beginning of metaphase.[2] Metaphase accounts for approximately 4% of the cell cycle's duration.[citation needed]

In metaphase, microtubules from both duplicated centrosomes on opposite poles of the cell have completed attachment to kinetochores on condensed chromosomes. The centromeres of the chromosomes convene themselves on the metaphase plate, an imaginary line that is equidistant from the two spindle poles.[3] This even alignment is due to the counterbalance of the pulling powers generated by the opposing kinetochore microtubules,[4] analogous to a tug-of-war between two people of equal strength, ending with the destruction of B cyclin.[5]

In order to prevent deleterious nondisjunction events, a key cell cycle checkpoint, the spindle checkpoint, verifies this evenly balanced alignment and ensures that every kinetochore is properly attached to a bundle of microtubules and is under balanced bipolar tension. Sister chromatids require active separase to hydrolyze the cohesin that bind them together prior to progression to anaphase. Any unattached or improperly attached kinetochores generate signals that prevent the activation of the anaphase promoting complex (cyclosome or APC/C), a ubiquitin ligase which targets securin and cyclin B for degradation via the proteosome. As long as securin and cyclin B remain active, separase remains inactive, preventing premature progression to anaphase.

Metaphase in cytogenetics and cancer studies

[edit]
Human metaphase chromosomes (normal male karyotype)

The analysis of metaphase chromosomes is one of the main tools of classical cytogenetics and cancer studies. Chromosomes are condensed (thickened) and highly coiled in metaphase, which makes them most suitable for visual analysis. Metaphase chromosomes make the classical picture of chromosomes (karyotype). For classical cytogenetic analyses, cells are grown in short term culture and arrested in metaphase using mitotic inhibitor. Further they are used for slide preparation and banding (staining) of chromosomes to be visualised under microscope to study structure and number of chromosomes (karyotype). Staining of the slides, often with Giemsa (G banding) or Quinacrine, produces a pattern of in total up to several hundred bands. Normal metaphase spreads are used in methods like FISH and as a hybridization matrix for comparative genomic hybridization (CGH) experiments.

Malignant cells from solid tumors or leukemia samples can also be used for cytogenetic analysis to generate metaphase preparations. Inspection of the stained metaphase chromosomes allows the determination of numerical and structural changes in the tumor cell genome, for example, losses of chromosomal segments or translocations, which may lead to chimeric oncogenes, such as bcr-abl in chronic myelogenous leukemia.

References

[edit]
  1. ^ "Chromosome condensation through mitosis". ScienceDaily. Retrieved 12 June 2007.
  2. ^ Alberts, Bruce; Hopkin, Karen; Johnson, Alexander; Morgan, David; Raff, Martin; Roberts, Keith; Walter, Peter (2019). Essential cell biology (Fifth ed.). New York London: W. W. Norton & Company. p. 632–633. ISBN 9780393680393.
  3. ^ "Metaphase plate". Biology Dictionary. Biology Online. Retrieved 9 December 2012.
  4. ^ "Metaphase". Nature Education. Retrieved 9 December 2012.
  5. ^ "The Cell Cycle". Kimball's Biology Pages. Archived from the original on 19 November 2012. Retrieved 9 December 2012.
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  • Media related to Metaphase at Wikimedia Commons