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Biology · Cell Architecture, Growth & Division

Chromosome segregation, mitosis and cytokinesis

Copying the chromosomes is only preparation. A dividing cell must distribute the copies so each destination receives the intended complement, then partition its cytoplasm. A picture of aligned chromosomes shows one stage of that work, not proof that copying and division have both succeeded.

The cells chapter introduces membranes and compartments; the DNA chapter distinguishes a DNA molecule from a chromosome and its replicated sister chromatids. Gene expression and signalling explain how proteins and regulatory responses arise. Here, nuclear DNA counts exclude organelle DNA. Every numerical movement, chromosome set and comparison case is an original teaching model, rather than a measured specimen.

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Explain sister-chromosome separation and cytoplasmic division, including the limits of attachment control and differences among division systems.

01

Mitosis separates sister copies

In a typical animal-cell mitosis, chromosomes condense, the nuclear envelope breaks down and a microtubule spindle interacts with the replicated chromosomes. Kinetochores are protein structures assembled at centromeric regions of the chromatids. They connect chromosomes to spindle microtubules and participate in regulatory signals. A kinetochore is not an entire chromosome or the DNA sequence of a centromere.

For ordinary mitotic segregation, the two sister kinetochores become attached towards opposite spindle poles. Chromosomes commonly align near the spindle's middle at metaphase. Release of sister cohesion permits anaphase separation; each separated chromatid is now called a daughter chromosome. Mitosis distributes sisters, rather than separating one whole homologous set from the other as in the first meiotic division.

Microtubule dynamics, motors and other interactions generate the movements. Unattached kinetochores can maintain signals that delay anaphase. Attachment and error-correction systems reduce segregation errors, but they are not perfect guarantees: some incorrect attachments can escape the checkpoint. A visually centred chromosome therefore does not establish correct connections on both sisters.

Attachment geometry matters as well as position

Attachment geometry matters as well as positionThree panels show one representative replicated chromosome in a larger mitotic spindle. First, its two distinct sister kinetochores are visible, with one unattached. Second, each sister kinetochore has a microtubule-bundle connection to the opposite spindle pole, with the joined sisters near the equator. Third, cohesion is released and the two daughter chromosomes move towards opposite poles. A checkpoint-delay callout belongs to the unattached panel, while a separate limitation note says some incorrect attachments evade control.An unattached sister kinetochore can sustain delay12341Left spindle pole2Right spindle pole3Sister kinetochore 14Sister kinetochore 2Microtubule-bundle attachmentUnattached-kinetochore signal can delay anaphaseSister kinetochores connected towards opposite poles12341Left spindle pole2Right spindle pole3Sister kinetochore 14Sister kinetochore 2Microtubule-bundle attachmentAfter cohesion release: daughter chromosomes separate12341Left spindle pole2Right spindle pole3Left daughter chromosome4Right daughter chromosomeTowards left poleTowards right poleCentromeric region of each sisterKinetochore: protein attachment structureSister cohesionSpindle equatorCheckpoint control does not detect every attachment error
One representative chromosome, not the complete 2n = 4 count diagram. Each connection denotes a bundle/relationship rather than a universal single microtubule. No exact attachment forces, duration or error probability.
Read the diagram in words

Three panels show one representative replicated chromosome in a larger mitotic spindle. First, its two distinct sister kinetochores are visible, with one unattached. Second, each sister kinetochore has a microtubule-bundle connection to the opposite spindle pole, with the joined sisters near the equator. Third, cohesion is released and the two daughter chromosomes move towards opposite poles. A checkpoint-delay callout belongs to the unattached panel, while a separate limitation note says some incorrect attachments evade control.

Sister cohesion and attachment state
IdentifierPanelSisters joinedUnattached kinetochore
unattachedAn unattached sister kinetochore can sustain delayYeskt-right · Sister kinetochore 2
biorientedSister kinetochores connected towards opposite polesYesNo unattached kinetochore is listed in this panel
separatedAfter cohesion release: daughter chromosomes separateNoNo unattached kinetochore is listed in this panel
Panel structure identities
PanelIdentifierStructure or state
An unattached sister kinetochore can sustain delayleft-poleLeft spindle pole
An unattached sister kinetochore can sustain delayright-poleRight spindle pole
An unattached sister kinetochore can sustain delaykt-leftSister kinetochore 1
An unattached sister kinetochore can sustain delaykt-rightSister kinetochore 2
Sister kinetochores connected towards opposite polesleft-poleLeft spindle pole
Sister kinetochores connected towards opposite polesright-poleRight spindle pole
Sister kinetochores connected towards opposite poleskt-leftSister kinetochore 1
Sister kinetochores connected towards opposite poleskt-rightSister kinetochore 2
After cohesion release: daughter chromosomes separateleft-poleLeft spindle pole
After cohesion release: daughter chromosomes separateright-poleRight spindle pole
After cohesion release: daughter chromosomes separateleft-daughter-chromosomeLeft daughter chromosome
After cohesion release: daughter chromosomes separateright-daughter-chromosomeRight daughter chromosome
Spindle attachment relationships
PanelSpindle pole identifierSpindle poleKinetochore identifierKinetochoreAttachment relationship
An unattached sister kinetochore can sustain delayleft-poleLeft spindle polekt-leftSister kinetochore 1Microtubule-bundle attachment
Sister kinetochores connected towards opposite polesleft-poleLeft spindle polekt-leftSister kinetochore 1Microtubule-bundle attachment
Sister kinetochores connected towards opposite polesright-poleRight spindle polekt-rightSister kinetochore 2Microtubule-bundle attachment
Separated chromosome movement
PanelMoving chromosome identifierMoving chromosomeTowards pole identifierTowards spindle poleMeaning in this model
After cohesion release: daughter chromosomes separateleft-daughter-chromosomeLeft daughter chromosomeleft-poleLeft spindle poleTowards left pole
After cohesion release: daughter chromosomes separateright-daughter-chromosomeRight daughter chromosomeright-poleRight spindle poleTowards right pole
Diagram terms and qualifications
Label identifierMeaning in this model
centromereCentromeric region of each sister
kinetochoreKinetochore: protein attachment structure
cohesionSister cohesion
equatorSpindle equator
checkpointUnattached-kinetochore signal can delay anaphase
limitCheckpoint control does not detect every attachment error
02

Dividing the cytoplasm is a separate event

Nuclear reorganisation follows segregation, while cytokinesis partitions cytoplasm and completes physical separation. In typical animal cells, an actin-and-myosin contractile system at the cortex contributes to an inward cleavage furrow. This is distinct from the spindle's chromosome-separation work.

In many plant cells, vesicles carrying membrane and wall material assemble a cell plate between the daughter nuclei. The plate expands outward and joins the cell boundary, helping establish new plasma membranes and a wall between daughters. It is not an animal-style furrow pinching through an existing rigid wall. The phragmoplast, a cytoskeletal array between forming daughters, helps organise delivery; its details are omitted from the simple comparison.

Mitosis can occur without completed cytokinesis, leaving two nuclei in one cell. Division systems also vary among eukaryotes: for example, not all mitoses break down the nuclear envelope as the animal example does. The intended daughter chromosome complements do not guarantee identical cell sizes, organelle distributions or later cell fates.

Partition cytoplasm through different structures

Partition cytoplasm through different structuresAn animal-cell row shows two already segregated nuclear groups and a cortex-associated actin/myosin ring driving an inward furrow, followed by two cell boundaries. A plant-cell row shows two daughter nuclei inside an existing wall, vesicles accumulating between them, then an expanding central cell plate joining the cell boundary. The plant row's new daughter plasma membranes lie on either side of new wall material. Arrows point inward for furrow closure and outward for cell-plate expansion.Typical animal cytokinesisCortical actin–myosin contractile systemTwo nuclear destinations in one cellInward cleavage furrowTwo separated cellsSelected plant cytokinesisVesicle delivery and fusion build cell plateVesicles between daughter nucleiCell plate expands outwardPlate joins boundary; daughters separatedDaughter nucleiPlasma membraneExisting cell wallNew separating wall materialContractile system at cortexMembrane-and-wall-material vesiclesCell plateCytokinesis is distinct from chromosome segregation
Original mechanism comparison after intended nuclear segregation. It omits detailed abscission and phragmoplast reactions and does not represent every eukaryotic lineage or require equal-sized daughters.
Read the diagram in words

An animal-cell row shows two already segregated nuclear groups and a cortex-associated actin/myosin ring driving an inward furrow, followed by two cell boundaries. A plant-cell row shows two daughter nuclei inside an existing wall, vesicles accumulating between them, then an expanding central cell plate joining the cell boundary. The plant row's new daughter plasma membranes lie on either side of new wall material. Arrows point inward for furrow closure and outward for cell-plate expansion.

Selected cytokinesis mechanisms
IdentifierSelected cell examplePartitioning structureDirection of furrow or plate progression
animalTypical animal cytokinesisCortical actin–myosin contractile systemInward
plantSelected plant cytokinesisVesicle delivery and fusion build cell plateOutward
Cytokinesis stage order
IdentifierSelected cell exampleStage orderStage description
animalTypical animal cytokinesis1Two nuclear destinations in one cell
animalTypical animal cytokinesis2Inward cleavage furrow
animalTypical animal cytokinesis3Two separated cells
plantSelected plant cytokinesis1Vesicles between daughter nuclei
plantSelected plant cytokinesis2Cell plate expands outward
plantSelected plant cytokinesis3Plate joins boundary; daughters separated
Diagram terms and qualifications
Label identifierMeaning in this model
nucleiDaughter nuclei
membranePlasma membrane
oldWallExisting cell wall
newWallNew separating wall material
ringContractile system at cortex
vesiclesMembrane-and-wall-material vesicles
plateCell plate
separateCytokinesis is distinct from chromosome segregation
03

Binary fission does not require a mitotic nucleus

Binary fission is a common mode of bacterial reproduction. In a simplified one-chromosome example, DNA is replicated and its copies are distributed while the cell grows; division machinery forms a septum that partitions the cell. There is no membrane-enclosed nucleus undergoing mitosis. Replication, segregation and constriction can overlap rather than wait in three entirely separate time blocks.

In many bacteria, FtsZ protein assemblies help organise the division site and coordinate other machinery, including wall synthesis. FtsZ alone is not a complete membrane-pinching machine. Bacterial chromosome number, shape and replication timing vary, and archaeal division systems are not all the same as this bacterial example. The small diagram assumes one circular chromosome and one copying round; it is not a universal prokaryotic cycle.

A selected bacterial fission cycle

A selected bacterial fission cycleThree rod-shaped bacterial outlines show one circular chromosome, an enlarged cell with two replicated chromosome circles at different locations and a central division site, then two daughter cells each with one circle. The division site is labelled as a coordinated machinery system, with FtsZ assemblies helping organise it and wall/membrane changes completing separation. There is no nucleus or mitotic spindle in this model.One chromosome before copyingReplication, growth and segregation can overlapCopies distributed; division site organisedCoordinated septum and membrane divisionTwo daughters, one chromosome eachSelected circular chromosomeCoordinated division machineryFtsZ helps organise the division siteDeveloping septumNo mitotic nucleus in this bacterial model
Simplified one-circular-chromosome example with a completed copying round. It omits multifork replication, plasmids, detailed segregation and varied bacterial/archaeal division mechanisms. FtsZ is not portrayed as the only machinery or a universal prokaryotic feature.
Read the diagram in words

Three rod-shaped bacterial outlines show one circular chromosome, an enlarged cell with two replicated chromosome circles at different locations and a central division site, then two daughter cells each with one circle. The division site is labelled as a coordinated machinery system, with FtsZ assemblies helping organise it and wall/membrane changes completing separation. There is no nucleus or mitotic spindle in this model.

Bacterial cell and chromosomal DNA counts
IdentifierPanelDisplayed cell countScope of the DNA countChromosomal DNA molecules
beforeOne chromosome before copying1Whole single cell1
predivisionCopies distributed; division site organised1Whole single cell2
afterTwo daughters, one chromosome each2Each of two daughter cells1
Selected directed routes
Route orderFrom identifierFrom state or compartmentTo identifierTo state or compartmentMeaning of the directed route
1beforeOne chromosome before copyingpredivisionCopies distributed; division site organisedReplication, growth and segregation can overlap
2predivisionCopies distributed; division site organisedafterTwo daughters, one chromosome eachCoordinated septum and membrane division
Diagram terms and qualifications
Label identifierMeaning in this model
chromosomeSelected circular chromosome
divisionCoordinated division machinery
ftszFtsZ helps organise the division site
septumDeveloping septum
scopeNo mitotic nucleus in this bacterial model

Worked example

State whether the count is per cell or per destination

Our diploid 2n = 4 model enters mitosis with four replicated chromosomes, eight chromatids and eight DNA molecules, totalling 4C. When sisters separate in anaphase, the still-undivided cell contains eight daughter chromosomes and eight DNA molecules. Each intended pole receives four chromosomes and 2C. Separation changes the chromosome counting units, not the total DNA amount.

After cytokinesis, each daughter cell has four unreplicated chromosomes, four DNA molecules and 2C, retaining diploidy. If both sisters of one chromosome instead reach the same pole while all other sisters segregate normally, destinations contain five and three chromosomes. That is an unequal chromosome complement, not ordinary conversion into complete triploid and haploid sets. No error frequency or DNA mass ratio is inferred.

Check your understanding

  1. Does the temporary count of eight daughter chromosomes in the undivided model's anaphase mean eight chromosomes will normally enter each daughter cell?

    Show reasoning

    No. Eight is the total across both destinations in one still-undivided cell. Normal segregation gives four to each daughter.

  2. A replicated chromosome appears at the spindle equator. Does that alone prove that its sister kinetochores are correctly attached?

    Show reasoning

    No. Position and attachment geometry are different observations. Correct biorientation and regulatory evidence matter, and some attachment errors can evade checkpoint control.

  3. Two nuclei are present inside one continuous cell boundary. Has cytokinesis necessarily finished?

    Show reasoning

    No. Nuclear division and cytoplasmic separation are distinct. The cell may have completed nuclear segregation without completing cytokinesis.

What to carry forward

Replication supplies copies; spindle-mediated segregation distributes them; cytokinesis partitions the cell. Count each stage and destination explicitly.

References

Resources supporting this lesson’s explanations and examples.

  1. Mary Ann Clark, Matthew Douglas, Jung Choi2 sources
    1. Biology 2e · The Cell Cycle (OpenStax, 2018)Mary Ann Clark, Matthew Douglas, Jung ChoiTextbookopenstax.orgRead source
    2. Biology 2e · Prokaryotic Cell Division (OpenStax, 2018)Mary Ann Clark, Matthew Douglas, Jung ChoiTextbookopenstax.orgRead source
  2. The checkpoint delaying anaphase in response to chromosome monoorientation is mediated by an inhibitory signal produced by unattached kinetochores (Journal of Cell Biology 130:941–948, 1995)Conly L. Rieder, Richard W. Cole, Alexey Khodjakov, Greenfield SluderResearch abstractrupress.orgRead source
  3. Merotelic kinetochore orientation is a major mechanism of aneuploidy in mitotic mammalian tissue cells (Journal of Cell Biology 153:517–527, 2001)Daniela Cimini, Bonnie Howell, Paul Maddox, Alexey Khodjakov, Francesca Degrassi, Edward D. SalmonResearch abstractpubmed.ncbi.nlm.nih.govRead source
  4. Molecular Biology of the Cell · Cytokinesis (4th edition, Garland Science, 2002; selected indexed text)Bruce Alberts and colleaguesPublisher-author textbookncbi.nlm.nih.govRead source
  5. Treadmilling by FtsZ filaments drives peptidoglycan synthesis and bacterial cell division (Science 355:739–743, 2017; abstract and selected author text)Alexandre W. Bisson-Filho and colleaguesOriginal researchpubmed.ncbi.nlm.nih.govRead source
  6. The Nucleus during Mitosis (The Cell: A Molecular Approach, 2nd edition, 2000)Geoffrey M. CooperPublisher-author textbook section · selected textncbi.nlm.nih.govRead source
  7. A role for the ESCRT system in cell division in archaea (Science 322:1710–1713, 2008)Rachel Y. Samson, Takayuki Obita, Stefan M. Freund, Roger L. Williams, Stephen D. BellResearch abstractpubmed.ncbi.nlm.nih.govRead source
  8. Chromosome replication and the division cycle of Escherichia coli B/r (Journal of Molecular Biology 31:519–540, 1968)Stephen Cooper, Charles E. HelmstetterResearch abstractsciencedirect.comRead source
  9. Genome structure of the genus Azospirillum (Journal of Bacteriology 182:4113–4116, 2000; indexed abstract)Claudia C. G. Martin-Didonet, Leda S. Chubatsu, Emanuel M. Souza, Margareth Kleina, Fabiane G. M. Rego, Liu U. Rigo, M. Geoffrey Yates, Fabio O. PedrosaResearch abstractpubmed.ncbi.nlm.nih.govRead source
  10. Dynamics of Escherichia coli chromosome segregation during multifork replication (Journal of Bacteriology 189:8660–8666, 2007)Henrik J. Nielsen, Brenda Youngren, Flemming G. Hansen, Stuart AustinResearch abstractpubmed.ncbi.nlm.nih.govRead source