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.
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
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.
| Identifier | Panel | Sisters joined | Unattached kinetochore |
|---|---|---|---|
| unattached | An unattached sister kinetochore can sustain delay | Yes | kt-right · Sister kinetochore 2 |
| bioriented | Sister kinetochores connected towards opposite poles | Yes | No unattached kinetochore is listed in this panel |
| separated | After cohesion release: daughter chromosomes separate | No | No unattached kinetochore is listed in this panel |
| Panel | Identifier | Structure or state |
|---|---|---|
| An unattached sister kinetochore can sustain delay | left-pole | Left spindle pole |
| An unattached sister kinetochore can sustain delay | right-pole | Right spindle pole |
| An unattached sister kinetochore can sustain delay | kt-left | Sister kinetochore 1 |
| An unattached sister kinetochore can sustain delay | kt-right | Sister kinetochore 2 |
| Sister kinetochores connected towards opposite poles | left-pole | Left spindle pole |
| Sister kinetochores connected towards opposite poles | right-pole | Right spindle pole |
| Sister kinetochores connected towards opposite poles | kt-left | Sister kinetochore 1 |
| Sister kinetochores connected towards opposite poles | kt-right | Sister kinetochore 2 |
| After cohesion release: daughter chromosomes separate | left-pole | Left spindle pole |
| After cohesion release: daughter chromosomes separate | right-pole | Right spindle pole |
| After cohesion release: daughter chromosomes separate | left-daughter-chromosome | Left daughter chromosome |
| After cohesion release: daughter chromosomes separate | right-daughter-chromosome | Right daughter chromosome |
| Panel | Spindle pole identifier | Spindle pole | Kinetochore identifier | Kinetochore | Attachment relationship |
|---|---|---|---|---|---|
| An unattached sister kinetochore can sustain delay | left-pole | Left spindle pole | kt-left | Sister kinetochore 1 | Microtubule-bundle attachment |
| Sister kinetochores connected towards opposite poles | left-pole | Left spindle pole | kt-left | Sister kinetochore 1 | Microtubule-bundle attachment |
| Sister kinetochores connected towards opposite poles | right-pole | Right spindle pole | kt-right | Sister kinetochore 2 | Microtubule-bundle attachment |
| Panel | Moving chromosome identifier | Moving chromosome | Towards pole identifier | Towards spindle pole | Meaning in this model |
|---|---|---|---|---|---|
| After cohesion release: daughter chromosomes separate | left-daughter-chromosome | Left daughter chromosome | left-pole | Left spindle pole | Towards left pole |
| After cohesion release: daughter chromosomes separate | right-daughter-chromosome | Right daughter chromosome | right-pole | Right spindle pole | Towards right pole |
| Label identifier | Meaning in this model |
|---|---|
| centromere | Centromeric region of each sister |
| kinetochore | Kinetochore: protein attachment structure |
| cohesion | Sister cohesion |
| equator | Spindle equator |
| checkpoint | Unattached-kinetochore signal can delay anaphase |
| limit | Checkpoint control does not detect every attachment error |
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
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.
| Identifier | Selected cell example | Partitioning structure | Direction of furrow or plate progression |
|---|---|---|---|
| animal | Typical animal cytokinesis | Cortical actin–myosin contractile system | Inward |
| plant | Selected plant cytokinesis | Vesicle delivery and fusion build cell plate | Outward |
| Identifier | Selected cell example | Stage order | Stage description |
|---|---|---|---|
| animal | Typical animal cytokinesis | 1 | Two nuclear destinations in one cell |
| animal | Typical animal cytokinesis | 2 | Inward cleavage furrow |
| animal | Typical animal cytokinesis | 3 | Two separated cells |
| plant | Selected plant cytokinesis | 1 | Vesicles between daughter nuclei |
| plant | Selected plant cytokinesis | 2 | Cell plate expands outward |
| plant | Selected plant cytokinesis | 3 | Plate joins boundary; daughters separated |
| Label identifier | Meaning in this model |
|---|---|
| nuclei | Daughter nuclei |
| membrane | Plasma membrane |
| oldWall | Existing cell wall |
| newWall | New separating wall material |
| ring | Contractile system at cortex |
| vesicles | Membrane-and-wall-material vesicles |
| plate | Cell plate |
| separate | Cytokinesis is distinct from chromosome segregation |
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
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.
| Identifier | Panel | Displayed cell count | Scope of the DNA count | Chromosomal DNA molecules |
|---|---|---|---|---|
| before | One chromosome before copying | 1 | Whole single cell | 1 |
| predivision | Copies distributed; division site organised | 1 | Whole single cell | 2 |
| after | Two daughters, one chromosome each | 2 | Each of two daughter cells | 1 |
| Route order | From identifier | From state or compartment | To identifier | To state or compartment | Meaning of the directed route |
|---|---|---|---|---|---|
| 1 | before | One chromosome before copying | predivision | Copies distributed; division site organised | Replication, growth and segregation can overlap |
| 2 | predivision | Copies distributed; division site organised | after | Two daughters, one chromosome each | Coordinated septum and membrane division |
| Label identifier | Meaning in this model |
|---|---|
| chromosome | Selected circular chromosome |
| division | Coordinated division machinery |
| ftsz | FtsZ helps organise the division site |
| septum | Developing septum |
| scope | No 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
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.
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.
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.
Mary Ann Clark, Matthew Douglas, Jung Choi2 sources
- 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 abstract
- 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 abstract
- Molecular Biology of the Cell · Cytokinesis (4th edition, Garland Science, 2002; selected indexed text)Bruce Alberts and colleaguesPublisher-author textbook
- 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 research
- The Nucleus during Mitosis (The Cell: A Molecular Approach, 2nd edition, 2000)Geoffrey M. CooperPublisher-author textbook section · selected text
- 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 abstract
- Chromosome replication and the division cycle of Escherichia coli B/r (Journal of Molecular Biology 31:519–540, 1968)Stephen Cooper, Charles E. HelmstetterResearch abstract
- 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 abstract
- 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 abstract