Name: chromosome movement during mitosis
Uploaded: 2021-04-07T12:12:01.000Z
Duration: 2 min 11 s
Channel: JoVE

During mitosis, the bioriented chromosomes oscillate to stabilize their spindle attachments and subsequently assemble along the metaphase plate at the equator of the mitotic spindle.
Multiple forces act on chromosomes after they are attached to the mitotic spindle.
A major poleward force exerted along the kinetochore-microtubule, as a result of plus-end de-polymerization, pulls the kinetochore and its associated chromosome towards the spindle pole. At the kinetochore, Ndc80 protein complexes link the kinetochore to the microtubule through multiple low-affinity attachments along microtubule sides.
During microtubule plus-end de-polymerization, the Ndc80 attachments break and reform at new sites to maintain the kinetochore-microtubule connection. The mechanism gradually pulls the chromosome towards the spindle pole, as the microtubule shortens in length.
A second poleward force results from microtubule flux. Microtubule minus-end depolymerization generates a minus-end directed flux, causing microtubule movement towards the spindle pole. Coordinating plus-end polymerization compensates for the minus-end depolymerization, allowing microtubules to maintain their length.
A third force, &#173;the polar ejection force or polar wind, &#173;generated by kinesin-4 and 10 motor proteins pushes the chromosomes away from the spindle poles. Kinesin-4 and 10 link chromosomal arms with interpolar microtubules. These plus-end directed motor proteins move the chromosome toward the spindle equator.
A balanced interplay of these multiple opposing forces enables the bioriented chromosomes to precisely align along the metaphase plate, in preparation for chromosomal segregation.

forces acting on chromosomes

Forces Acting on Chromosomes

During mitosis, chromosome movements occur through the interplay of multiple piconewton level forces. In prometaphase, these forces help in chromosome assembly or congression at the equatorial plane, eventually leading to their alignment at the metaphase plate. The forces acting on the chromosomes are space and time-dependent; therefore, they vary with the position of the chromosomes as the cell progresses through mitosis.&#160;
Microtubules and motor proteins exert two types of forces on chromosomes&#8212;poleward and anti-poleward, also known as polar-ejection forces. Kinetochore microtubule depolymerization generates the poleward force and pulls the chromosome towards the spindle pole. In contrast, polymerization of the kinetochore-microtubule leads to polar-ejection forces, which push the chromosome towards the cell&#8217;s equator.&#160; Microtubule plus-end directed motor proteins, like chromokinesins and kinesin-7, also produce polar-ejection forces by propelling chromosomes towards the cell&#8217;s equator.
The simultaneous but unequal action of poleward and polar-ejection forces cause the oscillation of chromosomes during prometaphase; however, during metaphase, the bioriented sister chromatids experience equal but opposing forces. This creates enough tension to silence the spindle assembly checkpoint pathway and allows cells to move into anaphase. In anaphase, poleward forces act on sister chromatids, resulting in their successful segregation to the daughter cells.
In addition to the above forces, chromosomes are also subjected to cohesive and resolving forces. The cohesive force exerted by cohesin holds the sister chromatids together until the end of the metaphase. On the other hand, the resolving force generated by condensins allows chromosomes to form distinct rod-shaped structures, which helps in their proper separation during anaphase.

During mitosis, chromosome movements occur through the interplay of multiple piconewton level forces. In prometaphase, these forces help in chromosome ...

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JoVE Core

Molecular Biology

Unit 2

Cell Division

KEY TERMS AND CONCEPTS

chromosome movement during mitosis

Chromosome movement during mitosis depends on several small forces that act at different times and positions in the cell. In prometaphase, these forces help chromosomes move to the equatorial plane and line up at the metaphase plate. Because the forces change with location and stage, chromosome behavior also changes as mitosis continues.

Microtubules and motor proteins create two main kinds of force on chromosomes: poleward force and anti-poleward force, also called polar-ejection force. Poleward force comes from kinetochore microtubule depolymerization, which pulls a chromosome toward the spindle pole. Polar-ejection force comes from kinetochore-microtubule polymerization, which pushes the chromosome toward the cell’s equator.

Motor proteins also help drive chromosomes toward the middle of the cell. Plus-end directed proteins such as chromokinesins and kinesin-7 generate polar-ejection forces. During prometaphase, the unequal balance between poleward and polar-ejection forces makes chromosomes oscillate. In metaphase, the sister chromatids are bioriented, so they experience equal and opposing forces.

That balanced tension is important because it silences the spindle assembly checkpoint pathway. Once this checkpoint is off, the cell can move into anaphase. In anaphase, poleward forces act on sister chromatids and separate them into the daughter cells.

Chromosomes are also controlled by cohesive and resolving forces. Cohesin provides the cohesive force that keeps sister chromatids together until the end of metaphase. Condensins generate the resolving force, which helps chromosomes form distinct rod-shaped structures and supports their proper separation during anaphase.

Chromosome Movement During Mitosis

mitosis and cytokinesis

Mitosis and Cytokinesis Stages

Mitosis and cytokinesis are the two main steps of cell division. Mitosis is the process that divides the nucleus, while cytokinesis splits the rest of the cell. During mitosis, the cell’s genetic material is separated into two sets. This helps form two new nuclei with the same genetic information. Cytokinesis then completes division by separating the cytoplasm, which creates two daughter cells. These stages work together to make cell division possible in growing and repairing tissues.

duplication of chromatin structure

Chromatin Rebuilding During DNA Replication

Chromatin rebuilding during DNA replication helps cells copy DNA and pass on chromosome structure to daughter cells. During cell division, the genome must be disrupted and then reassembled across the whole chromosome. This process allows chromatin to be inherited, rebuilt, and maintained in new cells. The basic unit of chromatin is the nucleosome. A nucleosome contains DNA wrapped around an octamer of histone proteins, with short linker DNA between neighboring nucleosomes. The histone proteins...

cohesins

Cohesins in Meiosis and Genetic Disorders

Cohesin protein complexes act like molecular glue that keeps sister chromatids together. They are important in both mitosis and meiosis. In mitosis, cohesin complexes are removed from chromosomes before anaphase begins. Meiosis has two rounds of chromosome separation. Meiosis I separates homologous chromosomes, while Meiosis II separates sister chromatids. This process produces four daughter cells. The Meiosis I cohesin complex is made of four subunits: Smc1, Smc3, Rec8, and Scc3. Rec8...

condensins

How Condensins Shape Mitotic Chromosomes

Condensins shape chromosomes during mitosis, the stage when a cell divides. These large protein complexes use ATP, the cell’s energy molecule, to help turn loose post-interphase DNA into compact, separate chromosomes. They organize and segregate chromosomal DNA so it can be passed on correctly. Plant and animal cells contain two condensin complexes: condensin I and condensin II. Each complex has five subunits. Two are SMC, or Structural Maintenance of Chromosomes, subunits. One is a kleisin...

the mitotic spindle

How the Mitotic Spindle Separates Chromosomes

The mitotic spindle is a microtubule-based structure that helps separate chromosomes during cell division. It forms in eukaryotic cells and moves sister chromatids to opposite ends of the parent cell. This leaves each new cell nucleus with the correct genetic material. A bipolar spindle has two poles, and that shape is important for chromosome segregation. In many animal cells, two centrosomes are present when mitosis begins. Each centrosome helps organize a circular set of microtubules at one...

centrosome duplication

Centrosome Duplication in the Cell Cycle

Centrosome duplication is a key part of the animal cell cycle. The centrosome is the main microtubule organizing center, or MTOC, in animal cells. It contains two cylindrical centrioles at its core. Each centriole has nine sets of three microtubules held together by proteins. The two centrioles sit at right angles and are surrounded by the pericentriolar matrix, or PCM, a shapeless protein cloud. Duplication starts early in the cell cycle so each daughter cell can receive a centrosome after...

microtubule instability

Microtubule Growth and Shrinkage

Microtubules are hollow, cylindrical filaments in the cell. They are about 25 nm wide and can range from 200 nm to 25 μm in length. Their main building blocks are GTP-bound tubulin subunits that form αβ-heterodimers, which then assemble into protofilaments. These protofilaments join by lateral bonds to create stable microtubule cylinders. Microtubules are highly dynamic because they repeatedly assemble and break down. They can even grow on one end while shrinking on the other at the same time.

spindle assembly

How the Mitotic Spindle Forms

The mitotic spindle forms through centrosome-mediated, chromatin-mediated, and microtubule-mediated pathways. These pathways often work at the same time. Together, they build a strong spindle apparatus for cell division. In most cells, centrosomes are the main sites where microtubules begin to grow. During the G2 to prophase transition, centrosomes mature and make more microtubules. The growing microtubules extend from both centrosomes, and their plus-ends search for chromosomes by attaching...

attachment of sister chromatids

Kinetochore-Microtubule Binding in Mitosis

Kinetochore-microtubule binding helps chromosomes line up and separate during mitosis. After the nuclear envelope breaks down, condensed chromosomes are exposed to the bipolar microtubules of the mitotic spindle. The kinetochore is a large, disc-shaped protein complex at the centromere region of each sister chromatid, and it serves as the microtubule binding site. Usually, the plus-end of a single microtubule inserts into the kinetochore. In some cases, a kinetochore first makes lateral...

Chromosome movement during mitosis depends on several small forces that act at different times and positions in the cell. In prometaphase, these forces help chromosomes move to the equatorial plane and line up at the metaphase plate. Because the forces change with location and stage, chromosome behavior also changes as mitosis continues. Microtubules and motor proteins create two main kinds of force on chromosomes: poleward force and anti-poleward force, also called polar-ejection force.

separation of sister chromatids

Anaphase Start and Chromatid Release

Anaphase starts when sister chromatids are released from one another and pulled toward opposite spindle poles. Before that happens, cohesion along the chromosomal arms is reduced from prophase to metaphase, which helps the sister chromatids resolve. A residual cohesin link still holds them together until the metaphase-to-anaphase transition. This leftover connection helps prevent early separation and reduces the risk of aneuploidy, or the wrong number of chromosomes, in daughter cells. At the...

the spindle assembly checkpoint

How Cells Safeguard Chromosome Separation

The spindle assembly checkpoint helps cells safeguard chromosome separation during anaphase. It acts as a molecular surveillance system. The checkpoint checks whether the steps needed for chromosome segregation are complete before the cell moves on. Many proteins work together to control this checkpoint. If mutations affect these proteins, a cell may enter anaphase too early. Chromosomes can then separate before the needed steps are finished. When this happens, the daughter cells may receive...

anaphase a and b

Chromatid Movement by Microtubule Forces

Chromatid movement during anaphase depends on microtubule forces at the spindle. Kinetochore microtubules grow from the spindle poles, and their plus ends attach to kinetochores on sister chromatids. The kinetochore protein complex Ndc80 makes low-affinity links with the microtubule plus end.

the contractile ring

How Cells Build the Contractile Ring

Contractile rings are built from microfilaments and help separate daughter cells during cytokinesis. This ring forms as the cell cycle moves forward, but the exact timing and coordination with other cell cycle events are still not fully understood. RhoA, a small GTPase in the Ras superfamily, controls contractile ring assembly and function. The inactive form of RhoA is bound to GDP. A guanine nucleotide exchange factor, or Rho-GEF, is located in the cortex region, where the next cell division...

determining the plane of cell division

How Cells Set the Division Plane

How cells set the division plane during cell division affects daughter cell size and, in many cases, cell fate. The plane of division is perpendicular to the plane of chromosome segregation. Different organisms use different cues to place it, based on their shape and needs. In animal cells, the cleavage furrow forms along the division plane from the cell cortex, the area just below the plasma membrane. The mitotic spindle is the main guide for this position. If the spindle axis changes, the...

the phragmoplast

How Plant Cells Build a New Wall

Plant cells build a new wall during cytokinesis with help from the phragmoplast. This plant structure forms from the central spindle and related cell parts as the cell finishes dividing. In animal cells, the comparable structure is the midbody, but in plants the phragmoplast expands outward to guide new wall formation. The mature phragmoplast has a donut-like shape. It has an outer leading region, a middle transition zone, and an inner lagging region. New microtubules are assembled...

distribution of cytoplasmic content

How Organelles Are Split During Cytokinesis

Cytokinesis splits a cell’s chromosomes and organelles into two daughter cells. Before cell division, organelles grow and divide. They cannot be made from scratch, so each daughter cell must inherit at least one copy to survive. Many details of this organelle sorting process are still not fully known. The cytoplasm contains organelles, salts, proteins, and water. Small organelles such as peroxisomes and lysosomes can be shared by simple diffusion. Because cells contain many copies of these...

anaphase promoting complex

APC Activates Chromosome Separation

The anaphase-promoting complex, or APC, helps control cell cycle progression by destroying specific proteins at the right time. These proteins are tagged with ubiquitin by ubiquitin ligases and then broken down by the proteasome. The SCF (Skp1/Cullin/F-box) complex and APC are two major ubiquitin ligases involved in this process. SCF stays active throughout the cell cycle, but APC turns on during the metaphase-to-anaphase transition. APC becomes active when Cdc20 or Cdh1 binds to it. These...

key terms and concepts

Learn how poleward and polar-ejection forces move chromosomes during mitosis, align sister chromatids at metaphase, and support separation in anaphase.

Chromosome Movement During Mitosis | JoVE K12

molecular biology

JoVE Core Molecular Biology showcases the molecular processes of the cell through clear and concise animated video lessons. Additionally, in scientist-in-action videos, these concepts' application is demonstrated through real experiments in modern laboratories.

Chromosome Movement During Mitosis

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