Cell Cycle and Cell Division

Cell division is a very important process, during this process DNA replication and cell growth
also takes place, they have to take place in a coordinated way to ensure correct division and
formation of progeny cells containing intact genomes. The sequence by which a cell
duplicates its genome, synthesizes other constituents of cell and eventually divides into two
daughter cells is called cell cycle.

Phases of Cell Cycle

A typical eukaryotic cell divides once in every 24 hours. However, the duration varies from
organism to organism and from cell type to cell type. E.g., Yeast can complete the same cell
cycle in 90 minutes. The cell cycle has two phases:

 Interphase
 M Phase (Mitosis Phase)
The M phase is the phase when actual cell division or mitosis occurs and interphase is the
phase between two successive M phases.
M phases starts with nuclear division corresponding to separation of daughter
chromosomes(karyokinesis) and ends at division of cytoplasm(cytokinesis). The interphase is
the resting time/phase during which the cell prepares for further division by undergoing
DNA replication and cell growth in an orderly manner.
The interphase has three phases:

 G1 phase (Gap-1)
 S phase (Synthesis)
 G2 phase (Gap-2)
G1 phase corresponds to interval between mitosis and initiation of DNA replication, during
this phase the cell is metabolically active and grows continuously but does not replicate
DNA.
S phase marks the initiation of synthesis of DNA replication, when it starts the DNA per cell
doubles. However, the DNA number does not increase, even after S phase the number of
chromosomes remain the same. In animal cells, DNA replication begins in the nucleus and
the centriole duplicates in the cytoplasm.
During G2 phase, proteins are synthesized for mitosis while cell growth continues. Some
cells don’t exhibit division(heart cells) while some and many other cells divide occasionally
in order to replace older cells. These cells do not divide after G1 stage and enter an inactive
stage called quiescent stage (G0 phase).

M phase

Karyokinesis
This is the phase in which almost all components of the cells are completely replaced, it
shows a drastic change. Since number of chromosomes remain same, it is called equational
division. Mitosis has been divided into four stages of nuclear division(karyokinesis).

 Prophase
 Metaphase
 Anaphase
 Telophase
Prophase: It is the first stage of karyokinesis of mitosis, it follows the S and G2 phase. In the
S and G2 phase, the new DNA molecules are not distinct but intertwined. Prophase marks
the condensation of chromosomal material; it becomes untangled during process of
chromatin condensation. The centrosome which had divided during S phase starts to move
towards the opposite poles of the cell.
Thus, Prophase is marked by two characteristic events:

 Chromosomal material condenses to form two compact mitotic chromosomes.

Chromosomes are composed of two chromatids attached together at the
centromere.
 Centrosome which had gone under duplication begins to move towards the poles of
the cell. Each centrosome radiates microtubules called asters. These two asters form
the mitotic apparatus.

Metaphase: The complete disintegration of the nuclear envelope marks the second step
in karyokinesis called metaphase such that the chromosomes are spread all around the
cell. At this stage, metaphase chromosomes are made of two sister chromatids which
are held together by centromere. Small-disc shaped structures at the surface of the
centromere are called kinetochores. These structures serve as sites of attachment of
spindle fibers to the chromosomes that are moved into position at the center of the cell.
The plane of alignment of the chromosomes connected by its sister chromatid is called
the metaphase plate. Key features of metaphase are:

 Spindle fibers attach to kinetochores of chromosomes.

 Chromosomes are moved to the spindle equator and get aligned along
metaphase plate through spindle fibers to both poles.
Anaphase: At the onset of anaphase, each chromosome arranged at the metaphase plate is
split simultaneously and now these sister chromatids are called daughter chromosomes. Of
future daughter nuclei.
As each chromosome moves away from the equatorial plate, the centromere of each
chromosome moves away from the equatorial plate but remain directed towards the pole
and connected to the astral.
Thus, anaphase is characterized by:

 Centromeres split and chromatids separate.

 Chromatids move to opposite poles.
Telophase: At the beginning of the final stage of karyokinesis, the chromosomes condense
and lose their individuality, the individual chromosomes cannot be seen and each set of
chromatin material is found at the poles. Thus, the key features are:

 Chromosomes collect at spindle poles and lose their individuality.

 Nuclear envelope develops around the chromosomes and each one forms a
daughter nucleus.
 Nucleolus, Golgi Apparatus and Endoplasmic Reticulum is formed.

Cytokinesis
Mitosis accomplishes not only the segregation of duplicated chromosomes into daughter
nuclei (karyokinesis), but the cell itself is divided into two daughter cells by the separation of
cytoplasm called cytokinesis at the end of which cell division gets completed.
In an animal cell, this is achieved by the appearance of a furrow in the plasma membrane.
The furrow gradually deepens and ultimately joins in the centre dividing the cell cytoplasm
into two.
In plant cells, wall formation starts in the centre of the cell and grows outward to meet the
existing lateral walls. The formation of the new cell wall begins with the formation of a
simple precursor, called the cell-plate that represents the middle lamella between the walls
of two
adjacent cells.
At the time of cytoplasmic division, organelles like mitochondria and plastids get distributed
between the
two daughter cells. In some organisms karyokinesis is not followed by cytokinesis as a result
of which multinucleate condition arises leading to the formation of syncytium (e.g., liquid
endosperm in coconut).

Significance of Mitosis
Mitosis or equational division occurs in diploid cells. It also takes place in some haploid cells,
mitosis results in two diploid daughter cells with identical genetic complement.
Mitosis results in cell growth
In multicellular organisms, it results in cell growth and maintains the nucleo-cytoplasmic
ratio.
It helps repair the cells in our body, if damaged. The cells of upper layer of epidermis, lining
of the gut and blood cells are constantly replaced.

Meiosis
Production of offspring by sexual reproduction includes fusion of two gametes, each with a
haploid set of chromosomes. These gametes are formed from diploid cells. This special kind
of division in which the chromosomal number reduces to half id called meiosis, it results in
formation of daughter haploid cells. Meiosis is a process used in plants and animals during
gametogenesis.
Features of Meiosis:
 Meiosis involves two sequential cycles of nuclear and cell division called meiosis-I
and meiosis-II.
 Meiosis is initiated after formation of identical sister chromatids have been formed
in the S phase.
 Meiosis involves pairing of homologous chromosomes and recombination of non-
sister chromatids of homologous chromosomes.
 Four haploid cells are formed after meiosis.

Meiosis-I
Prophase-I: Prophase of meiosis is complex and ling as compared to prophase of mitosis. It
has been further divided into five phases: Leptotene, Zygotene, Pachytene, Diplotene and
Diakinesis.
During leptotene, the chromosome starts to become visible, the compaction of
chromosome continues during leptotene. During zygotene, the chromosomes start to pair
and this process of association is called synapsis. It is also accompanied by formation of
structures called synaptonemal complex. The complex formed by a pair of chromosomes is
called a bivalent or a tetrad.
The first two stages of prophase-I are rather short lived compared to the next one called
pachytene. During this stage, the four chromatids become distinct and clearly visible and
appear as tetrads. This stage is characterized by formation of recombination nodules. These
are the sites at which crossing over occurs, it is the exchange of genetic material between
two homologous chromosomes, it is mediated by an enzyme called recombinase.
Recombination is completed by the end of pachytene, leaving chromosomes at the sites of
crossing over. The initiation of diplotene is marked by the dissolution of synaptonemal
complex and tendency of homologous bivalents to separate from each other. These X-
shaped structures are called chiasmata.
The final stage of meiotic prophase I is diakinesis. This is marked by terminalisation of
chiasmata. During this phase the chromosomes are fully condensed and the meiotic spindle
is assembled to prepare the homologous chromosomes for separation. By the end of
diakinesis, the nucleolus disappears and the nuclear envelope also breaks down. Diakinesis
represents transition to metaphase.
Metaphase-I: The bivalent chromosomes align to the equatorial plate, the microtubules of
the opposite poles of the spindle attach to the kinetochore of homologous chromosomes.
Anaphase-I: The homologous chromosomes separate while the sister chromatids remain
associated at their centromeres.
Telophase-I: The nuclear membrane and nucleolus reappear; cytokinesis follows and this is
called as dyad of cells. The stage between two meiotic divisions is called interkinesis, there is
no DNA replication in this process and this process is short lived.

Meiosis-II
Prophase-II: Meiosis II is initiated immediately after cytokinesis, usually before the
chromosomes have fully elongated. In contrast to meiosis I, meiosis II resembles a normal
mitosis. The nuclear membrane disappears by the end of prophase II. The chromosomes
again become compact.
Metaphase-II: At this stage the chromosomes align at the equator and the microtubules from
opposite poles of the spindle get attached to the kinetochores of sister chromatids.
Anaphase-II: It begins with the simultaneous splitting of the centromere of each chromosome
(which was holding the sister chromatids together), allowing them to move toward opposite
poles of the cell by shortening of microtubules attached to kinetochores.
Telophase-II: Meiosis ends with telophase II, in which the two groups of chromosomes once
again get enclosed by a nuclear envelope; cytokinesis follows resulting in the formation of
tetrad of cells i.e., four haploid daughter cells.

Significance of Meiosis
Meiosis is the mechanism by which conservation of specific chromosome number of each
species is achieved across generations in sexually reproducing organisms, even though the
process, per se, paradoxically, results in reduction of chromosome number by half. It also
increases the genetic variability in the population of organisms from one generation to the
next. Variations are very important for the process of evolution.