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CELL CYCLE And CELL DIVISION

The cell cycle is the life cycle of a cell. During its life cycle, cell
grow, perform its assign functions, replicate, and divide. For
eukaryotic cells, the cell cycle consists of two general phases:
interphase and the mitotic phase. Cell division is part of the
life cycle for almost every cell. It is a more complicated process
in eukaryotic than prokaryotic cells. Eukaryotes have multiple
chromosomes and many organelles, all of which must be duplicated
and separated for the cell to divide.

Cell Cycle
yy The sequence of events by which a cell duplicates its
genome, synthesises other cell constituents, and eventually
divides into two daughter cells is known as cell cycle.
yy The events of the cell cycle are under genetic control.

Phases of Cell Cycle

yy Duration of the cell cycle varies from organism to organism,
and from cell to cell.
yy Duration of the cell cycle in humans is 24 hrs, and in yeast
is 90 min.
yy Cell cycle consists of two phases:
Interphase (Non-dividing phase)
M phase (Dividing phase)

1. Interphase:
yy The interval between two successive cell divisions is termed
as Interphase.

yy Interphase involves a series of changes that prepare a cell

for division.
yy It is the period during which the cell experiences growth
and DNA replication in an orderly manner.
yy Interphase is the longest period in the cell cycle and occupies
more than 95% of the duration of the cell cycle.
yy It is divided into 3 phases: G1 phase, S phase and G2
phase
G1 (Gap 1) phase: It is the first phase of the growth. It
is the stage during which the cell grows and prepares
its DNA for replication. In this phase, the cell is
metabolically active. A lot of protein and RNA are
synthesized during this phase. A large number of
nucleotides, amino acids for histone synthesis and
energy rich compounds are formed.
S phase (Synthetic phase): It is the stage during which
DNA synthesis occurs. In this phase, the amount of
DNA (per cell) doubles, but the chromosome number
remains the same. Each chromosome now consists of
two sister chromatids. The sister chromatids are held
together by a structure called a centromere. The cell is
now committed to division.
G2 phase: In this phase, the cell continues to grow
and prepares itself for division. The proteins and RNA
required for mitosis are synthesised during this stage.

Cell cycle arrest

Fig. Cell cycle

2
G0 or quiescent phase: Is the stage wherein cells remain
metabolically active, but do not proliferate unless called
to do so. Such cells are used for replacing the cells lost
during injury.
yy Interphase is followed by a shorter phase of the cell cycle
called the mitotic phase.
M Phase
yy A cell reproduces during the mitotic phase. The mitotic
phase has two stages: Karyokinesis and Cytokinesis.
Karyokinesis: It involves nuclear division, producing
two daughter nuclei.
Cytokinesis: It involves cell division, producing two
roughly equal cells called as daughter cells.
yy The mitotic (M) phase of the cell cycle alternates with the
much longer interphase.

Length of a Cell Cycle

yy The time it takes a cell to complete the cell cycle depends
on the type of cell that is dividing.
yy Most of the cells in the human body can complete the cell
cycle in about 24 hours.
yy Of this time:
The M phase would occupy < 1 hour
The S phase would last 10-12 hours (~ the cell
cycle)
The G2 phase typically takes 4-6 hours
The G1 phase usually occupies 5-6 hours. This phase
is, however, the most variable in length in different
types of cells

MITOSIS (also known as equational division)

yy Mitosis was first discovered in animal cells by Walter
Fleming (1878). He also coined the term mitosis.
yy Strasburger (1879) identified mitosis in plant cells
yy Mitosis is called somatic cell division since it occurs most
commonly in vegetative cells or somatic cells.
yy In plants, active centres of mitosis are root apex and stem
apex.
yy The best region to observe mitosis in plants is root apex.
yy Mitosis is also called equational division as number of
chromosomes in parent and progeny remain the same.
yy Mitosis is the process of cell division wherein the
chromosomes replicate and get equally distributed into two
daughter cells. The chromosome number in each daughter
cell is equal to that in the parent cell, i.e., diploid. Hence,
mitosis is known as equational division.
Phases of Mitosis
yy Mitosis (M phase) is divided into 4 stages: Prophase
(1st stage), Metaphase, Anaphase, Telophase (Last stage)
yy Cytokinesis completes the mitotic phase.

Fig. Diagrammatic representation of Interphase and M

phase

1. PROPHASE: It can be divided into three stages:

(i) Early prophase
Centrioles start moving towards opposite poles of the
nucleolus cell. Both the centriole pairs radiate out fine
microtubular fibrils called astral rays. Each pair of astral
rays along with centriole pair is called aster.
The chromatin reticulum disappears.
Chromosomes appear as long threads.
Nucleus become less distinct
(ii) Middle prophase
The chromatin condenses into chromosomes by
dehydrating and coiling.
The chromosomes consist of two identical sister
chromatids, joined together by a centromere. For the
first time, they can be seen with a light microscope.
(iii) Late prophase
A structure known as the centrosome duplicates itself to
form two daughter centrosomes that migrate to opposite
ends of the cell.
The centrosomes organise the production of microtubules
that form the spindle fibres that constitute the mitotic
spindle.
The nuclear membrane breaks down to form a number
of small vesicles and the nucleolus disintegrates.

Fig. Prophase

3
2. Metaphase
(i) Pro-metaphase
The chromosomes, led by their centromeres, migrate to
the equatorial plane in the midline of cell. This region of
the mitotic spindle is known as the metaphase plate.
Each chromatid now has a specialized protein structure
located at the centromere called a kinetochore through
which spindle fibres attach.
The chromosomes continue to condense.
(ii) Metaphase
It is the longest phase of the cell cycle. It last for about
20 minutes.
The spindle fibres pull and push the duplicated
chromosomes to the middle of the cell during
metaphase.
Notice in the figure that the chromosomes line up along
the middle of the cell. This makes sure that each new
cell will receive one copy of each chromosome.

3. Anaphase
It is the shortest stage of mitosis.
It has two sub stages, A and B. Both anaphase A and
anaphase B contribute to the movement of chromosome
toward the spindle poles
Anaphase A:
In anaphase A, chromosomes are pulled pole ward.
In anaphase A, the centromere divide, separating the
two sister chromatids from each other.
Each chromatid is pulled toward the pole along the path
of their chromosome fibres.
During anaphasic movement of chromosomes,
centromeres lead the arms.
As a result of movement, chromosomes acquire V, L, J,
I shapes based on the position of centromere.
Once they are separated, the sister chromatids are now
referred to as daughter chromosomes.
Anaphase B:
In anaphase B, poles move apart.
As the single-stranded chromosomes move to opposite
sides of the cell, the cell begins to get longer.
Anaphase ends when the two sets of identical
chromosomes reach opposite ends of the cell.
Most spindle fibres disappear from near the poles but
remain intact near the middle.
The number and type of chromosomes at each pole is
same as present in the parent nucleus.
It is the alignment and separation in metaphase and
anaphase that is important in ensuring that each daughter
cell receives a copy of every chromosome.

Fig. Metaphase
Anaphase A

Anaphase B

Fig. Anaphase

4
4. Telophase
yy During telophase, the spindle fibres that helped divide
chromosomes begin to disappear.
yy The chromosomes begin to uncoil and form chromatin
which together appears as reticulum.
yy Nucleolus becomes visible again.
yy Nuclear envelope is re-organised from RER by a process
called nucleogenesis around each set of chromosomes at
either pole of the cell.
yy Two new identical nuclei forms.
5. Cytokinesis
yy Cytokinesis completes the process of cell division.
yy Cytokinesis is the process in which the cell actually divides
into two. With the two nuclei already at opposite poles of
the cell, the cell cytoplasm separates, and the cell pinches
in the middle, ultimately leading to cleavage.
yy The two cells formed are roughly equal in size.
yy The process of cytokinesis is different in animal and plant cells.

yy Cytokinesis in plants, which have cell walls, involves a

completely different mechanism.
During telophase, vesicles from the Golgi apparatus
move to the middle of the cell, where they fuse with each
other to form a cell plate or middle lamella.
Cell wall materials carried in the vesicles collect in the
cell plate as it grows.
The plate enlarges until its membranes fuse with the
plasma membrane at the perimeter.
The contents of the vesicles form new cell wall material
between the daughter cells.

Fig. Cell plate formation in a plant cell

Fig. Cytokinesis

yy In animal cells, cytokinesis occurs by a process called

cleavage.
The first sign of cleavage is the appearance of a cleavage
furrow in the cell surface near the old metaphase plate.
On the cytoplasmic side of the cleavage furrow is a
contractile ring of actin filaments. They induce the cell
membrane to invaginate.
The furrow deepens centripetally and cleaves the cell
into two.

Fig. Cytokinesis in animal cell

Significance of Mitosis
yy It is an equational division, and the two daughter cells are
identical in all respects.
yy They receive the same number and kind of chromosomes
as were in the mother cells.
It is the only mode of multiplication in unicellular
organisms.
It is the process by which growth takes place in animals
and plants by constantly adding more and more cells.
Somatic cells are formed by mitosis.
It maintains the genetic continuity and equality. This
helps in proper coordination among different cells.
It also plays a role in repair by growth, example in
wound healing, regeneration of damaged parts (as in
the tail of lizard), and replacement of cells lost during
normal wear and tear (as the surface cells of the skin or
the red blood cells).

Meiosis (also called as Reductional division)

yy Meiosis was discovered in plant cells by Strasburger.
yy The term Meiosis was coined by J.B., Farmer and J.E.
Moore.
yy Meiosis is a specialized kind of cell division that reduces
the chromosome number by half (thus called reductional
division) resulting in production of haploid daughter cells
(gametes). For example, in human beings, the normal
chromosome number is 46 (23 pairs), but as a result of
meiosis this number is halved to 23 in daughter cells.
yy Meiosis-I initiates after the parental chromosomes replicates
to produce identical sister chromatids at the S-Phase of
Interphase.

5
yy Meiosis occurs in reproductive cells, e.g. in the testes of male
and in the ovaries of female animals; in the pollen mother
cell of the anthers (male organs) and in the megaspore
mother cells of the ovary (female organ) of the flowers.
yy Cells which undergo meiosis are called meiocytes.
yy It involves sequential cycle of two nuclear division called
Meiosis-I and Meiosis-II but only a single cycle of DNA
replication takes place.
yy It results in four haploid daughter cells.
yy The cells produced by meiosis are not all alike. Each cell
is unique and this variation is produced by two processes
independent segregation of the chromosomes and by
crossing-over. Both take place during Meiosis I.
yy Importance of meiosis: Every living organism has a definite
number of chromosomes in its body cells. Therefore to keep
the chromosome number constant, the reproductive cells of the
parents (ovaries and testis in animals, and pollen mother cells
and cells of ovary in plants) divide through meiosis.
Stages of Meiosis
Meiosis consists of two divisions:
(i) Meiosis I (Reduction Division): Meiosis I start after DNA
has been replicated in S phase. During meiosis I, the number
of chromosome is reduced to half. The homologous pairs
of chromosomes get separated and cells become haploid.
The two chromatids of the chromosome become genetically
different due to crossing over.
Meiosis I is followed by meiosis II at the end of which four
haploid cells are formed.
(ii) Meiosis II (Equational division): During meiosis II the
sister chromatids of each chromosome are separated (as
in mitosis). It is called equational division because the
chromosome number remains the same as produced after
the end of the first division.
Meiosis I
Like mitosis, meiosis also consists of four stages; prophase,
metaphase, anaphase and telophase.

 rophase I: It is the longest phase and is further sub-divided into

P
5 phases.
(i) Leptotene (Gk. leptos - thin; tene thread):
yy The chromosomes become distinct and compact due to
condensation and thickening of chromosomes. They possess
a string of swollen areas called chromomeres.
yy Each chromosome consists of two chromatids held together
by a centromere but these are not easily visible.

(ii) Zygotene (Gk. Zygos-pairing)

yy Homologous chromosomes become closely associated due
to the development of nucleoprotein between them.
yy The process of attachment of the homologous chromosomes
to form a complex structure called synaptonemal complex
is known as synapsis.
yy Synapsis may be
Proterminal, starting from ends and prceeding toward
centromeres.
Procentric, starting from centromeres and proceeding
towards ends.
Random, at various places between centromeres.
yy Two synaptonemal complexes further form a complex called
bivalent or tetrad.
yy Bivalent or tetrad is a pair of synapsed homologous
chromosomes
(iii) Pachytene (Gk. pachus - thick)
yy It is the longest phase of prophase I.
yy The chromosome becomes shorter and thicker due to
contraction.
yy During pachytene, each chromosome shows two chromatids
so that bivalent shows 4 chromatids. Hence they are called
pachytene tetrads.
yy The two chromatids belonging to the same chromosomes
are called sister chromatids while chromatids belonging to
the two different chromosomes of a homologous pairs are
termed as non-sister chromatid.
yy Crossing-over occurs at the end of pachytene. It involves
break and exchange of parts (genes) between non-sister
chromatid i.e. chromatids of a homologous pair.
yy Crossing is mainly responsible for the genetic difference.
yy Crossing over is the exchange of genetic material between
two homologous chromosomes with the help of enzyme
recombinase.
yy The point of interchange and rejoining appears X-shaped
and is known as chiasma or points of crossing over.
yy Crossing over results in recombinations of linked genes,
which ultimately lead to evolution.
(iv) Diplotene (Gk. Diplous-double)
yy The nucleoprotein fusion complex of the synapsed
chromosomes dissolves partially. As a result, the homologous
chromosomes begin to separate.
yy The two non-sister chromatids of a homologous pair remain
attached at one or two points, called the chiasmata.
yy It is at the chiasmata that exchange of segments of
chromatids (genes) between homologous chromosomes
has taken place. The process of gene exchange is known
as genetic recombination.
yy Chiasmata may be terminal or interstitial. Depending on
their position, the homologous chromosomes appear crosslike, ring like or chain like.
(v) Diakinesis (GK dia = through, in different directions,
kinesis = motion)
yy Bivalents become very thick and short and migrate to the
periphery of the nucleus.

6
yy The homologous chromosomes of a bivalent move apart
from each other by a phenomenon called terminalisation.
yy Movement of intermediary chiasmata to ends of
chromosomes due to repulsion between homologous
chromosomes is called terminalisation.

yy Nuclear membrane and nucleolus disappear during

diakinesis.
yy Formation of spindle also gets completed by the end of
diakinesis.

Fig. Different stages in the meiotic division of a plant cell

Metaphase-I
yy The bivalents (homologous pairs of chromosomes) arrange
themselves as a double row at the equator or metaphase
plate.
yy The spindle fibres get attached at the centromere of the
chromosomes.
yy Centromeres bivalents are away from equatorial plane, and
arms are on equatorial plane.

yy The homologous chromosomes remain attached by the

chiasmata at the telomeric ends.
Anaphase-I
yy The spindle fibres shorten.
yy Homologous chromosomes of each bivalent segregate and
move to opposite poles during anaphase -I. (Note: There is
no division of centromere).Thus, half of the chromosome
(each with two chromatids) of the parent cell goes to one
pole and the remaining half to the opposite pole.

7
yy Segregation of homologous chromosomes during anaphase
I is called disjunction.
yy Random segregation of chromosomes during anaphase I is
a cause for recombinations of non-linked genes.
yy The number of chromosomes is reduced to half by the end
of anaphase - I.
yy Each set of chromosomes that moves to one pole consists
of a mixture of paternal and maternal chromosome parts
(new gene combination).
Telophase-I
yy The separated chromosomes form two haploid nuclei by
the end of telophase - I.
yy The daughter cells are now called haploid (n) as they have
one set of chromosomes.
yy The nucleolus reappears and nuclear membrane reforms.
yy The daughter nuclei begin the second meiotic division.
Interkinesis
yy It is the time gap between telophase of meiosis I and
prophase of meiosis II.
yy Chromosomes elongate but there is no formation of
chromatin reticulum.
yy Synthesis of RNA and protein may take place.
yy Centrosome or centriole pairs undergo replication.
 eiosis II
M
It has the same four stages; Prophase II, Metaphase II,
Anaphase II, Telophase II.
Though it is similar to mitosis, but meiosis II is not mitosis
because:
yy It always occurs in haploid cells.
yy It is not preceded by DNA replication.
yy The daughter cells formed at the end of meiosis II is neither
similar to each other nor to the parent cell.
Prophase II
yy The chromosomes shorten and reappear. The two chromatids
are attached to the single centromere.
yy The formation of spindle starts.
yy Nucleolus and nuclear membrane begin to disappear.
yy In animal cells, the centriole pairs develop asters and move
to the region of future spindle poles.
Metaphase II
yy The chromosomes arrange themselves along the equator.

yy The formation of spindle apparatus gets completed.

yy Each chromosome gets connected with both the spindle
poles by chromosome fibres that develop from both the
surfaces of its centromere.
Anaphase II
yy The centromere in each chromosome divides into two, so
that there is one centromere for each chromatid.
yy The chromatids get their centromere and become daughter
chromosomes and begin to move towards the opposite
poles.
yy At the end of anaphase II, four groups of chromosomes are
produced, each having haploid number.
Telophase II
yy On reaching the poles the chromosomes organize themselves
into haploid daughter nuclei.
yy The nucleolus and the nuclear membrane reappear.
yy Chromosomes elongate to form chromatin network.
yy Spindle fibres degenerate during telophase II.
Cytokinesis
yy It can be of two types: Successive and simultaneous type.
In successive type, cytokinesis occurs twice following
meiosis -I and meiosis - II.
In simultaneous type, cytokinesis occurs only after
meiosis II.
yy Meiosis results in four haploid cells.
Significance of Meiosis
yy Meiosis maintains the chromosome number from generation
to generation. It reduces the chromosome number to half so
that the process of fertilisation restores the original number
in the zygote.
yy Meiosis occurs during gamete formation (gemetogenesis)
and reduces the number of chromosomes from diploid (2n)
to haploid (n) in the gametes. These haploid gametes fuse to
form diploid zygote during fertilization. The diploid zygote
develops into a normal diploid individual.
yy Variations are caused by the cross-over and the random
distribution of homologous chromosomes between daughter
cells. Variations play an important role in evolution.
yy Chromosomal mutations are br ought about by the
introduction of certain abnormalities. These chromosomal
mutations may be advantageous for an individual.

Difference between Mitosis and Meiosis

Mitosis
In mitosis, a single division results in two daughter cells.

Meiosis
Meiosis involves two successive divisions meiosis I and meiosis II. These
divisions result in four daughter cells.

Mitosis is known as equational division. This is because the daughter Meiosis I is known as reductional division. This is because the chromosome
cells have the same diploid number of chromosomes as the parent.
number is reduced to half.
Meiosis II is known as equational division. This is because the sister
chromatids separate and the chromosome number remains the same.
Prophase is short and does not comprise any phase.

Prophase I is very long and comprises 5 phases leptotene, zygotene,

pachytene, diplotene, and diakinesis.

188
There is no pairing of chromosomes, crossing-over, or chiasmata- In the zygotene stage of prophase, the pairing of chromosomes occurs.
formation during prophase.
During pachytene, the crossing-over occurs. The chiasmata are formed in
the diplotene stage.
Synaptonemal complex is not formed.

Synaptonemal complex is formed during the zygotene stage of prophase I.

Anaphase involves the separation of the chromatids of each During anaphase I, the homologous chromosomes separate, while the
chromosome.
chromatids remain attached at their centromeres.
During anaphase II, the chromatids separate as a result of the splitting of
the centromere.
Mitosis plays a significant role in the healing, repair, and growth of a Meiosis brings about variation and maintains the chromosome number from
cell.
generation to generation.

Types of Meiosis
There are three types of meiosis, depending upon the stage when
meiosis occurs: Gametic meiosis, zygotic meiosis and sporic
meiosis.
1. Gametic meiosis
yy In gametic meiosis, the diploid zygote undergoes mitosis to
produce multicellular diploid cells. Cells from the diploid
individuals then undergo meiosis to produce haploid
gametes.
yy The haploid gametes fuse and produce the diploid zygote
with gametes of the opposite type.
yy In the whole cycle, gametes are the only haploid cells;
mitosis occurs only in the diploid phase.
2. Zygotic meiosis
yy A zygotic meiosis is meiosis of a diploid zygote immediately
after karyogamy, the fusion of two cell nuclei.

yy In this way, the organism ends its diploid phase and produces
several haploid cells. These cells undergo mitosis (divide
mitotically) to form either larger, multicellular individuals,
or more haploid cells. Two opposite types of gametes (e.g.,
male and female) from these individuals or cells fuse to
become a diploid zygote.
yy In the whole cycle, zygotes are the only diploid cell; mitosis
occurs only in the haploid phase.
3. Sporic meiosis
yy In sporic meiosis mitosis occur in both the diploid and haploid
phases. The organism exhibits alternation of generations,
which features spore-producing multicellular sporophytes
(which are diploid) and gamete-producing multicellular
gametophytes (which are haploid). Diagramatically, sporic
meiosis looks like the complex halves of gametic meiosis
and zygotic meiosis are merged into one.