CELL CYCLE AND CELL DIVISION Although cell growth (in terms of

cytoplasmic increase) is a continuous

Are you aware that all organisms, even
process, DNA synthesis occurs only during
the largest, start their life from a single
one specific stage in the cell cycle. The
cell?
replicated chromosomes (DNA) are then
You may wonder how a single cell then distributed to daughter nuclei by a
goes on to form such large organisms. complex series of events during cell
division.
Growth and reproduction are
characteristics of cells, indeed of all living These events are themselves under
organisms. genetic control.

All cells reproduce by dividing into two, 10.1.1 Phases of Cell Cycle
with each parental cell giving rise to two
A typical eukaryotic cell cycle is illustrated
daughter cells each time they divide.
by human cells in culture.
These newly formed daughter cells can
These cells divide once in approximately
themselves grow and divide, giving rise to
every 24 hours.
a new cell population that is formed by
the growth and division of a single However, this duration of cell cycle can
parental cell and its progeny. vary from organism to organism and also
from cell type to cell type.
In other words, such cycles of growth and
division allow a single cell to form a Yeast for example, can progress through
structure consisting of millions of cells. the cell cycle in only about 90 minutes.

10.1 CELL CYCLE The cell cycle is divided into two basic
phases: Interphase and M Phase (Mitosis
Cell division is a very important process in
phase)
all living organisms.
The M Phase represents the phase when
During the division of a cell, DNA
the actual cell division or mitosis occurs
replication and cell growth also take place.
and the interphase represents the phase
All these processes, i.e., cell division, DNA between two successive M phases.
replication, and cell growth, hence, have
It is significant to note that in the 24 hour
to take place in a coordinated way to
average duration of cell cycle of a from
ensure correct division and formation of
one cell human cell, cell division proper
progeny cells containing intact genomes.
lasts for only about an hour.
The sequence of events by which a cell
The interphase lasts more than 95% of the
duplicates its genome, synthesises the
duration of cell cycle.
other constituents of the cell and
eventually divides into two daughter cells The M Phase starts with the nuclear
is termed cell cycle. division, corresponding to the separation
of daughter chromosomes (karyokinesis)
and usually ends with division of How do plants and animals continue to
cytoplasm (cytokinesis). grow all their lives?

The interphase, though called the resting Do all cells in a plant divide all the time?
phase, is the time during which the cell is
Do you think all cells continue to divide in
preparing for division by undergoing both
all plants and animals?
cell growth and DNA replication in an
orderly manner. Can you tell the name and the location of
tissues having cells that divide all their life
The interphase is divided into three
in higher plants?
further phases: G1 phase (Gap 1), S phase
(Synthesis) and G2 phase (Gap 2). Do animals have similar meristematic
tissues?
G1 phase corresponds to the interval
between mitosis and initiation of DNA You have studied mitosis in onion root tip
replication. cells.

During G1 phase, the cell is metabolically It has 16 chromosomes in each cell.

active and continuously grows but does
Can you tell how many chromosomes will
not replicate its DNA.
the cell have at G 1 phase, after S phase,
S or synthesis phase marks the period and after M phase?
during which DNA synthesis or replication
Also, what will be the DNA content of the
takes place.
cells at G 1, after S and at G 2 , if the
During this time the amount of DNA per content after M phase is 2C?
cell doubles.
Some cells in the adult animals do not
If the initial amount of DNA is denoted as appear to exhibit division (e.g., heart cells)
2C then it increases to 4C. and many other cells divide only
occasionally, as needed to replace cells
However, there is no increase in the
that have been lost because of injury or
chromosome number.
cell death.
If the cell had diploid or 2n number of
These cells that do not divide further exit
chromosomes at G1, even after S phase
G1 phase to enter an inactive stage called
the number of chromosomes remains the
quiescent stage (G0) of the cell cycle.
same, i.e., 2n.
Cells in this stage remain metabolically
In animal cells, during the S phase, DNA
active but no longer proliferate unless
replication begins in the nucleus, and the
called on to do so depending on the
centriole duplicates in the cytoplasm.
requirement of the organism.
During the G2 phase, proteins are
In animals, mitotic cell division is only
synthesised in preparation for mitosis
seen in the diploid somatic cells.
while cell growth continues.
However, there are few exceptions to this The chromosomal material becomes
where haploid cells divide by mitosis, for untangled during the process of
example, male honey bees. chromatin condensation.

Against this, the plants can show mitotic The centrosome, which had undergone
divisions in both haploid and diploid cells. duplication during S phase of interphase,
now begins to move towards opposite
From your recollection of examples of
poles of the cell.
alternation of generations in plants
identify plant species and stages at which The completion of prophase can thus be
mitosis is seen in haploid cells. marked by the following characteristic
events:
10.2 M PHASE
Chromosomal material condenses to form
This is the most dramatic period of the
compact mitotic chromosomes.
cell cycle, involving a major reorganisation
of virtually all components of the cell. Chromosomes are seen to be composed
of two chromatids attached together at
Since the number of chromosomes in the
the centromere.
parent and progeny cells is the same, it is
also called as equational division. Centrosome which had undergone
duplication during interphase, begins to
Though for convenience mitosis has been
move towards opposite poles of the cell.
divided into four stages of nuclear division
(karyokinesis), it is very essential to Each centrosome radiates out
understand that cell division is a microtubules called asters.
progressive process and very clear-cut
The two asters together with spindle
lines cannot be drawn between various
fibres forms mitotic apparatus.
stages.
Cells at the end of prophase, when viewed
Karyokinesis involves following four
under the microscope, do not show golgi
stages: Prophase, Metaphase, Anaphase
complexes, endoplasmic reticulum,
and Telophase
nucleolus and the nuclear envelope.
10.2.1 Prophase
10.2.2 Metaphase
Prophase which is the first stage of
The complete disintegration of the
karyokinesis of mitosis follows the S and
nuclear envelope marks the start of the
G2 phases of interphase.
second phase of mitosis, hence the
In the S and G2 phases, the new DNA chromosomes are spread through the
molecules formed are not distinct but cytoplasm of the cell.
intertwined.
By this stage, condensation of
Prophase is marked by the initiation of chromosomes is completed and they can
condensation of chromosomal material. be observed clearly under the
microscope.
This then, is the stage at which daughter nuclei, begin their migration
morphology of chromosomes is most towards the two opposite poles.
easily studied.
As each chromosome moves away from
At this stage, metaphase chromosome is the equatorial plate, the centromere of
made up of two sister chromatids, which each chromosome remains directed
are held together by the centromere. towards the pole and hence at the leading
edge, with the arms of the chromosome
Small disc-shaped structures at the
trailing behind.
surface of the centromeres are called
kinetochores. Thus, anaphase stage is characterised by
the following key events:
These structures serve as the sites of
attachment of spindle fibres to the Centromeres split and chromatids
chromosomes that are moved into separate.
position at the centre of the cell.
Chromatids move to opposite poles.
Hence, the metaphase is characterised by
10.2.4 Telophase
all the chromosomes coming to lie at the
equator with one chromatid of each At the beginning of the final stage of
chromosome connected by its karyokinesis, i.e., telophase, the
kinetochore to spindle fibres from one chromosomes that have reached their
pole and its sister chromatid connected respective poles decondense and lose
by its kinetochore to spindle fibres from their individuality.
the opposite pole.
The individual chromosomes can no
The plane of alignment of the longer be seen and each set of chromatin
chromosomes at metaphase is referred to material tends to collect at each of the
as the metaphase plate. two poles.

The key features of metaphase are: This is the stage which shows the
following key events:
Spindle fibres attach to kinetochores of
chromosomes. Chromosomes cluster at opposite spindle
poles and their identity is lost as discrete
Chromosomes are moved to spindle
elements.
equator and get aligned along metaphase
plate through spindle fibres to both poles. Nuclear envelope develops around the
chromosome clusters at each pole
10.2.3 Anaphase
forming two daughter nuclei.
At the onset of anaphase, each
Nucleolus, golgi complex and ER reform.
chromosome arranged at the metaphase
plate is split simultaneously and the two 10.2.5 Cytokinesis
daughter chromatids, now referred to as
Mitosis accomplishes not only the
daughter chromosomes of the future
segregation of duplicated chromosomes
into daughter nuclei (karyokinesis), but However, in some lower plants and in
the cell itself is divided into two daughter some social insects haploid cells also
cells by the separation of cytoplasm called divide by mitosis.
cytokinesis at the end of which cell
It is very essential to understand the
division gets completed.
significance of this division in the life of an
In an animal cell, this is achieved by the organism.
appearance of a furrow in the plasma
Are you aware of some examples where
membrane.
you have studied about haploid and
The furrow gradually deepens and diploid insects?
ultimately joins in the centre dividing the
Mitosis usually results in the production
cell cytoplasm into two.
of diploid daughter cells with identical
Plant cells however, are enclosed by a genetic complement.
relatively inextensible cell wall, thererfore
The growth of multicellular organisms is
they undergo cytokinesis by a different
due to mitosis.
mechanism.
Cell growth results in disturbing the ratio
In plant cells, wall formation starts in the
between the nucleus and the cytoplasm.
centre of the cell and grows outward to
meet the existing lateral walls. It therefore becomes essential for the cell
to divide to restore the nucleo-
The formation of the new cell wall begins
cytoplasmic ratio.
with the formation of a simple precursor,
called the cell-plate that represents the A very significant contribution of mitosis is
middle lamella between the walls of two cell repair.
adjacent cells.
The cells of the upper layer of the
At the time of cytoplasmic division, epidermis, cells of the lining of the gut,
organelles like mitochondria and plastids and blood cells are being constantly
get distributed between the two daughter replaced.
cells.
Mitotic divisions in the meristematic
In some organisms karyokinesis is not tissues – the apical and the lateral
followed by cytokinesis as a result of cambium, result in a continuous growth of
which multinucleate condition arises plants throughout their life.
leading to the formation of syncytium
10.4 MEIOSIS
(e.g., liquid endosperm in coconut).
The production of offspring by sexual
10.3 125 Significance of Mitosis
reproduction includes the fusion of two
Mitosis or the equational division is gametes, each with a complete haploid
usually restricted to the diploid cells only. set of chromosomes.

Gametes are formed from specialised

diploid cells.
This specialised kind of cell division that Prophase of the first meiotic division is
reduces the chromosome number by half typically longer and more complex when
results in the production of haploid compared to prophase of mitosis.
daughter cells.
It has been further subdivided into the
This kind of division is called meiosis. following five phases based on
Meiosis ensures the production of haploid chromosomal behaviour, i.e., Leptotene,
phase in the life cycle of sexually Zygotene, Pachytene, Diplotene and
reproducing organisms whereas Diakinesis.
fertilisation restores the diploid phase.
During leptotene stage the chromosomes
We come across meiosis during become gradually visible under the light
gametogenesis in plants and animals. microscope.

This leads to the formation of haploid The compaction of chromosomes

gametes. continues throughout leptotene.

The key features of meiosis are as follows: This is followed by the second stage of
prophase I called zygotene.
Meiosis involves two sequential cycles of
nuclear and cell division called meiosis I During this stage, chromosomes start
and meiosis II but only a single cycle of pairing together and this process of
DNA replication. association is called synapsis.

Meiosis I is initiated after the parental Such paired chromosomes are called
chromosomes have replicated to produce homologous chromosomes.
identical sister chromatids at the S phase.
Electron micrographs of this stage,
Meiosis involves pairing of homologous indicate that chromosome synapsis is
chromosomes and recombination accompanied by the formation of complex
between non-sister chromatids of structure called synaptonemal complex.
homologous chromosomes.
The complex formed by a pair of
Four haploid cells are formed at the end synapsed homologous chromosomes is
of meiosis II. called a bivalent or a tetrad.

Meiotic events can be grouped under the However, these are more clearly visible at
following phases: the next stage.

Meiosis I Meiosis II Prophase I Prophase II The first two stages of prophase I are
Metaphase I Metaphase II Anaphase I relatively short-lived compared to the next
Anaphase II Telophase I Telophase II stage that is pachytene pachytene.

10.4.1 Meiosis I During this stage, the four chromatids of

each bivalent chromosomes becomes
Prophase I:
distinct and clearly appear as tetrads.
This stage is characterised by the By the end of diakinesis, the nucleolus
appearance of recombination nodules, disappears and the nuclear envelope also
the sites at which crossing over occurs breaks down.
between non-sister chromatids of the
Diakinesis represents transition to
homologous chromosomes.
metaphase.
Crossing over is the exchange of genetic
Metaphase I:
material between two homologous
chromosomes. The bivalent chromosomes align on the
equatorial plate.
Crossing over is also an enzyme-mediated
process and the enzyme involved is called The microtubules from the opposite poles
recombinase. of the spindle attach to the kinetochore of
homologous chromosomes.
Crossing over leads to recombination of
genetic material on the two Anaphase I:
chromosomes.
The homologous chromosomes separate,
Recombination between homologous while sister chromatids remain associated
chromosomes is completed by the end of at their centromeres.
pachytene, leaving the chromosomes
Telophase I:
linked at the sites of crossing over.
The nuclear membrane and nucleolus
The beginning of diplotene is recognised
reappear, cytokinesis follows and this is
by the dissolution of the synaptonemal
called as dyad of cells.
complex and the tendency of the
recombined homologous chromosomes Although in many cases the chromosomes
of the bivalents to separate from each do undergo some dispersion, they do not
other except at the sites of crossovers. reach the extremely extended state of the
interphase nucleus.
These X-shaped structures, are called
chiasmata. The stage between the two meiotic
divisions is called interkinesis and is
In oocytes of some vertebrates, diplotene
generally short lived.
can last for months or years.
There is no replication of DNA during
The final stage of meiotic prophase I is
interkinesis.
diakinesis.
Interkinesis is followed by prophase II, a
This is marked by terminalisation of
much simpler prophase than prophase I.
chiasmata.
10.4.2 Meiosis II
During this phase the chromosomes are
fully condensed and the meiotic spindle is Prophase II:
assembled to prepare the homologous
chromosomes for separation.
Meiosis II is initiated immediately after It also increases the genetic variability in
cytokinesis, usually before the the population of organisms from one
chromosomes have fully elongated. generation to the next.

In contrast to meiosis I, meiosis II Variations are very important for the

resembles a normal mitosis. process of evolution.

The nuclear membrane disappears by the SUMMARY

end of prophase II.
According to the cell theory, cells arise
The chromosomes again become from preexisting cells.
compact.
The process by which this occurs is called
Metaphase II: cell division.

At this stage, the chromosomes align at Any sexually reproducing organism starts
the equator and the microtubules from its life cycle from a single-celled zygote.
opposite poles of the spindle get attached
Cell division does not stop with the
to the kinetochores of sister chromatids.
formation of the mature organism but
Anaphase II: continues throughout its life cycle.

It begins with the simultaneous splitting of The stages through which a cell passes
the centromere of each chromosome from one division to the next is called the
(which was holding the sister chromatids cell cycle.
together), allowing them to move toward
Cell cycle is divided into two phases:
opposite poles of the cell by shortening of
microtubules attached to kinetochores. (i) Interphase – a period of preparation for
cell division.
Telophase II:
(ii) Mitosis (M phase) – the actual period of
Meiosis ends with telophase II, in which
cell division.
the two groups of chromosomes once
again get enclosed by a nuclear envelope. Interphase is further subdivided into G1, S
and G2.
Cytokinesis follows resulting in the
formation of tetrad of cells i.e., four G1 phase is the period when the cell
haploid daughter cells. grows and carries out normal metabolism.

10.5 SIGNIFICANCE OF MEIOSIS Most of the organelle duplication also

occurs during this phase.
Meiosis is the mechanism by which
conservation of specific chromosome S phase marks the phase of DNA
number of each species is achieved across replication and chromosome duplication.
generations in sexually reproducing
G2 phase is the period of cytoplasmic
organisms, even though the process, per
growth.
se, paradoxically, results in reduction of
chromosome number by half.
Mitosis is also divided into four stages Meiosis is divided into two phases –
namely prophase, metaphase, anaphase meiosis I and meiosis II.
and telophase.
In the first meiotic division the
Chromosome condensation occurs during homologous chromosomes pair to form
prophase. bivalents, and undergo crossing over.

Simultaneously, the centrioles move to Meiosis I has a long prophase, which is

the opposite poles. divided further into five phases.

The nuclear envelope and the nucleolus These are leptotene, zygotene, pachytene,
disappear and the spindle fibres start diplotene and diakinesis.
appearing.
During metaphase I the bivalents arrange
Metaphase is marked by the alignment of on the equatorial plate.
chromosomes at the equatorial plate.
This is followed by anaphase I in which
During anaphase the centromeres divide homologous chromosomes move to the
and the chromatids start moving towards opposite poles with both their
the two opposite poles. chromatids.

Once the chromatids reach the two poles, Each pole receives half the chromosome
the chromosomal elongation starts, number of the parent cell.
nucleolus and the nuclear membrane
In telophase I, the nuclear membrane and
reappear.
nucleolus reappear.
This stage is called the telophase. Nuclear
Meiosis II is similar to mitosis.
division is then followed by the
cytoplasmic division and is called During anaphase II the sister chromatids
cytokinesis. separate.

Mitosis thus, is the equational division in Thus at the end of meiosis four haploid
which the chromosome number of the cells are formed.
parent is conserved in the daughter cell.

In contrast to mitosis, meiosis occurs in

the diploid cells, which are destined to
form gametes.

It is called the reduction division since it

reduces the chromosome number by half
while making the gametes.

In sexual reproduction when the two

gametes fuse the chromosome number is
restored to the value in the parent.