CHAPTER – CELL CYCLE AND CELL DIVISION

Cell division is defined as a process in which a single parent cell divides into two daughter
cells.
9.1 CELL CYCLE
• The sequence of events by which a cell duplicates its genome, synthesises other
constituents of the cell and eventually divides into two daughter cells is termed as cell
cycle.
• All the events of the cell cycle occur in a coordinated manner and are under genetic
control.
• Although cell growth is a continuous process, DNA synthesis occurs only during one
specific stage in the cell cycle.
• The replicated chromosomes (DNA) are distributed to daughter nuclei by a complex
series of events during cell division.
• During the division of a cell, DNA replication and cell growth take place.
• All these processes, i.e. cell division, DNA replication and cell growth, have to take
place in a coordinated way to ensure correct division and formation of progeny cells
containing intact genomes.

9.1.1 PHASES OF CELL CYCLE

G1 phase

Interphase S phase
Prophase
G2 phase
Cell Cycle
Metaphase
Karyokinesis
M or Mitotic
Anaphase
phase
Cytokinesis
Telophase
Fig: A diagrammatic view of the cell cycle indicating the formation of two cells from
one cell

INTERPHASE

• It is a long, non-dividing, growing phase of the cell cycle.

• It represents the most active stage of the cell cycle, where both cell growth and DNA
replication occur in an orderly manner.
• It is the period of intense growth and synthesis in which a cell stocks all the
biomolecules required during cell division.
• It is called the resting phase because there is no apparent activity related to cell division.
Rather, it deals with the changes that occur in a cell and the nucleus before it enters into
cell division.
• In the 24-hour average duration of the cell cycle of a human cell, cell division properly
lasts for only about an hour. The interphase lasts more than 95% of the duration of the
cell cycle.

Interphase is further divided into three stages:

Stage of Interphase Characteristic features

G1 phase • It is the longest phase of the interphase between mitosis and
initiation of DNA replication.
• Continuous growth of cells.
• Cells become metabolically active.
 • Prepares machinery for DNA replication.
• Active synthesis of RNA and proteins.
• Most of the organelles duplication occurs in this phase.
S phase • DNA replication takes place.
• Amount of DNA per cell doubles.
• Chromosome number does not increase.
• Synthesis of histone proteins also occurs
• In animal cells, during the S phase, DNA replication begins in
the nucleus, and the centrioles duplicate in the cytoplasm
G2 phase • It is the second gap phase between S and M phases.
• Cell growth continues.
• Synthesis of RNA and proteins continues.
• Mitochondria, Chloroplast and Golgi bodies duplicate in this
phase.
• Cell is prepared for mitosis.
G0 phase:

• Some cells in adult animals do not appear to exhibit division (e.g., heart cells), and
many other cells divide only occasionally as needed to replace cells that have been lost
because of injury or cell death. These cells that do not divide further exit the G1 phase
to enter an inactive stage called the quiescent stage (G0) of the cell cycle.
• Cells in this stage remain metabolically active but no longer proliferate unless called on
to do so depending on the requirement of the organism.

9.2 M PHASE

M phase or mitotic phase is the phase where actual cell division occurs. It is the most
dramatic period of the cell cycle. It consists of two processes.

I. Karyokinesis – where the division of the nucleus occurs (Karyon → Nucleus, Kinesis
→ Movement)

II. Cytokinesis – where the division of cytoplasm occurs (Kytos → Cell/ hollow, Kinesis
→ Movement)
9.3 MITOSIS (Equational Division)

• Mitosis was first observed by Strasburger in plant cells and by Walter Flemming in
animal cells.
• The term mitosis was given by Walter Flemming in 1882.
• Mitosis is a process in which a parent cell divides into two identical daughter cells.
These cells are identical to each other as well as to the parent cell.
• It occurs in the somatic cells (body cells) of the animals and in the gonads for the
multiplication of undifferentiated germ cells. In plants, it occurs in the dividing
meristematic tissue and also in leaves, flowers and fruits during growth.

Karyokinesis of Mitosis has 4 stages:

a) Prophase
b) Metaphase
c) Anaphase
d) Telophase

PROPHASE

• It is the first stage of karyokinesis of mitosis.

• The condensation of chromatin material takes place and during condensation, the DNA
strands get untangled to form compact mitotic chromosomes.
• Each chromosome appears double and consists of two coiled sister chromatids joined
by a centromere. Their ends are not visible in the early prophase.

• Centrosome which had undergone duplication during interphase, begins to move

towards the opposite poles of the cell.
• Each centrosome radiates out microtubules called asters. The two asters, together with
spindle fibres, form a mitotic apparatus.
• Cells at the end of prophase, when viewed under the microscope, do not show the
Golgi complex, ER, nuclear membrane, nucleolus and nuclear envelope.

METAPHASE

• Metaphase is the stage at which the morphology of chromosomes is most easily studied.
• The disintegration of the nuclear envelope marks the beginning of the metaphase. The
nuclear envelope disappears, and chromosomes spread through the cytoplasm of the
cell.
• The chromosomes contain two chromatids attached to each other through the
centromere. The chromosomes during this stage are the thickest and shortest.

• Mitotic spindle formation is complete. The phenomenon of bringing the chromosomes

to the equator of the spindle is called congression.
• All the chromosomes align themselves at the equator. The plane of alignment of
centromeres of all the chromosomes is referred to as the metaphase plate or equatorial
plate.
• The centromere is surrounded by a small disc-shaped structure called kinetochore. The
kinetochores form the site of attachment of microtubules (Spindle fibres).

Fig: A metaphase chromosome with kinetochores joined by spindle microtubules

ANAPHASE

• The centromere which holds the two chromatids together splits and separates daughter
chromatids are now referred to as chromosomes of the future daughter nuclei.
• The spindle fibres attached to the kinetochore now shorten and daughter chromosomes
begin to migrate towards the opposite poles.
• During migration, chromosomes always move away from the equatorial plate. The
centromere of the daughter chromosomes moves towards the pole, and the arms of
chromosomes trail it. The anaphase ends when the chromosomes reach the poles. It is
the best stage to study shape of the chromosomes (V, L, J, I).

TELOPHASE

• The chromosomes (sister chromatids) reach their respective poles. The mitotic spindle
disappears.
• After reaching the poles, the chromosomes decondense and lose their individuality. The
individual chromosomes can no longer be seen, and each set of chromatin material tends
to collect at each of the two poles.
• Nucleolus, endoplasmic reticulum, Golgi bodies, and other organelles reappear in the
daughter cells.
• The nuclear envelope develops around the chromosome clusters at each pole, forming
two daughter nuclei.

CYTOKINESIS

• This phase marks the end of cell division. After nuclear division, i.e., karyokinesis, the
cytoplasm of the parent cell divides into two daughter cells so that two daughter cells
have their own nucleus and cytoplasm. The cell organelles present in the cytoplasm also
distribute in two daughter cells.
• In animal cells, cytokinesis is achieved by the formation of a furrow. The furrow in the
plasma membrane of the cell deepens gradually. It moves centripetally and ultimately
joins in the centre, dividing the cell cytoplasm into two.
• In plant cells, cytokinesis is achieved by cell plate formation in a centrifugal manner.
Phragmoplast is formed by the Golgi complex to form a cell plate. The formation of a
new cell wall begins with the formation of a simple precursor, called a cell plate, that
represents the middle lamella.
SIGNIFICANCE OF MITOSIS

• Growth: Mitosis is essential for the growth and development of multicellular

organisms.
• Maintenance of cell size: Mitosis maintains the size of the cell. An overgrown somatic
cell is induced to divide so that it maintains a proper surface area to volume ratio, which
is essential for the proper functioning of the cell. If a cell becomes large, then it enters
into mitosis.
• Maintenance of chromosome number (genetic stability): Mitosis maintains the same
type and number of chromosomes in two daughter cells.
• Repair: Mitosis is a mechanism for replacing old dead and worn-out cells with new
cells.
• Reproduction: Mitosis brings about reproduction (multiplication) in unicellular
organisms.
• Healing and regeneration: Mitosis produces new cells for healing the wounds and for
regeneration.
9.4 MEIOSIS (REDUCTIONAL DIVISION)

• The term meiosis was coined by Farmer and Moore in 1905. The division was studied
by Van Benedin Winiwarter and Strassburger.
• The division that reduces the chromosome number by half results in the production of
haploid daughter cells. This kind of division is called meiosis.
• Meiosis ensures the production of the haploid phase in the life cycle of sexually
reproducing organisms, whereas fertilisation restores the diploid phase.
• Meiosis involves two sequential cycles of nuclear and cell division called meiosis I and
meiosis II, but only a single cycle of DNA replication.
• Meiosis I is initiated after the parental chromosomes have replicated to produce
identical sister chromatids at the S phase.
• Meiosis involves the pairing of homologous chromosomes and recombination between
them.
• Four haploid cells are formed at the end of meiosis II.

Meiosis consists of two divisions i.e., Meiosis I and Meiosis II

Meiosis I Meiosis II

1. Prophase I 2. Prophase II

3. Metaphase I 4. Metaphase II

5. Anaphase I 6. Anaphase II

7. Telophase I 8. Telophase II

9.4.1 MEIOSIS I

It is the reductional division in which the number of chromosomes is reduced to half. It is

studied in four stages:

PROPHASE I

Prophase I is more elaborate, prolonged, and complex than the prophase of mitosis.
Prophase I is further subdivided into five stages, viz., Leptotene, Zygotene, Pachytene,
Diplotene and Diakinesis.
 Stage of Prophase I Characteristic features
Leptotene • Also known as the bouquet stage.
• Condensation and coiling of chromatin fibres begin
to form distinct chromosomes.
• Chromosomes become gradually visible under the
microscope.
Zygotene • Chromosomes become more condensed.
• The two chromosomes which are similar in form,
size, and structure are called homologous
chromosomes.
• During zygotene, these homologous chromosomes
start pairing together. These homologous
chromosomes come to lie side by side in pairs, and
this pairing is known as synapsis.
• The complex formed by a pair of synapsed
chromosomes is called bivalent.
• Chromosomal synapsis is accompanied by a
structure called synaptonemal complex, which is
thought to stabilise the two homologous
chromosomes till the crossing over is completed.
Pachytene • Bivalent chromosomes split into similar chromatids.
This stage is called tetrad.
• Genetic material is exchanged between non-sister
chromatids of the homologous chromosomes. The
process is called crossing over.
• The site where crossing over occurs forms a
recombination nodule. The recombination is an
enzyme-mediated process. An enzyme called
recombinase is involved during this process.
 • By the end of pachytene, the recombination between
the homologous chromosomes is complete and the
two chromatids are linked at the site of crossing over.
Diplotene • Longest phase of Prophase I
• During this stage, the crossing-over is completed,
and the two homologous chromosomes begin to
separate from each other.
• There is the dissolution of the synaptonemal
complex, so the homologous chromosomes will start
separating
• They remain attached at one or more points where
crossing over has occurred. These points of
attachment are called Chiasmata, which are seen as
X-shaped structures.
• Terminalisation of chiasma starts in this stage.
Diakinesis • It represents the transition to metaphase I.
• Spindle assembles to prepare homologous
chromosomes for separation.
• Terminalisation of chiasmata: Due to tight
condensation of chromosomes, the chiasmata
disappear from the chromosomes by slipping off or
sliding from the tip of the chromosomes.
• Nucleolus disappears, and the nuclear envelope
disintegrates

METAPHASE I

The bivalent chromosome aligns on the equatorial plate. The microtubules from the
opposite poles of the spindle attach to the kinetochore of homologous chromosomes.

ANAPHASE I
Due to the shortening of kinetochore/chromosomal fibres, homologous chromosomes
separate from each other and move towards the opposite poles. Sister chromatids remain
associated at their centromeres.

TELOPHASE I

• The chromosomes reach the poles. The spindle fibres completely disappear.
• The nucleolus and nuclear membrane reappear.
• The chromosomes uncoil and elongate but remain straight in this phase. They do not
reach the extremely extended state of the interphase nucleus.
• It produces two daughter nuclei, each containing half the number of chromosome but
double the amount of nuclear DNA as both the chromatids move together to a single
pole. The separation of these two chromatids occurs during meiosis II.

9.4.2 MEIOSIS II

• Meiosis II is similar to mitosis, i.e., equational division, but not an exact copy of
mitosis because mitosis occurs in diploid somatic cells, but meiosis II always occurs in
haploid germ cells.
• Mitosis is always followed by DNA replication, but meiosis II is not followed by DNA
replication.
• The daughter cells formed after meiosis II are neither similar to each other nor
similar to the parent cell.
• The main event that occurs during meiosis II is the separation of the chromatid of the
univalent chromosomes present in the daughter cells formed after meiosis I.
• The chromatids present in the univalent chromosome differ from each other due to
crossing over.

Meiosis II is divided into four phases, namely – Prophase II, Metaphase II, Anaphase II
and Telophase II.

• Prophase II: Prophase II is not as long and complicated as Prophase I. It is a short

phase where the chromatids of the univalent condense. The chromatin material again
becomes compact. The nucleolus and the nuclear envelope disintegrate and disappear.
• Metaphase II: The univalents, i.e., chromosomes, align themselves at the equator (on
the equatorial plane) in metaphase II. The microtubules from the opposite poles extend
towards the equator and attach at the kinetochore of the chromatids.
• Anaphase II: During this phase, the centromere holding the two chromatids splits and
allows the separation and movement of the two chromatids to the opposite poles of the
cell by shortening of microtubules attached to kinetochores.
• Telophase II: This is the last stage of meiosis II. During this phase, the chromatids
reach the poles and start uncoiling. They decondense and become thin. The spindle
fibres degenerate. The nuclear membrane and nucleolus reappear and four haploid
nuclei are formed. The telophase II is now followed by cytokinesis which divides the
cytoplasm and forms four individual haploid cells.

SIGNIFICANCE OF MEIOSIS

• Formation of gametes: Meiosis produces gametes for sexual reproduction.

• Maintenance of chromosome number: Meiosis reduces the chromosome number to
half in the gametes so that fertilization restores the original diploid number in the
zygote.
• Introduction of variations: Meiosis provides a chance for the formation of new
combinations of chromosomes. This brings out variations.

DIFFERENCE BETWEEN MITOSIS AND MEIOSIS

S.No MITOSIS MEIOSIS

GENERAL
1. Division occurs in somatic cells It occurs in reproductive cells
2. Both haploid and diploid cells show Found only in haploid cells.
mitosis
3. It is a single division It is a double division
4. It produces two cells It produces four cells
5. It does not introduce variation It introduces variations due to gene
exchange
6. Number of chromosomes same as Chromosome number is reduced/halved
mother cells.
7. It is required for growth, repair and It is involved only in sexual reproduction.
healing.
PROPHASE
8. It is simpler It is complicated
9. The bouquet stage and synapsis are The bouquet stage and synapsis occur
absent.
10. Crossing over does not occur, and Chiasmata and crossing-over formation
chiasmata are not seen occur.
METAPHASE
11. Centromeres are present over the Centromeres projects towards the poles.
equator or metaphase plate while
arms facing towards the poles.
ANAPHASE
12. Centromere divides during Division of centromere is absent in
anaphase Anaphase I
13. Chromosomes have a single Chromosome has two chromatids in
chromatid. Anaphase I
14. The two chromatids of a Chromatids do not separate in anaphase-I
chromosome separate and become
daughter chromosomes.
15. Chromosomes moving towards the Chromosomes are dissimilar.
opposite poles are similar
TELOPHASE
16. Chromosomes become completely Chromosomes not completely
decondensed to form chromatin decondensed to form chromatin network
network
17. Daughter cells possess same genetic All four daughter cells are differ from each
constitution as in mother cell. other and also from mother cell.