Cell Cycle And Division

[Notes]
Created @January 21, 2023 12:05 PM

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INTRODUCTION
Growth and reproduction are two major characteristics of
living organisms which are also shown by their individual
cells. A young cell grows in size. There is replication of
genetic material followed by division of the cell into two.
The number of cells increases with each cycle of growth and
division. A large sized cell population is produced and
differentiation leads to the formation of tissues, organs
and organ systems.

Prevost and Dumas (1824) are the first to study cell

division during the cleavage of zygote of frog.

Rudolf Virchow (1855) observed that new cells always

develop from pre-existing cells. He also gave cell lineage
theory or law of cell lineage and doctrine of genetic
continuity.

Cell division is the process by which a mature cell divides

and forms two nearly equal daughter cells which resemble
the parental cell in a number of characters.

The cell which undergoes division is called mother cell or

parent cell. The newly formed cells are known as daughter
cells.

In unicellular organisms, cell division is the means of

reproduction by which the mother cell produces two or more

Cell Cycle And Division [Notes] 1

 new cells. In multicellular organisms also, new individual
develops from a single cell.

CELL CYCLE
Cell division is a biological process in all living
organisms.

Although cell growth (in terms of cytoplasmic increase) is

a continuous process, DNA synthesis occurs only during one
specific stage in the cell cycle. The replicated
chromosomes (DNA) are then distributed to daughter nuclei
by a complex series of events during cell division. These
events are themselves under genetic control.

The sequence of events which occur during cell growth and

cell division are collectively called cell cycle.

PHASES OF CELL CYCLE

The period required to complete one cell cycle (from beginning
of one cell division to the beginning of next) is called

Cell Cycle And Division [Notes] 2

 generation time. It is 24 hours in human cells and 90 minutes
in yeast. Cell cycle is simpler in prokaryotes and more
complex in eukaryotes.

The cell cycle is divided into two basic phases :

Interphase

M Phase (Mitosis phase)/Dividing phase

INTERPHASE
It is the period between the end of one cell division to
the beginning of the next cell division.

It is a highly metabolically active phase in which cell

prepares itself for the next cell division.

All mature cells of the body, therefore, occur in

interphase. Nerve cells of mammals have the longest
interphase and thus, do not divide after their formation at
birth.

Interphase is completed into three successive stages.

(1) G1 phase/Post mitotic/Pre-DNA synthetic phase/gap-I

G1 phase corresponds to the interval between mitosis and

initiation of DNA replication.

Following events take place during this phase

Intensive cellular synthesis.

Synthesis of rRNA, mRNA ribosomes and proteins.

Metabolic rate is high.

Cell size increases.

Synthesis of enzymes, amino acids, nucleotides etc. but

there is no change in DNA amount.

(2) S-phase/Synthetic phase
S or synthesis phase marks the period during which DNA
synthesis or replication takes place.
Following events take place during this phase

Cell Cycle And Division [Notes] 3

 DNA replicates and its amount becomes double. If the
initial amount of DNA is denoted as 2C then it increases to
4C.

Synthesis of histone proteins and NHC (non-histone

chromosomal proteins).

Duplication of centrioles in the cytoplasm.

(3) G2-phase/Pre mitotic/Post synthetic phase/Gap-II

Following events take place during this phase

Mitotic spindle protein (tubulin) synthesis begins.

Chromosome condensation factor appears.

Synthesis of 3 types of RNA, NHC proteins, and ATP

molecule.

Repair of damaged DNA occurs.

The cells that do not divide further exit G1 phase to enter

an inactive stage called 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.

M -PHASE/DIVIDING PHASE/MITOTIC PHASE

It is the phase of actual cell division.

It is divided into two phases - karyokinesis (division of

nucleus) and cytokinesis (division of the cytoplasm).

Cell Cycle And Division [Notes] 4

 Time period for G1, S, G2 and M-phase is species specific
under specific environmental conditions e.g., 20 minutes
for bacterial cell, 8-10 hours for intestinal epithelial
cells, onion root tip cells may take 20 hours. 24 hours for
human cells and 90 minutes for yeast cells to complete
these phases of cell cycle.

It is of three types – amitosis, mitosis and meiosis.

AMITOSIS
Amitosis is also called as direct cell division.

In this division, there is no differentiation of

chromosomes and spindle. The nuclear envelope does not
degenerate. The nucleus elongates and constricts in the
middle to form two daughter nuclei. This is followed by a
centripetal constriction of the cytoplasm to form two
daughter cells.

Examples : Prokaryotes, protozoans, yeasts, foetal membrane

of mammals, cartilage of mammals etc.

MITOSIS
Mitosis is also called indirect cell division or somatic
cell division or equational division.

In this, mature somatic cells divide in such a way that

chromosome number is kept constant that is, number of
chromosomes in daughter cells equal to those in the parent
cell.

The growing regions of plants have meristematic cells (e.g.

these cells are found in apical portion of root and stem
and in the expanding leaf) in which mitosis takes place.

KARYOKINESIS
It comprises of four phases i.e., Prophase, Metaphase,
Anaphase, Telophase.
(1) Prophase

It is the longest phase of karyokinesis.

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 Chromatin fibres thicken and shorten to form chromosomes
which may overlap each other and appear like a ball of
wool.

Each chromosome divides longitudinally into 2 chromatids

which remain attached to centromere.

Nuclear membrane starts disintegrating except in

dinoflagellates.

Nucleolus starts disintegrating.

Spindle formation begins.

In cytoskeleton, golgi complex, ER, etc. disappear.

In animal cells, centrioles move towards opposite sides.

(2) Metaphase

Cell Cycle And Division [Notes] 6

 Chromosomes become maximally distinct i.e., size can be
measured.

Chromosomes move towards equatorial plane of spindles

called congression and become arranged with their arms
directed towards pole and centromere towards equator.

Spindle fibres attach to kinetochores.

Metaphase is the best stage for studying chromosome

morphology (structure, size, number).

(3) Anaphase

Cell Cycle And Division [Notes] 7

 Centromere splits from the middle and two chromatids get
separated.

Both the chromatids move towards opposite poles due to

repulsive force called anaphasic movement.

Anaphasic movement is brought about by the repolymerisation

of continuous fibres and depoly-merisation of chromosomal
fibres. Formation and expansion of interzonal fibres occur.

The centromere faces towards equator.

Shape of chromosome is best studied at anaphase.

(4) Telophase

Chromosomes reach the poles by the spindle fibers and form

two groups.

Chromosomes begin to uncoil and form a chromatin net.

The nuclear membrane and nucleolus reappear.

In animal cells, astral rays and spindle fibres completely

disappear in telophase. The two centriole pairs organise
themselves into centrosomes. In plant cells, the spindle

Cell Cycle And Division [Notes] 8

 fibres disappear from near the poles but remain intact
towards the equator.

Golgi complex and ER etc., reform.

CYTOKINESIS
Cytokinesis is the division of cell having undergone
karyokinesis to produce two daughter cells each with a
daughter nucleus. It begins in mid-anaphase and is
generally completed along with the completion of telophase.

Cytokinesis is by 2 methods:

Cell furrow method : This is characteristic of animal

cells.

A dense vesicular and fibrous structure is formed in the

equatorial region of the spindle simultaneously with its
depolymerisation. It is called mid body.

A circular constriction or invagination appears at the centre

or equator and forms a furrow. The constriction or furrow
deepens centripetally and divides the mother cell into two
daughter cells.

Cell plate method : This is characteristic of plant cells.

Here, vesicles provided by Golgi apparatus unite to form
phragmoplasts, which join to form the cell plate. Cell
plate is first laid down in the centre and then proceeds
towards periphery (i.e., centrifugal plate-formation). Cell
wall materials are now laid down on both sides of cell
plate thus, forming two daughter cells.

TYPES OF MITOSIS

Intranuclear or Promitosis : Nuclear membrane is not

disintegrated and spindle is formed inside the nuclear
membrane e.g., Protozoans (Amoeba) and yeast. It is so as
centriole is present within the nucleus.

Extranuclear or Eumitosis : In this, nuclear membrane is

disintegrated and spindle is formed outside nuclear
membrane e.g., in plants and animals.

Cell Cycle And Division [Notes] 9

 Endomitosis : Chromosomes and their DNA duplicate but fail
to separate which leads to polyploidy e.g., in liver of
man, both diploid (2N) and polyploid cells (4N) have been
reported. It is also called endoduplication and
endopolyploidy.

Dinomitosis : In this, nuclear envelope persists but

microtubular spindle is not formed. During movement, the
chromosomes are attached with nuclear membrane, e.g.,
dinoflagellates.

SIGNIFICANCE OF MITOSIS

It keeps the chromosome number constant and maintains

genetic stability in daughter cells. All the cells have
similar genetic constituents.

It provides new cells for repair and regeneration and for

healing of the wounds. (e.g., cells of the upper layer of
the epidermis, cells of the lining of the gut, and blood
cells).

It helps in asexual reproduction by fragmentation, budding,

stem cutting, etc.

Somatic variations when maintained by vegetative

propagation can play an important role in speciation.

MITOGENS AND MITOTIC POISONS

The agents which stimulate cell division are called mitogens,

e.g., cytokinins, some steroids, however, some chemicals
inhibit cell division, these are called mitotic poisons, e.g.,
azides, cyanides, colchicine, chalones, etc. Colchicine is an
alkaloid derived from Colchicum autumnale. It interferes with
spindle formation and arrests cell division at metaphase.
Polyploidy in plants can be induced by the application of
colchicine.

Table : Difference between the Cytokinesis of Animal cell and

Plant cell.

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 MEIOSIS
It is a division that occurs in mature diploid reproductive
cells (2x) in which the nucleus divides twice but
chromosome (DNA) replicates only once to form four haploid
cells, each having half the number of chromosomes present
in the parent cell. As it causes a reduction in the number
of chromosomes, it is known as reduction division. Meiosis
in a cell occurs only once. The so formed haploid cells do
not further undergo meiosis because there is no
synaptonemal complex in haploid genome.

It is reported in diploid germ cells of sex organs (e.g.,

primary spermatocytes of testes to form male gametes called
spermotozoa and primary oocytes to form female gametes

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 called ova in animals) and in pollen mother cells
(microsporocytes) of anther and megasporocyte of ovules of
ovary of flowers in plant to form the haploid spores. The
study of meiosis in plants can be done in young flower
buds.

Meiosis I is initiated after the parental chromosomes have

replicated to produce identical sister chromatids during
the S phase.

Meiosis involves pairing of homologous chromosomes and

recombination between them.

Four haploid cells are formed at the end of meiosis II.

Process of meiosis : Meiosis is completed in two steps,

Meiosis I and Meiosis II. Each of them is further divisible
into different stages as in mitosis.

Meiosis I  Meiosis II
Prophase I  Prophase II

Metaphase I  Metaphase II

Anaphase I  Anaphase II
Telophase I  Telophase II

MEIOSIS-I

It results in the formation of two haploid cells from one

diploid cell. The daughter cells are, therefore, haploid but
with 2n DNA content. It is divided into four phases i.e.,
prophase, metaphase, anaphase, telophase.

(1) Prophase-I
It is the longest phase of karyokinesis of meiosis. It is
again divisible into five subphases i.e., leptotene, zygotene,
pachytene, diplotene and diakinesis.

(a) Leptotene/Leptonema

Chromosomes are long thread like with chromomeres (i.e.

linear series of darkly stained swollen areas) on it.

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 Each chromosome has its similar structure or homologue
known as homologous chromosomes derived from different
parents either paternal or maternal.

Leptonema chromosomes have a definite polarization and form

loops whose ends are attached to the nuclear envelope at
points near the centrioles, contained within an aster. Such
peculiar arrangement is termed as bouquet stage (in
animals) and synzetic knot (in plants like Lilium).

Lampbrush chromosome found in oocyte of amphibians is seen

in leptotene.

(b) Zygotene/Zygonema

Pairing or "Synapsis" of homologous chromosomes takes place

in this stage.

Paired chromosomes are called bivalents which by further

molecular packing and spiralization become shorter and
thicker.

Pairing of homologous chromosomes in a zipper-like fashion.

On observing under an electron microscope, a filamentous

ladder like nucleoproteinous complex, called synaptonemal
complex helps in adherence of the visible sister
chromatids.

(c) Pachytene/Pachynema

In the tetrad, two similar chromatids of the same

chromosome are called sister chromatids while chromatids
belonging to different chromosomes of the homologous pairs
are termed as non-sister chromatids.

Crossing over i.e., exchange of segments between non-sister

chromatids of homologous chromosomes occurs at this stage.

It takes place by breakage and reunion of chromatid segments.

Breakage called nicking is assisted by an enzyme endonuclease
and reunion termed annealing is done by an enzyme ligase.

A tetrad consists of two sets of homologous chromosomes

each with two chromatids. Chromatids of pachytene

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 chromosomes are attached with centromere. Each tetrad has
four kinetochores (two sisters and two homologous).

A number of electron dense bodies about 100 nm in diameter

are seen at irregular intervals within the centre of the
synaptonemal complex known as recombination nodules. These
are believed to be sites having multienzyme recombinase
complex required for crossing over.

(d) Diplotene/Diplonema

At this stage, the paired chromosomes begin to separate

(desynapsis) and terminalisation starts.

Cross is formed at the place of crossing over between non-

sister chromatids.

Homologous chromosomes move apart and remain attached to

one another at specific points called chiasmata.

At least one chiasma is formed in each bivalent.

Cell Cycle And Division [Notes] 14

 This stage remains as such for a long time. For example,
all the oocytes of human female reach the diplotene stage
in the fifth month of foetus and remain so for many years
till ovulation is to occur. In oocytes of many fishes,
amphibians, reptiles and birds, the bivalents elongate and
become converted into lampbrush chromosomes for synthesis
of specific biochemicals.

(e) Diakinesis

Terminalization of chiasmata takes place.

Nuclear membrane and nucleolus degenerates.

Chromosomes recondense and tetrad moves to the metaphase

plate.

Formation of spindle occurs.

When the diakinesis of prophase-I is completed then cell

enters into metaphase-I.
(2) Metaphase-I

Chromosomes align at the equator.

Bivalents arrange themselves in two parallel equatorial or

metaphasic plates. Each equatorial plate has one genome.

Each homologous chromosome has two kinetochores and both

the kinetochores of a chromosome are joined to the
chromosomal or tractile fibre of same side.

(3) Anaphase-I

It involves separation of homologous chromosomes which

start moving towards opposite poles so that each tetrad is
divided into two daughter dyads. So, anaphase-I involves
the reduction of chromosome number, this is called
disjunction.

Segregation of Mendelian factors or independent assortment

of chromosomes takes place, in which the paternal and
maternal chromosomes of each homologous pair segregate
during anaphase-I thus, introducing genetic variability.

(4) Telophase-I

Cell Cycle And Division [Notes] 15

 Two daughter nuclei are formed but the chromosome number is
half the chromosome number of mother cell.

Nuclear membrane reappears.

After telophase I, cytokinesis may or may not occur.

Interkinesis (Intrameiotic interphase)

Generally there is no interphase between meiosis-I and
meiosis-II. A brief interphase called interkinesis, or
intrameiotic interphase occurs. There is no replication of
chromosomes because chromosomes are already in replicated
state. Centrosome may replicate in animal cells.
Significance of Meiosis-I

It separates the homologous chromosomes to reduce the

chromosome number to the haploid state, a necessity for
sexual reproduction.

It introduces variation by forming new gene combinations

through crossing over and random assortment of paternal and
maternal chromosomes.

MEIOSIS-II

It is also called equational or homotypical division

because the number of chromosomes remain same as after
meiosis-I.

It involves the separation of two chromatids of each

chromosome and their movement to separate cells.

It is divided into four phases i.e., Prophase-II,

Metaphase-II. Anaphase-II and Telophase-II.

(1) Prophase-II

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.

(2) Metaphase-II

Cell Cycle And Division [Notes] 16

 At this stage, the chromosomes align at the equator and
microtubules from opposite poles of the spindle get attached
to kinetochores of sister chromatids.
(3) 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 towards opposite poles of the
cell.

(4) Telophase-II

Meiosis ends with telophase II, in which 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.

Cytokinesis-II is always present and occurs by cell furrow

formation in animal cells and cell plate formation in plant
cells.

Types of meiosis II

Gametic/Terminal meiosis : In many protozoans, all animals

and some lower plants, meiosis takes place before
fertilization during the formation of gametes.

Zygotic or Initial meiosis : In fungi, certain protozoan

groups, and some algae, fertilization is immediately
followed by meiosis in the zygote, and the resulting adult
organisms are haploid. Such a meiosis is said to be zygotic
or initial. This type of life cycle with haploid adult and
zygotic meiosis is termed the haplontic cycle.

Sporogenetic / Intermediate meiosis : Spore mother cells of

sporophytic plant undergo meiosis to form haploid spores in
the sporangia. Haploid spore germinates to form haploid
gametophyte which produces haploid gametes by
mitosis. Haploid gametes fuse to form a diploid zygote
which develops into diploid sporophyte by mitotic
divisions. e.g., in higher plants like pteridophytes,
gymnosperms and angiosperms.

Cell Cycle And Division [Notes] 17

 Significance of Meiosis-II

Constancy of chromosome number in successive generations is

brought by this process.

It helps in introducing variations and mutations.

It maintains the amount of genetic material.

The four daughter cells will have different types of

chromatids.

Cell Cycle And Division [Notes] 18

 Fig. : Stages in Meiosis

Cell Cycle And Division [Notes] 19