NANYANG JUNIOR COLLEGE H2 Biology

Cell and Nuclear Division

TOPIC F: CELL & NUCLEAR DIVISION

(MITOSIS & MEIOSIS)

Learning Outcomes
Core Idea 2 – Genetics and Inheritance
Candidates should be able to:
(n) Describe the events that occur during the mitotic cell cycle and the main stages of mitosis
(including the behaviour of chromosomes, nuclear envelope, cell membrane and centrioles).
(o) Explain the significance of the mitotic cell cycle (including growth, repair and asexual
reproduction) and the need to regulate it tightly. (Knowledge that dysregulation of checkpoints
of cell division can result in uncontrolled cell division and cancer is required, but detail of the
mechanism is not required.)
(s) Describe the events that occur during the meiotic cell cycle and the main stages of meiosis
(including the behaviour of chromosomes, nuclear envelope, cell surface membrane and
centrioles). (Names of the main stages are expected, but not the sub-divisions of prophase.)
(t) Explain the significance of the meiotic cell cycle (including how meiosis and random fertilisation
can lead to variation).

Content Outline
1. The Role of Nucleus and Chromosomes in Cell Division
(a) Chromatin Structure
(b) Chromosome Structure
(c) Chromosome Number
(d) Homologous Chromosomes
2. The Cell Cycle
(a) Interphase
(b) Mitosis
(c) Cytokinesis
(d) Significance of Mitosis
(e) The control of the mitotic cell cycle
(f) Meiosis
(g) Significance of Meiosis
(h) Non-disjunction

References
1. Campbell, N. A. And Reece J.B. (2011) Biology. Chapter 12 :The Cell Cycle and Chapter 13:
Meiosis and Sexual Life Cycles. 9th ed. Benjamin Cummings Publishing, Inc.

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1. The Role of Nucleus and Chromosomes in Cell Division

All new cells are derived from pre-existing cells through cell division, a part of the cell cycle
which includes:
 Interphase - cell growth and synthesis of cell materials
 Nuclear division - nucleus divides to form two nuclei
 Cytokinesis - cytoplasm divides to form two daughter cells
The nucleus is responsible for carrying out cell division.

(a) Chromatin Structure (X-ref to: Org and Control of Euk Genome notes)
 is the complex of DNA and proteins.
 is the less condensed form of chromosome.
 dispersed as a mass of long, thin fibres when cell is not dividing and during interphase.
 Two types of chromatin: Heterochromatin (highly condensed) and euchromatin (less
condensed).

 The degree of condensation may change:

o Chromatin condenses by coiling to form chromosomes during cell division.
o Chromatin decondenses for polymerases to gain access to specific regions of DNA
for replication, gene expression or repair.

10nm

DNA packed into chromatin and chromosome

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(b) Chromosome Structure

During cell division, chromatin condenses into chromosomes, which may exist as
unduplicated chromosomes or duplicated chromosomes.

Diagram showing duplicated chromosome and unduplicated chromosome

(i) Unduplicated chromosome

 consists of a single DNA molecule

(ii) Duplicated chromosome (also known as replicated chromosome)

 Four-arm structure visible under light microscope
 consists of two DNA molecules
 consists of two identical sister chromatids joined at the centromere
 Identical sister chromatids are formed due to semi-conservative DNA
replication during synthesis phase of interphase.

 Centromere is a constricted region of satellite DNA (highly repetitive DNA

sequence).
o Bound to kinetochore proteins which allow for attachment of spindle
fibres.
o The centromere holds two sister chromatids together and are involved
in chromosome movement during cell division.

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(c) Chromosome Number

(i) Diploid
 Diploid cells contain two sets of chromosomes.
 Diploid cell has a diploid number of chromosomes, 2n, where n represents the
number of chromosomes in a single set.
 The number of chromosomes in somatic cells (i.e. all cells except sex cells) is fixed
for a species. Each sexually reproducing species has a characteristic diploid number
and haploid number.
 For example, each human somatic cell has 46 chromosomes and the diploid
number is 46.
 Each somatic cell contains two sets of chromosomes (2n = 46). One set
contains 23 different chromosomes (n = 23).
 This can be clearly seen in a karyotype, an ordered display of chromosomes.

Identical DNA molecule

Identical DNA molecule

Karyotype of a human male

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(ii) Haploid
 Gametes (sex cells) contain a single set of chromosomes
 Haploid cells have a haploid number of chromosomes (n).
 For example, a human egg or human sperm has 23 chromosomes and the
haploid number is 23.
 Each sex cell contains one set of chromosomes (n = 23).
 The 23 different chromosomes consist of the 22 autosomes plus a single sex
chromosome.
 A human egg contains 22 autosomes and an X chromosome,
 A human sperm contains 22 autosomes and either an X or a Y chromosome.

Haploid Number = 3 Diploid Number = 6

Table showing chromosome numbers of human, fruit fly and dog

Human Fruit Fly Dog
Diploid number 46 8
Number of chromosomes in a gamete
Number of different types of chromosomes 39

Diagram describing chromosome numbers

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(d) Homologous Chromosomes

In a diploid cell, each type of chromosome exists as a pair of homologous chromosomes,
with each chromosome known as a homolog.
In human diploid cell, we inherit one chromosome of each pair from each parent. Thus,
the 46 chromosomes in our somatic cells are actually two sets of 23 chromosomes – a
maternal set (from the female parent) and a paternal set (from the male parent).

Homologous chromosomes:
 have the same length,
 have the same position of centromeres.
 have the same genes at the same loci.
 Homologous chromosomes may have different alleles from each other.
o Alleles are alternative forms of the same gene with a slight different nucleotide
sequence.
 One homolog is derived from maternal parent while the other is derived from the
paternal parent.
 Homologous chromosomes pair to form bivalents during prophase I of meiosis.
 Each homolog consists of double structure containing two identical sister
chromatids during prophase I of meiosis.

The two sex chromosomes are an important exception to the general pattern of homologous
chromosomes.

Homologous chromosomes
For example,
 When the two alleles coding for the gene A, on a pair of homologous chromosome are
the same, this organism is homozygous for the trait.
 When the two alleles coding for the gene D, on a pair of homologous chromosome are
different, the organism is heterozygous for the trait.
 When the genes that code for A, D and F are found on the same chromosome, they are
said to be linked (i.e. linked genes).

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2. The Cell Cycle

The cell cycle is the life of a cell from the time it is first formed from a dividing parent cell until
its own division into two daughter cells.
 Three stages: interphase, nuclear division and cytokinesis.
 Two types of nuclear division:
o mitosis for all somatic cells and
o meiosis in sexual reproductive organs to form gametes / sex cells.
 The duration of cell cycle varies for different types of tissue within a single organism.

(a) Interphase
Cells prepare for mitosis or meiosis in 3 sub-phases:
 First growth phase or G1 phase
 Synthesis or S phase
 Second growth phase or G2 phase

Phase Events that occur

G1  Synthesis of organelles e.g. endoplasmic reticulum and mitochondria / chloroplasts
(by binary fission)
 Nucleolus actively synthesises ribosomal RNA for formation of ribosomes.
 Synthesis of proteins
 Increase in cytoplasmic volume, resulting in an increase of cell size.
Synthesis  DNA semi-conservative replication occurs, resulting in doubling of DNA content.
 Histone proteins are synthesized.
 Newly synthesized DNA molecules are wound tightly around the histone proteins.
 Each chromatin fibre is composed of two identical DNA molecules.
G2  Intensive synthesis of organelles e.g. endoplasmic reticulum and mitochondria /
chloroplasts (by binary fission)
 2 centrosomes have formed by duplication of centrosome (started in G1). Each
centrosome is made up of a pair of centrioles (in animal cells).
 Synthesis of proteins – e.g. tubulins for assembly of microtubule in spindle fibre
 Synthesis of ATP for energy.

Cell cycle

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(b) Mitosis
Nuclear division:
 Nucleus of parent cell divides once to produce two genetically identical daughter
nuclei with same number and types of chromosomes as parent nucleus.
 Four main phases: prophase, metaphase, anaphase, telophase

(i) Prophase
 Chromatin fibre becomes more tightly coiled, condensing into chromosomes.
 Each duplicated chromosome appears as two identical sister chromatids joined
at their centromeres.
 Centrosome organise microtubules into spindle fibres and the radial array of short
microtubules extending from each centrosome are called asters.
 Centrosomes migrate to opposite poles of the cell by lengthening of microtubules.
o In animal cells, each pair of centrioles migrates to opposite poles of the cell.
 Nucleolus disperses and seems to disappear.
 Nuclear envelope fragments.

Centrosome:
Animals have a pair of centrioles

Chromatin
condensing

Diagrams of Prophase

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(ii) Metaphase (longest phase in mitosis)

 Microtubules from centrosome are attached to kinetochore at the centromere of
each chromatid of chromosome, becoming kinetochore microtubules.
 Centromeres of chromosomes are aligned along the metaphase plate (an
imaginary plane equidistant between the two poles of the cell) by microtubules.
 Non-kinetochore microtubules interact with those from the opposite pole of the
spindle.

Diagrams showing Metaphase and chromosomes attached to kinetochore

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(iii) Anaphase (shortest phase in mitosis)

 Centromere separates and two sister chromatids separate, thus becoming two
daughter chromosomes.
 Daughter chromosomes migrate toward opposite poles of the cell, with the
centromere leading the way as kinetochore microtubules shorten.
 Non-kinetochore microtubules lengthen, leading to cell elongation.
 By the end of anaphase, the two poles of the cell have equivalent and complete set
of chromosomes.

Diagrams showing Anaphase

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(iv) Telophase
 Nuclear envelopes reform from the fragments of the endomembrane system to
form two nuclei.
 Nucleolus reappears.
 Chromosomes become less condensed to form chromatin.
 Microtubules disperse by depolymerising.

Diagram showing telophase with cytokinesis

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Early prophase

Mid prophase

Late prophase

Metaphase

Early anaphase

Late anaphase

Telophase

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(c) Cytokinesis
Cytokinesis usually occurs by late telophase, dividing the cytoplasm evenly between the
two daughter cells that appear shortly after the end of mitosis.

In animal cells,
 Cell surface membrane invaginates towards the metaphase plate
 A ring of actin microfilaments contracts by interacting with myosin molecules.
 The cleavage furrow deepens until the parent cell is pinched into two, producing two
daughter cells.

In plant cells,
 Golgi vesicles that contain cell wall material (cellulose) move along microtubules,
towards the metaphase plate and fuse together to produce a cell plate.
 Cell plate enlarges as more vesicles fuse with it, until its surrounding membrane
fuses with cell surface membrane along the perimeter of the cell.
 A new cell wall is formed between the two daughter cells.

Diagram of animal and plant cytokinesis

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(d) Significance of mitosis

 Mitosis confers genetic stability between generations of cells.
 Each parent cell produces two genetically identical daughter cells with same number
and types of chromosomes as parent nucleus (no variation in genetic information).
 The daughter cells have identical genetic information as the parent cell, due to semi-
conservative replication of parental DNA during synthesis phase of interphase.
 This enables growth, repair and asexual reproduction.

(i) Organism Growth

 To grow from one cell to a multicellular organism, all daughter cells must be
genetically identical to the parent cell. However, genetically identical cells are able
to differentiate to different cell types due to differential gene expression.

(ii) Tissue Repair

 Cells that are lost, damaged or worn-out are replaced by genetically identical
cells.

(iii) Asexual reproduction

 Involves one single parent, producing offsprings that are genetically identical to
parent known as clones.
 This is common in plants - vegetative propagation, e.g. bulbs in onions,

(Asexual reproduction includes budding in yeasts, fragmentation in starfish and binary

fission in bacteria).

Diagram showing organisms that reproduce asexually

The organism on the left is the hydra, which reproduces by budding. The bud is a localised mass of
mitotically dividing cells that develops into a small hydra and detaches from its parent.
The organism on the right is the redwood tree. All the trees in this circle of redwood arose
asexually from a single parent tree, whose stump is in the centre of the circle.

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(e) The Control of the Mitotic Cell Cycle

The rate of cell division is highly regulated at various checkpoints in the cell cycle –
control points to determine if the cell will proceed to the next stage.

G1

Diagram showing Cell Cycle Control

There are three checkpoints during the mitotic cell cycle at G1, G2 and metaphase.
 G1 checkpoint ensures that the cell size is adequate and there are sufficient nutrients
available and growth factors present for cell to undergo mitosis.
o If the cell does not pass the G1 checkpoint, the cell will exit the cycle, switching into a
nondividing state called the G0 phase.
 G2 checkpoint ensures that the cell size is adequate and semi-conservative DNA
replication has been completed successfully.
 Metaphase checkpoint ensures that all chromosomes are attached to spindle fibres /
microtubules at the metaphase plate before proceeding to anaphase.
 When all checkpoints are passed, cell cycle continues to the next stage.

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(f) Meiosis
Nuclear division:
 Nucleus of a parent cell divides twice to produce four genetically non-identical
haploid daughter nuclei.
 Each daughter nuclei contains half the number of chromosomes in the parent nucleus
by reducing two sets of chromosomes to 1 set of chromosomes.
 It is vital for sexual reproduction and takes place in the reproductive organs of plants
and animals.
 Meiosis occurs after interphase, followed by cytokinesis

Overview of Meiosis

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(i) Prophase I
 Chromatin fibre becomes more tightly coiled, condensing into chromosomes.
 Each duplicated chromosome appears as two identical sister chromatids joined
at their centromeres.
 Homologous chromosomes pair to form bivalents in a state called synapsis.
 Crossing over occurs between non-sister chromatids of homologous
chromosomes resulting in chiasmata formation.
o the exchange of corresponding DNA segment between non-sister
chromatids.
 Centrosome organise microtubules into spindle fibres and the radial array of short
microtubules extending from each centrosome are called asters.
 Centrosomes migrate to opposite poles of the cell by lengthening of microtubules.
o In animal cells, each pair of centrioles migrates to opposite poles of the cell.
 Nucleolus disperses and seems to disappear.
 Nuclear envelope fragments.

Diagram of Prophase I

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(ii) Metaphase I
 The bivalents (pairs of homologous chromosomes) align themselves at the
metaphase plate, with one chromosome in each pair facing each pole.
 The arrangement of chromosome of each bivalent is independent of the
arrangement of the other bivalents (i.e. independent assortment).
 Both chromatids of one homolog are attached to kinetochore microtubules from
one pole; those of the other homolog are attached to microtubules from the opposite
pole.
 Non-kinetochore microtubules interact with those from the opposite pole of the
spindle.

Diagram of Metaphase I

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(iii) Anaphase I
 Each homolog of a bivalent separates / Homologous chromosomes separate
but not sister chromatids. Centromere of each chromosome does not separate.
 Homologous chromosomes migrate toward opposite poles of the cell, with
the centromere leading the way as kinetochore microtubules shorten.
 Non-kinetochore microtubules lengthen, leading to cell elongation.

Diagram showing Anaphase I

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(iv) Telophase I
 In some species, nuclear envelopes reforms from the endomembrane system,
forming two nuclei.
 Each nucleus has a haploid set of chromosomes (duplicated chromosomes).
 Thus meiosis I is known as the reductional division because it halves the number
of chromosome sets per cell.
 Nucleolus reappears.
 Chromosomes become less condensed to form chromatin.
 Microtubules disperse by depolymerising.
 Cytokinesis usually occurs simultaneously with telophase I, forming two haploid
daughter cells. No replication of DNA occurs between meiosis I and meiosis II.

 In other species, these processes (including the reforming of nuclear envelope) are
skipped and the cell carries on directly to Prophase II.

Comparison between Mitosis and Meiosis I

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(v) Prophase II
 Chromatin fibre becomes more tightly coiled, condensing into chromosomes.
 Nucleolus disperses and seems to disappear.
 Nuclear envelope fragments.

(vi) Metaphase II
 Microtubules from centrosome attach to kinetochore at the centromere of each
chromatid of chromosome, becoming kinetochore microtubules. (Sister
chromatids may no longer be identical as crossing over may have occurred in
Prophase I).
 Centromeres of chromosomes are aligned along the metaphase plate,
perpendicular to the metaphase plate in metaphase I by microtubules.
 Non-kinetochore microtubules interact with those from the opposite pole of the
spindle.

Diagram of Meiosis II

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(vii) Anaphase II
 Centromere separates and two non-identical sister chromatids separate, thus
becoming two daughter chromosomes.
 Daughter chromosomes migrate toward opposite poles of the cell, with the
centromere leading the way as kinetochore microtubules shorten.
 Non-kinetochore microtubules lengthen, leading to cell elongation.

(viii) Telophase II
 Nuclear envelopes reform from the fragments of the endomembrane system to
form two nuclei.
 Nucleolus reappears.
 Chromosomes become less condensed to form chromatin.
 Microtubules disperse by depolymerising.
 Four haploid daughter nuclei will arise from one diploid parent nuclei.
 Cytokinesis then occurs as in mitosis.

Comparison between Mitosis and Meiosis

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(g) Significance of meiosis

 The production of haploid gametes by meiosis is important prior to fertilisation in sexual
reproduction, where fusion of gametes from different parents occurs to form a zygote.
 Meiosis ensures maintenance of chromosomal number in offspring and prevent
doubling of chromosomal numbers during fertilisation
 Meiosis generates genetic variation in offspring by producing recombinant gametes
through:
(i) Crossing over between non-sister chromatids of homologous chromosomes
(ii) Independent assortment of homologous chromosomes

There are three processes that contribute to the genetic variation arising from sexual
reproduction:
(i) Crossing over between non-sister chromatids of homologous chromosomes
(ii) Independent assortment of homologous chromosomes and
(iii) Random fertilisation

 Note: The source of new genetic variation is due to spontaneous mutations of DNA
(e.g. errors that occur during semi-conservative replication), which results in different
nucleotide sequences of a gene (alleles).
 Genetic variation in a population (genetically different individuals of the same species)
is the basis for natural selection.

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(i) Crossing over

 Crossing over occurs between non-sister chromatids of homologous
chromosomes resulting in chiasmata formation, during prophase I of meiosis.
 Chiasmata (singular = chiasma), the X-shaped regions exist at the point where
crossing over has occurred.
 Crossing-over result in the exchange corresponding DNA segment of chromatids,
thus separating alleles of linked genes and creating new allelic combinations
in the chromatid.

 Crossing-over is a process that involves:

o Breakage of a corresponding DNA segment from each non- sister chromatid,
o Exchange of the corresponding DNA segment between non-sister chromatids,
and
o Rejoining of the DNA segment to the other chromatid, resulting in chiasmata.

Diagram of crossing over

In the diagram above,

 Parental chromosome contains linked genes, V and B. One homolog contains
dominant V and B alleles while the other contains recessive v and b alleles.
 Crossing over occurs between two non-sister chromatids, resulting in an exchange
of allele for the gene B. This result in two recombinant chromosomes; one
contains V and b alleles and the other contains v and B alleles.
 Four different gametes with different allelic combinations are produced: VB, Vb,
vB and vB.

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(ii) Independent assortment of homologous chromosomes

 Independent assortment is the random orientation of pairs of homologous
chromosomes along the metaphase plate independent of other bivalents during
metaphase I
 Independent assortment of homologous chromosomes occurs during metaphase I
leading to independent segregation during anaphase I.

Diagram showing independent assortment and independent segregation

In the diagram above,

 During prophase I: Bivalents are aligned where the maternal homolog and paternal
homolog faces each side of the pole
 At metaphase I: The two pairs of homologous chromosomes randomly orientate
themselves along the metaphase plate. Thus, the first arrangement is equally as
likely as the second arrangement.
 During anaphase I: Homologous chromosomes separate independently of other
bivalents (i.e. independent segregation).
 Note: Only two of the four combinations of daughter cells shown would result from
meiosis of a single diploid cell, because a single parent cell would have one or the
other possible chromosomal arrangement at metaphase I, but not both.
 However, the population of daughter cells resulting from meiosis of a large number
of diploid cells contains all four types of gametes in approximately equal
numbers.

 The number of possible chromosomal combinations in a gamete is 2n, where n =

haploid number of the organism.
o In humans, the number of possible chromosomal combination is 223, thus 223 different
combinations of gametes can be produced by an individual.

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(iii) Random fertilisation

 During fertilisation, genetic material from two different individuals is combined by
random fusion of gametes.
o In humans, the number of possible combinations of zygote is 223 x 223.

(h) Non-disjunction (KIV: Topic G Mutation and cancer notes)

 An error during meiosis that result in chromosomal aberration, aneuploidy.
 Non-disjunction occurs when:
o A pair or pairs of homologous chromosome fail to separate during Anaphase I of
meiosis or
o sister chromatids fail to separate during Anaphase II of meiosis.
 This results in aberrant gametes:
o gamete receives two of the same type of chromosome / extra chromosome (n+1)
or
o gamete with no copy of a particular type of chromosome / lacks a chromosome (n-1)

 If an aberrant gamete fuses with a normal gamete, the zygote will have an abnormal
number of chromosomes (2n+1) or (2n-1), a condition known as aneuploidy.
For example,
o Down syndrome (trisomy 21),
o Turner syndrome (XO),
o Edwards syndrome (trisomy 18) and
o Patau syndrome (trisomy 13).

Diagram showing Non-disjunction

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