3 Cell replication

Students:
INQUIRY QUESTION •• model the processes involved in cell replication, including but not limited to:
– mitosis and meiosis (ACSBL075) CCT ICT
How important is it for
– DNA replication using the Watson and Crick DNA model, including nucleotide composition, pairing and
genetic material to be bonding (ACSBL076, ACSBL077)
replicated exactly?
•• assess the effect of the cell replication processes on the continuity of species (ACSBL084) ICT
Biology Stage 6 Syllabus © NSW Education Standards Authority for and on behalf of the Crown in right of the State of New South Wales, 2017
Shutterstock.com/Spectral-Design

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 Have you ever wondered why it is so

Nature Picture Library/Andrew Parkinson

important for genetic material to be
replicated exactly? Each living creature
has a different genetic code that carries
instructions for its own production. How
do the genetic instructions for wombats
differ from those for koalas, dogs, penguins
or even mushrooms? Parents give rise to
offspring of the same species, with a similar
genetic makeup to their own, so they can
survive in the same environment. It is
essential that organisms pass on their own FIGURE 3.1 Genetic instructions for a sheep differ from those of
unique genetic instructions or information a bird, grass or flowers.
from one generation to the next, to ensure
the continuity of the species.
How can something as tiny as a cell carry all the instructions needed to make a large, complex, living
organism? When scientists began searching for the ‘code of life’ in cells, they realised that the genetic
code would need to be in molecular form to be small enough to fit all the instructions into a cell. As
chromosomes are composed of DNA (deoxyribonucleic acid) and protein, scientists deduced that the
genetic code must be carried by one of these two macromolecules – either in proteins or in DNA.
It was expected that the genetic code would be complex, because it needs to store large amounts of
information accurately over long periods of time. The genetic code would also need to be copied exactly
and any errors that arose during copying would need to be easily fixed. Most scientists favoured protein
as the carrier of the genetic instructions, because proteins are complex and varied, being made up of at
least twenty different amino acids.
In 1953, James Watson and Francis Crick discovered the structure of DNA and how it held the ‘code
of life’. What surprised most biologists was that genetic information could be carried by a molecule that
uses an ‘alphabet’ made up of only four ‘letters’: the bases adenine (A), cytosine (C), guanine (G) and
thymine (T). In the decade following 1953, the combined work of many scientists led to an understanding
of how the information in DNA can lead to specific proteins being produced.
In this chapter, you will find out how Watson and Crick worked out the structure of DNA, as well as
when and how genetic material is replicated and why it is so important for replication to be exact.

3.1 Cell division – mitosis and meiosis

Why do cells need to divide? What mechanisms are in place for the faultless transmission of genetic
information from parent to daughter cells during division? When a cell divides, how does every daughter
cell get an exact copy of the genetic instructions? As you study the mechanisms of cell division, you will
discover how this is achieved.
There are two types of cell division: mitosis and meiosis.
In unicellular protist organisms, cell division by asexual binary fission is common – one organism
becomes two and no sex cells or gametes are involved. In multicellular organisms, cell division by mitosis
leads to the formation of two new identical cells that contribute to the growth of the organism.
Meiosis is a different type of cell division. It gives rise to gametes that transmit genetic material
from one generation to the next during sexual reproduction. Meiosis will be dealt with in more detail in
Chapter 5.

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 The role and importance of mitosis
Mitosis plays an important role in:
◗◗ growth of multicellular organisms – from fertilised egg to embryo, from infant to adult, growth relies
on mitotic division followed by cell assimilation, enlargement and differentiation
◗◗ repair of damaged tissue and replacement of worn-out cells
◗◗ asexual reproduction – for example, growth of plants from cuttings, and the cloning of organisms
◗◗ genetic stability – mitosis ensures the precise and equal distribution of chromosomes to each daughter
nucleus, so that all resulting cells have the same chromosome number and genetic information as
each other and the original parent cell.
Most multicellular organisms start life as a single fertilised egg cell or zygote, which grows into an
See Chapter 4, embryo. Embryonic cells are able to divide repeatedly and are pluripotent (that is, they have the potential
page 129 for
information on to develop into any type of tissue) (Fig. 3.2). These cells are termed embryonic stem cells. You may have
stem cells and gene heard of these already, because the use of embryonic stem cells is a subject that often draws media
expression.
attention and discussion among people. This is because the use of these cells in therapeutic medicine
and for research involves the destruction of the embryo, a contentious issue. This will be dealt with in
greater detail in Chapter 8.
In mature organisms, not all cells continue to divide. As cells differentiate and specialise, they form
tissues, and some of these cells lose the ability to divide by mitosis. Dividing cells remain at particular
locations within the adult body; for example, dividing
2-cell stage 4-cell stage 64-cell stage cells in the basal layer of the skin replace dead
surface skin cells, and cells in bone marrow give rise
to blood cells. Adult stem cells occur in almost every
type of tissue but they are pre-specialised; for example,
brain stem cells can only make brain tissue and heart
stem cells can only make cardiac tissue. Cells in bone
marrow can give rise to all types of blood cells, and so
they are termed ‘multipotent’. These adult stem cells
Blastula can therefore not be used as widely in all types of tissue
regeneration as pluripotent embryonic cells. Researchers
are continuing to investigate how adult stem cells can
be made pluripotent, so they can be used in tissue
Cells have the and organ regeneration. Unfortunately, reversing pre-
potential to
become any
specialisation of cells in tissue culture tends to make
cell type. cells turn cancerous, so there is still much research to be
done in this area.
FIGURE 3.2 Early stages of mammalian embryonic development: mitotic In plants, tissue that can divide to form any other
division of pluripotent embryonic stem cells tissue is known as meristem and occurs at particular
locations in the plant, such as the tip of the stem and
the tip of the root.

The role and importance of meiosis

Refer to Chapter 2 Meiosis is the type of cell division that occurs in the sexual reproductive organs of a plant or animal, and
for information on results in the formation of gametes (sex cells).
asexual and sexual
reproduction, and During sexual reproduction, two parents are involved in passing genetic material to the offspring.
to Chapter 9 for To prevent the chromosome number from doubling in each successive generation, a mechanism must
information on
cloning. exist to ensure that each parent contributes only half of his or her chromosomes to the new offspring.
This mechanism is meiosis, a type of reduction division in cells in the reproductive organs of both plants
and animals. Meiosis ensures that the chromosome number of each species is maintained (not doubled)
during sexual reproduction (Fig. 3.3).

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 FIGURE 3.3 Life
cycle of humans
showing the
importance of meiosis
Fusion to (formation of haploid
form zygote Zygote (2n) gametes) and mitosis
(n to 2n) (differentiation and
growth)

Egg (n) Sperm (n)

Juvenile (2n)
Meiosis
(2n to n)

Meiosis Mitosis
(2n to n) (differentiation
and growth)

Adults (2n)
Haploid stages
Diploid stages

When a cell divides by meiosis, DNA replicates before division. The cell then undergoes two
successive divisions:
◗◗ meoisis I, where the diploid cell divides into two haploid cells and the chromosome number is halved
◗◗ meiosis II, where the two cells each divide again, resulting in four haploid daughter cells (called
a tetrad).
Each daughter cell has half the original number of chromosomes that the parent cell had.
These resulting daughter cells, or gametes in plants and animals, are:
◗◗ egg cells (in ovaries) and sperm cells (in testes) in animals
◗◗ pollen grains (in anthers) and egg cells (in ovules) in seed-producing plants.
Gametes are often referred to as ‘vehicles of inheritance’ because they carry genes from one generation
to the next.
Meiosis is also the process by which genetic variation is introduced into a species. The process of
meiosis and its contribution to genetic variation is dealt with in detail in Chapter 5.

Cell division: why does DNA need to replicate exactly?

An organism’s genetic code contains instructions for every feature of the individual, from its
biochemistry, which allows it to function properly, to its physical features and body size. The genetic
code even holds instructions for how its own genes are expressed and therefore for how all biological
information in the organism is read, processed and converted (translated) from its coded form (DNA)
to its end product (protein).
When a cell divides into two, it is vital that the DNA passed on to each daughter cell is an exact
copy, so that each generation of cells contains the same genetic instructions. In unicellular organisms,
the genetic code passed from parent cells to offspring maintains the genetic stability of the species. In
multicellular organisms, all cells making up the body must contain the same genetic code so that they
can function in a controlled and coordinated way.

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 Along the chromosome are long sequences of base pairs in specific regions of DNA that form units
called genes. Genes code for the production of proteins, which in turn make up a large proportion of the
structure of cells and also regulate the functioning of cells. For example, many body parts such as bones,
hair, skin, muscles and connective tissue contain proteins, which are coded for by genes. Some proteins
are enzymes (as you learned in Year 11), and these carry out and control all cell functions. Each step of
every biochemical reaction in cells, such as chemical respiration and photosynthesis, is controlled by
enzymes.
It is therefore vital for the functioning of cells and the stability of species that the genetic code is
passed on without error. During mitosis, DNA is replicated precisely and the copies of all genes are
divided up equally between the resulting cells. The cells are therefore clones of each other. In exploring
the process of mitosis, you will see how this happens in somatic (body) cells.
All cells need to make particular end-products and function in a coordinated manner. DNA codes for
proteins used inside the cell, as well as proteins such as hormones and enzymes that are exported and
coordinate functioning or direct protein synthesis in other cells. A change in DNA, known as a mutation,
may change the end product and, as a result, may have a harmful effect on the cell, by interfering with
its structure and/or functioning. Sometimes mutations have no effect, and occasionally they may be
beneficial. You will learn more about mutations and genetic variation in chapters 5 and 7.
KEY CONCEPTS

●● Binary fission is cell division that occurs in prokaryotes for asexual reproduction.
●● Mitosis is the nuclear division of eukaryotic cells for asexual reproduction (in unicellular
organisms and propagation in plants) and for growth and repair (in multicellular organisms).
●● Meiosis is the nuclear division of eukaryotic cells for gamete formation in sexual reproduction.
●● Many protists, plants and fungi can reproduce by mitosis (asexual reproduction).
●● Mitosis in multicellular organisms is a mechanism for tissue growth, maintenance and repair.
●● Meiosis is the type of cell division that occurs in the sexual reproductive organs of a plant
or animal and it results in the formation of gametes (sex cells) with the haploid number of
chromosomes. It also introduces genetic variation into a population.
●● In 1953, James Watson and Francis Crick discovered and modelled the double-helical structure
of DNA.
●● The sequence/order of the bases A, T, C and G in DNA holds information in a coded form.
●● DNA replication is vital for the continuation of a species, as it allows an organism to reproduce
its genetic code and pass it on.

The cell cycle

Cell cycle
Play the animation Cell division and enlargement occur in a repetitive sequence called the cell cycle (Fig. 3.4). Mitosis is only
and do the quiz
to check your one part of this cycle and usually takes about an hour or two. A complete cell cycle in actively dividing
understanding.
cells may take about 18–22 hours. The cell spends a large amount of time preparing for division (a stage
called interphase, which precedes mitosis).
The cell cycle has four main phases.
◗◗ G1 is a gap phase for cell growth before DNA replication. During this phase, cell enlargement takes
Cell cycle game place. Metabolic changes prepare the cell for division and the cell reaches a point at which it is
committed to division, then enters the next phase.
◗◗ S is a synthesis phase during which DNA replicates – that is, the DNA in the cell is copied so that each
dividing cell has two identical copies at the start of mitosis. At the end of cell division, one full copy
Cell cycle
ends up in each resulting daughter cell.
animations ◗◗ G2 is a second gap phase after replication, when enzymes in the cell check the duplicated chromosomes
Watch the five short for any errors and correct these, and the cytoplasmic materials accumulate in preparation for division.
animations on the cell
cycle and cancer. ◗◗ Mitosis (division of the nucleus) then occurs, followed by cytokinesis (division of the cytoplasm).
Cytokinesis marks the separation of one cell into two.

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 G0 FIGURE 3.4 The cell
Cell cycle stops cycle

G1 – cell growth

INT
Cellular contents,

ERP
excluding the
S – synthesis

HASE
chromosomes,
Each of the 46
are duplicated
chromosomes is
duplicated by the cell

Cytokinesis

Cytoplasm
divides
M – mitosis
Nucleus G2 – 9proof reading9
divides The cell checks the
duplicated chromosomes
for errors, making
any needed repairs
IT
M

O SI S

In actively dividing tissue, following cytokinesis, each cell will enter G1 again to repeat the cell cycle.
This begins with a period of assimilation and cell enlargement, where cells increase in size by assimilating
new materials into the cell boundaries and cytoplasm. This new material is made from the nutrients that Cell cycle
an organism acquires. For example, these nutrients are the end products of digestion in animal cells and revision
Explore in more
of photosynthesis in plant cells. detail the cell
Not all cells continue to go through cyclical divisions. The basal layers of cells that need to be replaced cycle and mitosis,
explained in
often, such as skin cells and cells lining the digestive tract, divide rapidly. Some adult cells do not divide diagrams and text.
frequently and others, such as nerve cells, may last a lifetime. These cells are said to be terminally
differentiated. They are in a phase called the G0 phase and only re-enter the cell cycle under special
circumstances.

Mitosis: division of the nucleus

Mitosis is the type of nuclear division that ensures that daughter cells receive the same number and
form (exact copies) of chromosomes as the parent cell. Therefore, before a parent cell can divide it must
make an accurate and complete replica of the genetic information encoded in its DNA. This needs to
be accurately and equally distributed to the resulting daughter cells. Mitosis is a gradual and continuous
process, but is usually described in four phases: prophase, metaphase, anaphase and telophase.
In order to understand the stages of mitosis, you need to be familiar with the following concepts:
◗◗ In a non-dividing cell, the DNA wound around protein is known as chromatin (Fig. 3.5).
◗◗ Chromosomes contain linear sequences of genes, the units of heredity that code for the inherited
characteristics of an organism; for example, there are genes on human chromosomes that determine
eye colour, hair colour and height.

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 ◗◗ Each organism has a set number of chromosomes ( for example, humans have 46 chromosomes, a
platypus has 52 and a lettuce has 18).
◗◗ During the S phase, chromosomes replicate and the two chromosome copies or chromatids are held
together by a structure called a centromere. These sister chromatids (Fig. 3.5) separate and move to
opposite poles during mitosis. They are now known as daughter chromosomes and are distributed
into two daughter nuclei. Following this, the cytoplasm divides, separating the two daughter nuclei
from each other (cytokinesis).

FIGURE 3.5
Structure of a
chromosome at the Sister Daughter
start of mitosis chromatids chromosomes
Centromere

Chromosomes

Supercoil within
chromosome
Chromatin fibre

Coiling within
supercoil

Chromatin
DNA
Central Nucleosome
histone
(protein)

Histone
(protein)

DNA DNA double helix (duplex)

Stages of mitosis
At the start of mitosis, DNA is so condensed or highly folded that the chromosomes are visible with a
light microscope.
Although mitosis is a continuous process, it is easier to understand if we divide it into identifiable stages.
There are four main stages of nuclear division in mitosis: following interphase (the S phase during which DNA
replication takes place) are prophase, metaphase, anaphase and telophase. The late stages of nuclear division
are accompanied by the start of cytokinesis. The process of cell division is summarised in Table 3.1. To keep
the diagrams simple, only two pairs of chromosomes (four chromosomes in total) are shown in each cell.

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 TABLE 3.1 Mitosis in an animal cell
STAGE DESCRIPTION DIAGRAM

Interphase occurs in the S phase of the cell cycle, where DNA Cell
Nucleolus
Parent
synthesis occurs: DNA replicates (makes an identical copy of membrane cell

itself ). It appears diffuse and spread out and is termed ‘chromatin’.

Interphase/early

It is not yet recognisable as individual chromosomes.

prophase

In early prophase (seen in this image), DNA begins to separate

into chromosomes.

Nuclear
membrane Centrioles
Chromatid

G2, early prophase: DNA replication complete

In prophase, the chromatin material shortens and thickens by Sister Chromosome splits
longitudinally
coiling and the DNA separates out into chromosomes, which are chromatids

now visible with a light microscope.

Each chromosome contains two copies of the DNA.
Prophase

Each copy is called a sister chromatid and these are joined by

a single centromere.
Nuclear
The nuclear membrane begins to break down and is no longer membrane
breaks down
visible; a spindle begins to form from the broken-down material Spindle fibres
Centromere begin to form
and extends across the cells (like the imaginary lines of longitude
that are drawn on a globe of the world). Prophase: chromosomes condense, become visible and
spindle apparatus forms

In metaphase, the chromosomes line up across the centre or Spindle fibres

‘equator’ of the cell, each attached to the spindle fibres by a

centromere. Each chromosome consists of two identical sister
chromatids.
Metaphase
Mitosis

Centromere Chromatid

Metaphase: chromosomes align along the equator of the

cell

Anaphase begins when proteins in the centromere are cleaved, which Centromere

allows the sister chromatids to separate. Each chromatid becomes a

chromosome. The spindle fibres contract and the chromosomes are
pulled by their centromeres to opposite ends of the cell.
Anaphase

The spindle fibres contract and, as the sister chromatids

begin to separate, they are now known as daughter
chromosomes. They are pulled towards opposite poles of the cell
Spindle
and their movement is assisted by the centromere. fibres Daughter
contract chromosome

Anaphase: sister chromatids separate to opposite poles

of cell

The daughter chromosomes gather at opposite poles of the cell.

The spindle breaks down.
The nuclear membrane and nucleolus reappear.
Nucleolus
Nuclear division or mitosis is now complete. The result is
two nuclei with chromosomes identical to each other and to the
original nucleus in the parent cell.
Daughter
The nuclear membrane forms around the two nuclei, now
Telophase

nuclei
called daughter nuclei. Mitosis is now complete.
Nuclear
membrane
re-forms

Chromosome
Nucleolus

Telophase: nuclear membranes assemble around two nuclei

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 STAGE DESCRIPTION DIAGRAM

Division of the cytoplasm occurs, separating

the two daughter nuclei so that each is in its
own cell.
Cytokinesis differs in plant and animal cells.
• Animal cells: the cytoplasm constricts in the
centre of the cell between the two daughter Cell Nuclei
nuclei, and ‘pinches off’ (similar to squeezing membrane
constricts
a round balloon in the centre until the edges Daughter
cells
meet and then giving it a twist to separate it
into two bubbles).

Cytokinesis in animal cells: division of cytoplasm into

two; G1, cell growth of daughter cells
Cytokinesis

• Plant cells: a cell plate forms while the Nucleus Cell plate Nucleus Chromatin
nucleus is still in telophase; thickenings forms material

appear on the spindle fibres in the region of

the equator and join up to form a cell plate
made of pectin compounds. Cellulose is
then deposited on either side, forming a cell Cell wall
wall to separate the two daughter nuclei.

Cytokinesis in plant cells

Mitosis poster
and video
Download or print
the poster of mitosis, Cytokinesis
Science Photo Library/Steve GSchmeissner

then scroll down and

watch the video clip of Cytokinesis is the final step in cell division (Table 3.1,
live plant cell mitosis
using time lapse
Fig. 3.6). It is the division of the cytoplasm and
photography. begins while the nucleus is completing its division.
Cytokinesis is important because it separates the
newly formed daughter nuclei and ensures that each
cell has only one nucleus. The outcome at the end of
Narrated video – mitosis and cytokinesis is two daughter cells with
mitosis chromosomes that are identical to each other and
Watch the video clip
of mitosis, presented to the original parent cell. The daughter cells then
diagrammatically. enlarge until they are the same size as the original
adult cell (during G1, assimilation as well as cell
enlargement occurs). The nucleus of each cell controls
FIGURE 3.6 Root tip cells undergoing mitosis and all cell activities. The ratio between the proportion
cytokinesis, showing mitotic stages (in order from top
Interactive left to bottom right) interphase, prophase, metaphase,
of nucleus and cytoplasm remains constant. If the
animation – anaphase, telophase and cytokinesis. Chromosomes cytoplasm exceeds a certain proportion of the cell,
mitosis have been stained with toluidine blue.
the ability of the nucleus to control its functioning
decreases and this may help trigger cell division.

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 Telomeres and ageing
As people age, some of the cells in their dividing layers reach Short arm
a point where they can no longer divide. Scientists have
Long arm
discovered that there is a change in the structure of the ends of
the chromosomes in these cells. A telomere is a DNA–protein
Centromere
region at each end of a chromosome that seems to function in
preventing chromosomes from sticking to each other (Fig. 3.7).
Telomeres seem to have additional functions in extending the Chromosome A AU C C C
TTAGGG TTAGGG TTAGGG TTAGGG
life of a cell, such as protecting the genome from degradation
and unnecessary recombination and repair. Children have AATCCC AATCCC AATCCC AATCCC

Telomeres
longer telomeres than older people. With successive divisions,
a small amount of DNA is lost from the telomeres in a cell
and they become shorter. Once the telomeres reach a certain FIGURE 3.7 A telomere is a region at the end of a chromosome that
shortens as we age.
predetermined length, the cell stops dividing and this leads to
cell senescence and/or death. There seems to be variation in
the length of telomeres that people are born with and in the rate at which they shorten. Lifestyle choices
may also influence telomere shortening. This has become an interesting area of research into ageing
and disease.

INVESTIGATION 3.1

A practical and secondary-source investigation of mitosis

INTRODUCTION
Mitosis can be observed relatively easily in cells in the growing region of a root tip. In prepared slides, these
cells have been dyed using a stain (such as acetic orcein or a counterstain with toluidine blue) that is taken up
specifically by chromosomes.
Because the tissue in the prepared slide is no longer living, you will see the cells in the stages of division
that they were in when the root tip was killed with a fixative.
The division of cells at any one time is not synchronised – different cells are in various stages of division. The
phase that is most commonly represented in the slide is the phase that cells remain in for the longest period of
time. Review the cell cycle and predict which phase you would expect this to be.
Using the formula for the mitotic index, you can calculate the proportion of cells in the field of view that are
undergoing mitosis compared with the number not undergoing mitosis (and instead are in the interphase or
growth phase).
The prepared slides that you will look at show stained tissues that are no longer living. Modern advanced
microscopy techniques (not usually available in schools) would allow you to view and record living tissue that is
actively dividing. Videos taken using this technology are available for viewing on the Internet, often on YouTube.
You are encouraged to view video clips taken in real time of cells dividing. Record all the references that you use.

AIM
To gather information on the sequence of changes in the nucleus of plant cells undergoing mitosis

PREDICTION
Review the cell cycle and predict which phase you would expect to see the most number of cells in.

MATERIALS

•• compound light microscope

•• prepared slides of onion root tip

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 RISK ASSESSMENT

! Complete the table to assess the risks and outline how you can safely manage these.
RISK
ASSESSMENT WHAT ARE THE HAZARDS? WHAT RISK DOES THIS HAZARD POSE? HOW CAN YOU SAFELY MANAGE THIS RISK?

Microscope

Glass microscope slides

METHOD
Revise microscope
safety and risk 1 Using correct microscope technique, observe a prepared slide of a root tip using the compound light
assessment from microscope.
Biology in Focus
Year 11. 2 Under low power, locate and identify the apical meristem. This is found just behind the root cap and is
easily identified by numerous small cells that appear square in shape and have nuclei that are large relative
to the whole cell area. Look for cells with darkly stained chromosomes, indicative of mitosis occurring.
Move the slide so that these are in the centre of your field of view and then change to high power to
observe them in detail.
3 Under high power, identify cells in at least four different stages of mitosis. Use figures 3.6 and 3.8 to assist you.

FIGURE 3.8
Cell cycle stages in an onion root tip
A photomicrograph
of onion root tip cells

Science Photo Library/Steve GSchmeissner

undergoing mitosis,
with examples of
the different phases
highlighted
Telophase
Prophase Telophase

Prophase

Metaphase

Metaphase
Cytokinesis
Cytokinesis Anaphase

Interphase

Interphase Anaphase

RESULTS

1 Record your results in the form of fully labelled diagrams. Remember to include a heading for each diagram
and state the magnification used. Write a brief description of what is happening at each stage in the
diagram.
2 You may wish to take photographs of your field of view using your mobile phone. Print the photograph and
label on it the various stages of cell division that you can see.
3 Calculate the mitotic index of your field of view (refer to Introduction) using the formula:
number of actively dividing cells in field of view
Numeracy Mitotic index =
total number of cells in field of view
4 Write a practical report using a procedure text type, under the headings Aim, Risk assessment, Materials,
Method, Results and Conclusion.

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 DISCUSSION
Discuss the accuracy, reliability and validity of data you gathered during the investigation. Estimate the
uncertainty of measurements in your data. (See Chapter 1, page 20 for estimating uncertainty.)

CONCLUSION
Write a summary statement that relates to the aim and your prediction in this investigation, including a brief
outline of what you found.

EX TENSION
Carry out one of the following optional extension activities.
1 Find a video clip of mitosis filmed under a light microscope or fluorescence microscope in real time. Record
the weblink so that your peers can view it.
2 Estimate the duration of each stage of mitosis by recording data on the observed frequencies of each stage
of cell division in your field of view. Present your data in table format, using Table 3.2 as a guide.

TABLE 3.2 Observed frequencies of stages of cell division in a root tip field of view

OBSERVED OCCURRENCE IN REPRESENTATIVE

MICROSCOPE FIELDS DURATION Mitosis
animation and
% OF TOTAL ESTIMATED TIME quiz
STAGE OF CELL DIVISION NUMBER OF CELLS % OF TOTAL CELLS TIME (MINUTES)
Work through the
quiz to check your
Interphase understanding.

Prophase

Metaphase

Anaphase

Telophase

Cytokinesis

Replication of DNA outside the nucleus

Although each cell has one nucleus, it will have many organelles such as mitochondria and chloroplasts
in the cytoplasm. The number of each type of organelle depends on the functions of the tissue. Cells
that require large amounts of energy have many millions of mitochondria. Photosynthesising cells in the
leaves of plants have many chloroplasts. Just as cells divide, these organelles must also divide to maintain
their numbers for normal tissue functioning.
A small amount of DNA is located in organelles such as mitochondria and chloroplasts, in the
cytoplasm. This DNA, called non-nuclear DNA or extrachromosomal DNA, carries genes that are
important to cell metabolism.
In cytokinesis, when the cytoplasm divides in half, organelles in the cytoplasm such as mitochondria
and chloroplasts reduce in number per cell, as they are now distributed between two daughter cells. To
avoid a repeated reduction in quantity with each cell division, mitochondria and chloroplasts need to be
able to replicate themselves. These organelles contain their own small amounts of DNA and replicate You will learn
independently of the nucleus, in this way maintaining their numbers in successive generations of cells. By more about the
structure of
the time the daughter cells have grown to the size of the original cell, organelle replication has occurred mtDNA, maternal
to restore the organelle number to that of the original cell. Assimilation is important in the growth of inheritance and
applications
many organelles, such as the endoplasmic reticulum. of mtDNA
The DNA in mitochondria is referred to as mtDNA (mitochondrial DNA). Molecules of mtDNA are sequencing in
Chapter 4.
much shorter than nuclear DNA molecules and are arranged in a single circle.
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 In gamete production in humans, all mitochondria inherited by offspring are located in the cytoplasm
of the egg cell. Sperm cells are tiny and have very little cytoplasm, because they are made up predominantly
of a nucleus and a tail for swimming. Therefore children inherit all their mtDNA from their mother. No
mtDNA is inherited from the father. Therefore, studies of mtDNA reflect maternal inheritance over many
generations.
KEY CONCEPTS

●● There are two types of nuclear division – mitosis and meiosis.

●● The repeating sequence of cell growth and division that a cell passes through is called the
cell cycle, with phases G1, S, G2 and M.
●● Specialised cells are differentiated and no longer go through the cell cycle.
●● Mitosis has five phases: interphase, prophase, metaphase, anaphase and telophase.
●● Interphase is G1, S and G2 of the cell cycle. DNA replication takes place during the S phase.
●● In metaphase, sister chromatids held together by a centromere line up on the equator of the
cell.
●● In anaphase, the sister chromatids separate from each other and move towards opposite poles,
drawn by contraction of the spindle fibres.
●● Cytokinesis is division of the cytoplasm and begins as nuclear division is ending.
●● Cells that form by mitosis have the same genetic material as their parent cell and each other.
●● Organelles in the cytoplasm that have their own DNA, such as mitochondria and chloroplasts,
replicate independently of the nucleus.
●● Telomeres decrease in length with each division, and short telomeres are a sign of ageing.

CHECK YOUR
UNDERSTANDING 1 Define each of the terms below and draw a fully labelled diagram to distinguish between them:
chromatin, chromosome, chromatid, centromere.
3.1 2 Explain the difference between a chromatid and a chromosome.
3 Distinguish between cell division and mitosis.
4 a Describe the main difference between unicellular organisms and multicellular organisms with respect
to the role of mitotic cell division.
b State two other important roles of mitosis in multicellular organisms.
5 Compare cytokinesis in plant cells with that in animal cells.
a Draw a fully labelled comparative diagram, to compare early prophase with metaphase in a cell that is
dividing by mitosis.
b Explain why more cells are found in interphase than in any other phase of mitosis in the field of view of
the dividing region of a root tip cell.

DNA structure – the Watson

3.2 and Crick model
Today we know that cells carry their biological information in a molecule called DNA (deoxyribonucleic
acid). However, the actual chemical nature and chemical structure of the hereditary material and
genes was a mystery until the middle of the 20th century.

A brief history of the discovery of DNA

In 1869, Swiss scientist Friedrich Miescher discovered a chemical he called ‘nuclein’, and showed that
this distinct chemical was commonly found in the nuclei of cells in a variety of tissues. His ‘nuclein’
proved to be made up of DNA and associated proteins from cell nuclei. Miescher is therefore credited
with being the first scientist to identify DNA as a distinct molecule.

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 By 1910, chromosomes had been discovered. At that stage, scientists knew that they were made up
of a mixture of DNA and protein and that they carried the units of heredity called genes (named in 1909
by William Johannsen).
Over the next 30 years or so, the common expectation among scientists was that the secret of
heredity would be found in proteins, because their building blocks were more varied – twenty amino
acids, compared with DNA’s four bases. However, by the mid-20th century, a number of scientists were
moving towards the idea that DNA carried the genetic information.
In 1944, Oswald Avery and his team, working on bacteria at the Rockefeller University hospital in New
York, first provided experimental proof that DNA, not protein, is the hereditary material.
Many biologists were not convinced, as they were suspicious of Avery’s methodology and thought he
may have obtained his results using DNA that had been ‘contaminated’ by protein. The race was on in
laboratories around the world to try to solve the puzzle – the structures of both protein and DNA were
being investigated.
Two leading teams in England were working on the molecular structure of biological molecules at
the time:
◗◗ James Watson and Frances Crick worked in the Cavendish laboratory at Cambridge University, under WS

the leadership of Lawrence Bragg. TheHomework

discovery
◗◗ Maurice Wilkins and Rosalind Franklin worked at King’s College in London, under the leadership of of DNA
John Randall.
By 1951, highly competitive teams of scientists in these British laboratories were doing similar
research, as was the American laboratory of Linus Pauling. There was a longstanding rivalry between
Lawrence Bragg and Linus Pauling (who had won a Nobel prize for his work on chemical bonds). Early in
1953, Linus Pauling proposed that DNA was the heredity molecule and that its structure was a triple helix.
Watson and Crick used chemical and X-ray evidence to construct a model of the molecular structure
of DNA (Fig. 3.9). The model revealed a stable molecule that could store large amounts of information. It
was a two-stranded molecule, with paired bases twisted into a spiral ladder. Because of its structure, they
called the DNA molecule a ‘double helix’ (and the name of the book that Watson later wrote, describing
their journey of discovery, was The Double Helix). An interesting and critical feature of their model was
that it could be used to explain how the molecule could self-replicate.
Science Photo Library/A. Barrington Brown, Gonville and Caius College

a Alamy Stock Photo/Science History Images

b

FIGURE 3.9 a James Watson (left) and Francis Crick (right) with their model of DNA structure (1953); b James Watson in later
years, with some of the many books he wrote about DNA and science

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 Watson and Crick’s model of DNA showed a large linear molecule arranged as a double helix or
‘twisted ladder’, with four nitrogenous bases that were held in pairs by hydrogen bonds. Their model
suggested that genetic information could be stored in the order or sequence of the bases that make up
the middle of the DNA structure. On 28 February 1953, Watson and Crick won the race, and announced
that afternoon, at a pub called The Eagle, that they had discovered ‘the secret of life’.

Alamy Stock Photo/Martin Parker

a

Getty Images/Chris Radburn-PA Images

b

FIGURE 3.10 a The Eagle pub in Cambridge, England, and b the plaque outside the pub to commemorate where Crick and Watson celebrated
their discovery

Watson and Crick had worked collaboratively, evaluating the processes, claims and conclusions
of other scientists. They had asked questions of colleagues such as Maurice Wilkins who, in response,
had shown them Rosalind Franklin’s X-ray crystallography photograph of DNA (without first asking her
TED talk: James
Watson speaking permission) to Watson when he visited the laboratory. Watson and Crick applied critical and creative
about how he
discovered DNA
thinking, both in predicting what to expect and in analysing the primary and secondary chemistry and
crystallography data they had gathered.
They considered the quality of all available evidence from a variety of scientists and used reasoning to
construct their scientific predictions and arguments, subjecting the model they were building to rigorous
testing and validation. As a result of their innovation, their attention to sound scientific detail and their
Discovery of the
structure of DNA collaborative approach, they came to the valid conclusion that won them a Nobel Prize in 1962. Avery
The full story of the and his team may also have been in line for a Nobel Prize but, like Rosalind Franklin, Avery died before
scientific advances
preceding Watson this prize could be awarded to him.
and Crick’s insightful
discovery

DNA structure – the Watson and Crick model

Crick realised that each DNA strand, comprising a sugar-phosphate backbone, is antiparallel to the
other, meaning that the twisting strands run parallel to each other but in opposite directions. X-ray
crystallography photographs of the DNA structure suggested a set distance between the ‘backbones’
of the molecule. Thus, each base could have one end attached to the backbone and the other end
hydrogen-bonded to another base. So the bases lined up between the backbones, like the rungs of a
ladder (Fig 3.11).

Nucleotide pairing and bonding

Watson and Crick realised that in their DNA model, to ensure the DNA backbones remain equidistant
from each other, a double-ringed purine base needs to bond with a single-ringed pyrimidine base. The
purine base adenine must therefore bond with the pyrimidine base thymine (A and T), and the purine
base guanine must bond with the pyrimidine base cytosine (G and C).

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 a b

Adapted from Pray, L. (2008) Discovery of DNA structure and function: Watson and Crick.
Nature Education 1(1):100 (Figure 4: Base pairing in DNA)
39 end 59 end
H
H N H O N O P O2
C C C C HC Sugar-phosphate
O O
39 H CH C N H N G C N O 59 backbone
H N C C N H2C
H
H H O H N H H
59 C H2 H H
O H 39
CH3 H
O O
2O
P O H O O P O2
N H C C G C
O N HC O
39 H CH C C H N T N O 59 C H A T
N C A N C 2
H H G C
H H N C O H H
H H
59 C H2 H 39 G C
O H
H H O
O
2O P O H N O P O2 C G
O C C HC
O N O T A
39 H CH C C N C N 59
H C O H2C C G
H H N C G N
H H N C O H H A T
N H H H
59 C H2 O 39 G C
H H
O H O T A
CH3 H N N
2O
P O O HC O P O2 C G
C C C C
O CH O
H T N H N A C N
39 C O 59 C
H N C N H2 G C
H
H H H H H
59 C H2 O O H H A T
Hydrogen bond 39
H G C
O O
2O P O
39 end Base pairs
59 end

FIGURE 3.11 The chemical structure of DNA, showing its antiparallel backbone and base pairing: a the molecular structure of DNA showing the
sugar-phosphate bonding and hydrogen bonding of base pairs; b a simplified version of DNA

From the chemistry of this pairing, they also predicted that the DNA strands would be held together
by weak hydrogen bonds between the paired bases: a double bond between A and T, and a triple bond
between G and C (Fig. 3.12). Watson and Crick found evidence for this pairing, as it fitted with Chargaff ’s
‘parity rule’. In 1951, Austrian biochemist Erwin Chargaff discovered that, in DNA, the ratio of the bases
A : T and C : G was always 1 : 1.
Watson and Crick realised that this complementary base pairing could also be used to predict how
DNA could be copied. They published their research paper, titled, ‘Molecular structure of nucleic acids’, in
1953. In this paper they stated: ‘It has not escaped our notice that the specific pairing we have postulated
immediately suggests a possible copying mechanism for the genetic material’.

DNA serves as a template for replication DNA double

helix
Watson and Crick’s DNA model showed all the requirements expected of hereditary material.
◗◗ DNA can carry, in coded form, all the instructions for the formation and functioning of cells, despite
the fact that its ‘alphabet’ consists of only four nitrogenous bases.
◗◗ The structure of DNA allows for its own replication. To copy DNA, each strand can serve as a template
for enzymes to synthesise a new complementary DNA strand.

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 ◗◗ DNA can be transferred from one generation to the next, packaged in the form of chromosomes and
carried by gametes. It was discovered that proteins associated with DNA in chromosomes have the
The double helix
role of keeping DNA ‘neatly packaged’. DNA is coiled around these proteins (called histones), in much
Watch this 16-minute the same way that cotton thread is wound around a cotton reel. This holds the long threads of DNA
film about the in a compact way so that DNA is easy to sort and separate during cell division and can be efficiently
discovery of DNA.
transported from one generation to the next. Histones are also thought to play a role in exposing
sections of DNA, so that genes can be expressed.

The vertical sides of the ladder are

made up of alternating sugar and
phosphate molecules (a
sugar-phosphate backbone) and
the ‘rungs’ of the ladder are pairs of
nitrogenous bases (adenine, thymine,
guanine and cytosine or A, T, C and
G).

Sugar-phosphate ‘backbone’

Each nucleotide is made up of a

Hydrogen bonds between O
phosphate group, a sugar
P
nitrogenous bases (deoxyribose sugar), and a
O
P nitrogenous base attached to the
G
sugar.
O C
P
C O

O G P

P A
O
O There are four types of nitrogenous
T
P bases: adenine, guanine, cytosine
P G and thymine.

O
O
C
P
OH T
3 end
Chemically, bases pair in a particular
O manner: adenine with thymine and
The DNA molecule is a long-chain A guanine with cytosine (i.e. A–T and
molecule consisting of two G–C). They are held together in the
complementary strands. Each P centre by hydrogen bonds, forming
strand is made up of a sequence of Phosphodiester two complementary strands.
many nucleotides and the strands are bond
held together by weak hydrogen O
bonds in the centre. The two strands
in the double helix model have an
‘antiparallel’ arrangement – they run
in opposite directions. P

5 end
FIGURE 3.12 An annotated diagram of the DNA double helix molecule, showing sugar-phosphate backbone and base pairing

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 TABLE 3.3 Validation of the model: Watson and Crick’s findings, and the evidence on which they were based

FINDING EVIDENCE-BASED REASONING

The whole DNA ‘ladder’ molecule, instead of being Evidence: X-ray crystallography suggested a helix measuring 3.4 nm for every turn,
flat, spirals and is therefore known as the ‘double and this fitted the model where exactly 10 base pairs would measure 3.4 nm in
helix’. length and make up one twist of the helix.

The four types of nitrogenous bases: adenine (A), Evidence: Chargaff’s rule stated that DNA from any cell has a 1:1 ratio of pyrimidine
guanine (G), cytosine (C) and thymine (T) always pair and purine bases and, more specifically, that the amount of guanine is equal to
as A–T and C–G. cytosine and the amount of adenine is equal to thymine. Watson and Crick found
complementary base pairing of pyrimidine–purine (that is, A–T and G–C) was
critical in keeping the backbones of DNA equidistant.

The two complementary strands of DNA are held Reasoning: Chemically, when A bonds to T, a double hydrogen bond may form and
together in the centre by hydrogen bonds that form when G bonds to C, a triple hydrogen bond may form.
between the complementary bases.

The strands of the backbone must be identical and Evidence: DNA crystal images, generated by Rosalind Franklin, looked the same
run in opposite directions (antiparallel). when they were turned upside down or backwards.
Reasoning: The backbones are made of sugar-phosphate molecules, based on the
ratio of these in the chemical analyses.

Each DNA strand serves as a template for the Reasoning: The two complementary strands of DNA could ‘unzip’ or open up if the
production of a complementary strand, allowing the hydrogen bonds break between the base pairs, allowing them to replicate.
double-stranded molecule to self-replicate.

Watson and Crick could form hypotheses about DNA’s structure by building physical models of how the atoms fit together. Sometimes
they used cardboard cut-outs to represent the four nitrogenous bases and other subunits of DNA. They played with these on a desk,
like a jigsaw puzzle. They initially made an error in the configuration of the rings in thymine and guanine. By changing the arrangement
of atoms and making new cut-outs, based on a suggestion from Jerry Donahue, an American scientist, they found the perfect fit:
complementary base pairing, ratios to reflect Chargaff’s rule and the hydrogen bonding of purines to pyrimidine bases.

Nucleic acids as we know them today

There are two types of nucleic acids – DNA (deoxyribonucleic acid) and RNA (ribonucleic acid). Nucleic
acids contain the chemical elements carbon, hydrogen, oxygen, nitrogen and phosphorous. WS
All nucleic acids are polymers made up of simple repeating units (monomers) called nucleotides.
Chromosomes
Homework –
Nucleotides may be linked together to form single chains, as in RNA, or double strands, as in DNA the key to
inheritance
(Fig. 3.13). These DNA strands may be very long. For example, human chromosome 1 contains 249 million
base pairs of DNA, representing approximately 8 per cent of the total DNA content of a human cell.
Each nucleotide is made up of three parts: a simple sugar (ribose in RNA, deoxyribose in DNA), a
phosphate and a nitrogenous base.
The genetic code is created in the consecutive sequence of bases. These base sequences differ in each
gene, providing the ‘genetic code’ for a cell.
Revise the
structure of
Comparing the roles of DNA and RNA in cells nucleic acids from
Biology in Focus
◗◗ DNA stores the genetic information that controls the cell and thereby the whole organism. Year 11, Chapter 3
◗◗ DNA is the main chemical making up chromatin in the nucleus, although small amounts of DNA are
also found in mitochondria and chloroplasts.
◗◗ DNA is responsible for transmitting inherited information from one cell to another during cell division
and from one generation to another during reproduction. WS
◗◗ RNA is a nucleic acid found in small amounts in the nucleus, and in larger amounts in the cytoplasm;
The Homework
structure of
it is usually associated with ribosomes. nucleic acids
◗◗ RNA has the base uracil (U) instead of thymine (T).

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 The three types ◗◗ There are three types of RNA:
of RNA and their – messenger RNA (mRNA), which is involved in carrying information from DNA, transporting a
functions are dealt
with in more detail transcribed copy from the nucleus to the cytoplasm
in Chapter 4. See – ribosomal RNA (rRNA), which is associated with proteins in ribosomes, bringing mRNA and
Fig. 4.12.
tRNA together during translation
DNA instructing
the formation of
– transfer RNA (tRNA), which assists in ‘translating’ the mRNA message into proteins. (You will
proteins is also dealt learn more about this in the next section.)
with in Chapter 4.

P
a b Coiled DNA
S B
P
P
S B P Nucleus
S
P S B B
P Chromosome
S B P

B B
S
P S
P
S B P
S B P DNA double helix
Nucleotide S
P S B B
P
S B P Guanine The four bases
S
P S B B Thymine
P Adenine
S B P Cytosine
39 59
P DNA
S B Sugar–phosphate
backbone of DNA
RNA

FIGURE 3.13 a The nucleotide monomer is made up of base–sugar–phosphate, a single-stranded RNA polymer molecule or polynucleotide, and
a double-stranded DNA molecule (a polymer of nucleotides). b The four bases and sugar-phosphate backbone make up the DNA, which forms a
chromosome. Here the chromosome is composed of two identical (sister) chromatids, held together by the centromere.

DNA instructs the formation of proteins

See page 125 for the
In 1955, Crick proposed that the order of the four bases (A, T, G, C) on the DNA molecule determines the
genetic codes for sequence of amino acids in a protein. Their ‘sequence hypothesis’ could explain how the order of bases
amino acids.
in genes along the chromosome can be translated into the language of proteins. By 1966, scientists had
cracked the genetic code and could show exactly which three-base sequences code for each amino acid.
KEY CONCEPTS

●● Each nucleotide consists of three parts – a phosphate, a sugar (deoxyribose sugar in DNA, ribose
sugar in RNA) and a nitrogenous base.
●● There are four types of nitrogenous bases and each nucleotide is named after the base that it
carries – adenine, thymine, guanine or cytosine nucleotides. These are often simply referred to
by their first letters – A, T, G and C.
●● A and G are the larger double-ringed purine bases, and C and T are the smaller single-ringed
pyrimidine bases.
●● Each DNA molecule is made up of two chains or strands that have an antiparallel arrangement
– that is, they are parallel but run in opposite directions.
●● Each strand is made up of a sequence of many nucleotides, and the strands are held together by
weak hydrogen bonds between the bases in the centre of the DNA molecule.
●● The advantage of these weak hydrogen bonds is that little effort is required to pull the bases
apart so that DNA can replicate or be decoded to form proteins.

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 ●● The vertical sides of the ladder are made up of alternating sugar and phosphate molecules (a
sugar-phosphate backbone) and the ‘rungs’ of the ladder are pairs of a purine base linked to a
pyrimidine base: adenine-thymine (A-T) and guanine-cytosine (G-C). The bases are attached to
the backbone through the sugar component.
●● The bases are arranged in a sequence along each strand of DNA (e.g. GGTCAGGCTTGAACGA)
and the length of a DNA molecule is presented as the number of base pairs (bp).
●● The two strands of DNA unzip (weak hydrogen bonds break) for replication and gene
expression.
●● RNA is a single-stranded molecule made of nucleotides, with a ribose sugar attached to each
phosphate to form the backbone; on the other end it attaches to a base, either A, U, C or G. RNA
does not have thymine, which is replaced by the base, uracil (U).

CHECK YOUR
1 Create a timeline to summarise the contributions made by scientists in the discovery of the hereditary
UNDERSTANDING
material in cells. List the scientists in the chronological order of their discoveries and summarise their
findings under headings such as: Date, Name(s), Laboratory/Place of work, Contribution (theory/law/ 3.2
hypothesis), and a brief description of findings in their research.
2 Describe the main features of DNA that Watson and Crick discovered, under the headings: Nucleotide
composition, Base pairing and Bonding.
3 Compare, using examples, the structure of a purine with a pyrimidine base.
4 Explain, in terms of the structure of DNA, why a purine had to combine with a pyrimidine to create a
double helix.
5 What is meant by the term ‘antiparallel’ in describing the backbones of DNA?
6 Outline the three main requirements of hereditary material for it to be able to carry out its functions.
7 How does the structure of DNA support the idea that it can self-replicate?
8 Compare the structure and functions of DNA and RNA.
9 Evaluate the benefits and limitations of the two DNA models shown in Fig. 3.11.

DNA replication – the Watson

3.3 and Crick model
DNA replication is the production of two identical double-stranded molecules of DNA from one original
double helix molecule. DNA replicates during interphase prior to mitosis, and each cell receives one
exact copy of the coded instructions that control the basic life functions of the cell.
Watson and Crick noted that their model suggested a possible copying mechanism for the genetic
material. If the strands separated, each strand could act as a template for the synthesis of a new,
complementary strand. This is indeed what happens. This process of DNA replication in cells is termed
‘semi-conservative’, as each resulting double-stranded DNA molecule has one new strand and one ‘old’ or
conserved strand. This mechanism should ensure that the genetic material is copied exactly every time.
However, DNA replication is not quite as simple as it appears. The following features add complexity
to the process:
◗◗ DNA has to be unwound from its spiral configuration before the strands can be separated.
◗◗ A large number of physical and chemical reactions take place simultaneously during DNA replication,
and so a collection of enzymes is required to control each reaction.
◗◗ The two strands of DNA run in opposite directions (antiparallel) and nucleotides can be added in one
direction only (onto the free 3′ end).
◗◗ DNA replication errors sometimes occur and these need to be corrected.

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 The process of DNA replication
DNA replication occurs in four main steps.
1 The DNA double helix unwinds.
Each DNA molecule is a double-stranded helix. An enzyme called helicase causes the DNA helix to
progressively unwind and the strands to separate (Fig 3.14a).

WS P P
P P P P Weak hydrogen bonds
DNA replication S G C S S G C S break, catalysed by
Homework S G C S
DNA helicase.
P P P P P P
S S C G S S C G S
P C G P P P P P
DNA is
S S unwound S T A S S T A S
P T A P by helicase. P P P P
1 2
S S S C G S S C G S
C G DNA polymerase
P P P P P P adds nucleotides
S S T A S S T A S to make new
DNA strands.
P T A P P P P P
S S S G C S S G C S
P P P P S
P C G P G
P
S
Free P
a The DNA double helix unwinds. C nucleotides
A S S

G
3 P A P
S
P
b DNA unzips and the two strands separate.

P P P P P P P P
S G C S S G C S G C G C
S S S S
Hydrogen
P P P P P P P P
bonds form
S C G S S C G S S S
P P P P P P P P
G C G C
S T A S S T A S S S S S
P P P P A T A T
P P P P
4
S C G S S C G S S G C S S G C S
P P P P P P P P
S T A S S T A S S S
P P P P P P P P
T A T A
S G C S S G C S S S S S
P P P P P G C P G C
P P

Original Newly Original

strand synthesised strand
strands

c Nucleotides are added alongside both strands, d The resulting DNA molecules each contain half
opposite their complementary bases, to create a new of the original DNA molecule and a newly
strand along each original strand. synthesised strand (semi-conservative replication).

FIGURE 3.14 DNA replication – a simplified diagrammatic sequence

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 2 DNA unzips – the two strands separate.
Complementary Original
Using ATP as the energy source, helicase disrupts the weak hydrogen chains of DNA DNA helix
bonds between the complementary bases of the nucleotides on
opposing strands. The DNA strands separate, exposing the nucleotide
bases (Fig. 3.14b). If you think of DNA as a ‘ladder’, each ‘rung’ splits
down the middle, starting at the bottom of the long DNA molecule Replication
fork
and creating a replication fork further up, where the DNA is still
Pool of
joined (Fig 3.15). Single-stranded binding proteins (SSBs) bind to and nucleotides
stabilise the newly separated single-stranded DNA.
3 Nucleotides are added against each single strand. Building
units
Each separate strand of the DNA molecule acts as a template for the DNA
T A T A
DNA
G C G C
production of a new strand of DNA. For synthesis to be initiated, a polymerase polymerase
short strand of RNA needs to be made and attaches to the DNA – this
is known as a primer. RNA primers are made by the enzyme primase.
Okazaki fragments
Next, the enzyme DNA polymerase III adds DNA nucleotides, Template joined on this
to continue the synthesis of the new strand. It picks up free strand by ligase
nucleotide units (made of sugar–phosphate–base) floating in the
liquid background of the nucleus and inserts them opposite their New
complementary base partner, using the existing strands as templates chain
and keeping the nucleotide pairing specific:
Leading Lagging strand
Adenine (A) always pairs with thymine (T) nucleotides. strand
5' end
3' end
Guanine (G) always pairs with cytosine (C) nucleotides. Original
complementary strands
Note: The two strands in a double-stranded DNA molecule run
in opposite directions – they are antiparallel. Each DNA strand has a FIGURE 3.15 DNA replication showing a replication fork,
three prime (3′) end and a five prime (5′) end. Nucleotides are always free nucleotides and DNA polymerase synthesising the
new complementary strands (drawn in yellow)
added from the three prime (3′) end of a DNA strand. Therefore
nucleotide insertion during replication is also antiparallel along the
two template strands.
On one strand of DNA, nucleotides are added in a long chain, growing in the same direction as the
replication fork opens up. This is called the leading strand and replication is continuous. Base mismatch
On the other strand (the lagging strand), nucleotides are added in ‘chunks’ (called Okazaki pairs are dealt
with in more
fragments), from the replication fork backwards (Fig. 3.15). Replication in this strand is therefore detail on page
discontinuous and the fragments are then joined up by an enzyme called ligase, to form one 104 (Fig. 3.18).

continuous strand.
4 Replication errors are identified and corrected.
DNA polymerase I is a complex enzyme that can backtrack to ‘proof read’ and ‘edit’ the strand. It
corrects any base pair errors by splicing out the incorrect base that was inserted into the new strand Interactive
explanation of
and replacing it with the correct base. DNA replication
Click on terms in
Finally, the two new strands are sealed together by the enzyme called ligase. The final base pairing the explanation
is checked by another DNA polymerase enzyme, which recognises base pairing errors and carries out of DNA replication
for simplified
base mismatch repairs, to ensure accuracy. meanings.

Despite these checking mechanisms, a small number of errors in DNA replication still occur –
about one in every ten billion base pairs. Incorrect base pairing results in a change in the DNA base
sequence, known as a mutation.
Cells may delay the progression of the cell cycle until DNA repair (during the G2 phase) is complete.
DNA replication
Each resulting DNA molecule contains one strand of the existing DNA molecule and a newly simulations
and other
synthesised strand. The replicated DNA molecules rewind into the double helix conformation, like interactives

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 the original molecule. The end result is that there are two molecules of DNA, each a double-stranded
helix, identical to each other and to the original molecule from which they formed.

Enzymes ensure exact replication of DNA

Replication of DNA is a complex process and each step is controlled by one or more enzymes (Table 3.4,
Fig. 3.16). Most spontaneous changes in DNA are temporary because they are immediately corrected by
repair enzymes.

TABLE 3.4 Enzymes that regulate DNA replication

ENZYME FUNCTION IN DNA REPLICATION

Topoisomerase
• Relaxes DNA from its supercoiled state, always working ahead of the replication fork
(e.g. gyrase)

Helicase • Follows topoisomerase and unwinds the double helix by breaking hydrogen bonds
between bases, causing the two strands to separate and creating a replication fork

Primase • Connects RNA primer to a strand to initiate DNA replication

• Synthesises a short complementary RNA molecule, which binds to DNA, serving as the
starting point for DNA synthesis by polymerase
DNA polymerase III • Synthesises new DNA strands, using existing strands as a template. Nucleotides are
added to a growing strand from the 3′ end
• Joins the phosphate group of each nucleotide to the previous one, creating
phosphodiester bonds between these molecules to make the sugar-phosphate backbone

DNA polymerase I • Mainly functions in ‘editing’ – recognises and repairs base pairing errors (exonuclease)
• Also has a function in replication (removing primers ahead of the main polymerising enzyme)

DNA polymerase II • Editing function, but no exonuclease activity

Ligase • Connects and seals the two strands of the DNA molecule and also connects Okazaki
fragments

• On the leading strand – DNA moves along in the same direction as the developing replication fork, and
nucleotides are added in one long chain.
• On the lagging strand, DNA is added one ‘chunk’ at a time – called Okazaki fragments (about 100–150
nucleotides long) and these are then joined up.

FIGURE 3.16
Enzymes involved in
DNA replication

Helicase unwinds Binding proteins stabilise Primase adds a short

parental double helix. the strands. primer to template strand.

DNA polymerase binds Exonuclease removes Ligase joins Okazaki

nucleotides to form RNA primer and inserts fragments and seals nicks in
new strands. correct bases. sugar-phosphate backbone.

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 INVESTIGATION 3.2

Modelling DNA replication using the Watson and Crick model

BACKGROUND: USING SCIENTIFIC MODELS AND WRITING A HYPOTHESIS
Following their landmark paper on DNA structure, in 1954 Watson and Crick proposed a mechanism by which Information and
communication
DNA could be copied. This mechanism was based on their DNA model in that each DNA strand could serve as technology
a template for the synthesis of a new strand. In their own words, ‘prior to duplication, the hydrogen bonds are capability
broken, and the two chains unwind and separate. Each chain then serves as a template for the formation … of Critical and
two pairs of chains, where we only had one before. Moreover, the sequence of the pairs of bases will have been creative
thinking
duplicated exactly.’ So the final result is two identical double helix DNA molecules (sister chromatids), each
made up of one old and one newly synthesised strand – a model of semi-conservative replication.
© 1953 Nature Publishing Group. Watson, J. D., and Crick, F. H. C., Nature volume 171, pages 964–967 (1953). doi:10.1038/171964b0. All rights reserved

PREDICTIONS AND VALIDATIONS OF THE WATSON AND CRICK MODEL OF SEMI-CONSERVATIVE REPLICATION
The semi-conservative model that Watson and Crick proposed was tested and validated by scientists using the
DNA of both prokaryotes and eukaryotes. Hypotheses
used to test
Scientists realised that, to test the Watson-Crick proposal for DNA replication, they needed to find a way to DNA replication
distinguish between the old and the newly synthesised strands. proposal
In 1958, Matthew Meselson and Franklin Stahl carried out experiments that supported and verified Watson Investigative
methods used to
and Crick’s proposal of semi-conservative replication of DNA. They used heavy isotopes of nitrogen, which were test the hypothesis
incorporated into the nitrogenous bases that would be added to the newly synthesised strand. A centrifuging that DNA
replication is semi-
technique was then used to determine the density of the DNA strands, determined by the buoyancy of DNA. conservative
The results of the buoyancy tests supported predictions made for semi-conservative replication.

VALIDATING SCIENTIFIC MODELS

Scientific models are developed to represent an idea, object, process or system that cannot be observed directly.
Models are simplified representations; they do not try to explain every detail in the ‘real world’ process, object or Writing
a formal
idea. Scientists are usually aware of the limitations of the models that they propose and have critically analysed. The hypothesis (self-
models that are most valuable in science are those that allow us to make and test predictions. If a prediction is made assessment)
and the model holds true, the prediction is said to validate the model. Sometimes, as new evidence is found, models
need to change to reflect a more accurate understanding of an idea or process. If a model cannot hold true in the
light of new evidence, the model may be rejected. Models are therefore indicative of the tentative nature of science.

AIM Cut and paste

DNA
To model DNA replication, showing semi-conservative replication
In this investigation, you are required to generate a model to represent the structure of DNA and the process
of DNA replication. Because each model created will demonstrate only some aspects of the structure and
process, the models built by students will all be different. You will be required to peer assess the models that
you view. You will need to identify the structural and conceptual differences between the models and use your DNA with
confectionary
critical analysis skills to discuss the advantages and limitations of each model.

METHOD

1 Working in groups of three to four, build a working model of DNA structure, showing the double helical
nature of the molecule. You need to build a working model with features and spare parts that will be used
Jewellery DNA
by another group to demonstrate how your DNA can act as a template to demonstrate semi-conservative
replication. Your teacher may assign one or two specific features (a to e below) that your model needs to
show, or each group may be allowed to select which features they wish to show.
2 Using the Internet, carry out research to review different types of DNA models that may be created to show
structure and/or replication. You will discover that models may be created from a wide variety of materials, Digital DNA
from edible models using lollies, to models that use materials such as pipe cleaners and beads, or easily
available recyclable materials such as cardboard and toothpicks, or waste materials such as styrofoam and
plastic. See the online resources linked to this page for ideas. You will also need to analyse models of DNA

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 replication to determine whether they can be used to demonstrate the feature of replication you have
selected or been assigned.
3 Your model of semi-conservative replication of DNA should demonstrate some or all of the following:
a complementary pairing of bases
b creation of Okazaki fragments
c unwinding of DNA with helicase action, creating a replication fork
d addition of nucleotides by DNA polymerase, using an old strand as a template and creating a new strand
e mutation during DNA replication.
4 Label the structures in the DNA model. Labels should include all the features listed in Table 3.5. In addition
to these labels, also label any additional features, such as enzymes or assigned features.
Common labels: sugar-phosphate backbone, nucleotide, nitrogenous base, hydrogen bond, 3′ end, 5′ end.

PREPARING YOUR MODEL FOR VALIDATION

5 Create the following to be placed with your model in the next lesson:
a instructions to your classmates on how to replicate your DNA molecule
b raw materials, so that another group can carry out the replication process on your model
c a copy of Table 3.5, completed to describe the features of your model
d a printout of a photograph of your model (before it is replicated).

TABLE 3.5 Key to DNA model

FEATURE MATERIAL COLOUR

Sugar-phosphate backbone

Nitrogenous bases

Hydrogen bonds

One nucleotide

Feature to be demonstrated during replication

Limitations

GROUP ROTATION ACTIVIT Y: VALIDATING MODELS

6 Set up your model and provide spare parts and clear instructions for the next group on how to replicate your
DNA molecule. Copy and complete Table 3.5 to accompany your model as a key, identifying what materials
have been used to represent each feature. You also need to provide a printout of a photo of your model.
7 Rotate groups: A → B; B → C; C → D; D → E; E → A.
a Identify the parts of the DNA model you have rotated to, using the key provided. Take a photo of the
model before you replicate it.
b Use the spare parts provided to carry out replication, showing the main feature listed at the bottom of
the table. You may use your textbook to revise exactly what this feature is. Take a photo of the replicated
model. Complete a peer assessment of the model, listing its advantages and limitations.
8 Move to the next group and take on the proof reading and editing role of DNA polymerase. Check
whether the model has any errors of replication (mutations). Take a photo of the replicated model. Turn the
explanatory table over (or cover it with a piece of paper) so that the next group cannot see it.
9 Move to the next group and complete a blank copy of Table 3.5 for the model you are viewing. Do not
cheat by turning over the key. Take a photo of the model and your completed table.
10 Move to the last model. Compare it with the model that your group made, writing down two similarities
DNA replication and one difference between this model and your model. Take a photo of the model.
Build strands of
DNA by inserting EX TENSION
complementary bases
DNA replication – try the DNA replication simulation in class or complete it for homework. (See the weblink.)

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 KEY CONCEPTS
●● The two DNA strands unzip for replication.
●● DNA polymerases are complex enzymes that move along the DNA molecule, linking nucleotides
to their complementary base partners to make new DNA chains.
●● Each strand of DNA is used as a template to make a new, complementary strand – this is known
as semi-conservative DNA synthesis.
●● Accurate DNA replication is important so that daughter cells have exact copies of the genetic
information for synthesising proteins.
●● Some DNA polymerase enzymes have the ability to ‘proof read and edit’ the DNA, correcting its
own errors. There are also complex enzymes to repair other damage to DNA caused by mutagens.

1 What is meant by ‘DNA replication’?

CHECK YOUR
UNDERSTANDING
2 Create a flow chart that accurately shows the steps in the process of DNA replication. Indicate any enzymes
that act at particular steps and what their function is. 3.3
3 What is an error in the DNA base pairing called and how does it arise?
4 Why is DNA replication called ‘semi-conservative’?

The importance of accuracy during

3.4 DNA replication
Because the sequence of bases in DNA makes up the genetic code of an individual, exact copying of this
sequence during replication is critical, for two main reasons:
◗◗ heredity (inheritance of genes) – the genetic material transmitted from cell to cell (by mitosis) and
from generation to generation (by gametes from meiosis) needs to be accurate
◗◗ gene expression (protein synthesis) – the genetic instructions given to a cell to create its structure Causes of
and ensure its correct functioning must be accurate. mutation and
types of mutation
It is therefore very important that no errors are made when this information is replicated. This is are dealt with in
termed fidelity of replication. more detail in
Chapter 7.
However, studies show that DNA is at constant risk of mutation. It is therefore not surprising to find
that cells have enzymes that are able to repair incorrect base insertions and other DNA damage that may
arise during replication.

Errors in replication
Natural errors that arise at random during DNA during replication are called spontaneous mutations. Other
errors that arise as a result of exposure of cells to environmental factors such as radiation or chemicals are
called mutagenic mutations. Environmental factors such as radiation, chemicals and viruses that change
DNA are called mutagens. As the length of time that the cells are exposed to mutagens increases, and the
intensity of exposure rises, so the risk of mutation also increases.
Shutterstock.com/nobeastsofierce

There are enzymes in cells to repair both types of mutation, but sometimes DNA
errors go undetected and this results in a permanent mutation. This uncorrected
mutation will be replicated in successive divisions and, if occurring in meiosis, passed
on to later generations of individuals.

DNA repair
The insertion of an incorrect base is common during DNA replication. When this
occurs, a repair enzyme recognises the mismatched base pair, excises the incorrect
base (cuts it out) and replaces it with the correct base. This is called DNA mismatch FIGURE 3.17 An uncorrected mutation in
DNA will be passed on to future generations.
repair and is a function of the enzyme DNA polymerase I.

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 AT AT AT However, if a mismatched pair is missed by the enzyme, harm
CG CG CG
GC AC AT
may arise the next time the cell divides and the DNA replicates. For
TA TA TA example, if an error occurs in which a ‘C’ nucleotide is accidentally
AT Error AT Replication AT paired with an ‘A’ nucleotide (instead of the correct ‘G’) (Fig. 3.18),
then the next time this DNA strand acts as a template and replicates,
FIGURE 3.18 Mismatch error in DNA
replication the error will cause a strand to form that has an incorrect base pair
(AT instead of GC).
This strand is now permanently mutated. The error will not be recognised by a DNA repair enzyme,
because the pairing of bases is not incorrect. However, the sequence of bases within the strand now
Variation and the differs from that of the original strand. A permanent mutation such as this often has a harmful effect,
advantages of
biodiversity in terms
but some mutations have a beneficial effect on the organism. Some mutations have no effect at all. It is
of evolution are dealt important to remember that mutation is a mechanism that can give rise to variation in organisms and
with in Chapter 5.
this may be beneficial to a species in terms of biodiversity and evolution.
There are different types of enzymes that carry out repairs on different types of DNA damage, to
ensure accurate replication and normal functioning of cells.

Gene expression and the need for accurate replication

Different types of
DNA damage and Now that you understand the structure of DNA and its ability to serve as a template during replication,
their effects, as well
as DNA repair are you might ask how the genetic code actually works. The answer lies in the production of proteins.
dealt with in Enzymes are proteins; they control the synthesis of cell materials and the biochemistry (metabolism)
Chapter 7.
within each cell and within the whole organism. Besides storing exact copies of instructions in every cell,
the sequence of bases in DNA also plays a vital role in the translation of the genetic code into proteins.
In multicellular organisms, different genes are activated and expressed in each type of cell. The entire
genetic code passed on to each cell must contain a full and accurate set of instructions, so that when the
A comparison of
two major DNA relevant gene is activated, it functions correctly to make proteins that determine the type of cell it will
repair pathways become. The activation of genes is regulated by other molecules, such as enzymes (which are proteins),
Analyse the
information in and therefore they also need to be accurately coded for by DNA. Errors in genes that control the cell cycle
the diagrams to may lead to changes in cell division and cell death, leading to cancer. Mismatch repair of these genes is
understand a visual
representation of DNA particularly important (Fig. 3.19). Replication errors in genes that code for DNA repair enzymes are also
mismatch repair.
linked to cancer.

Genes that regulate FIGURE 3.19

cell division to The cell cycle, Oncogenes
prevent cancer are showing the stage G1 promote cell growth
discussed in more at which mismatch
(cell growth)
detail in Chapter 15, repair enzymes M (mitosis)
pages 525–6. (coded by genes)
correct errors in DNA
during replication
G2

S (synthesis)
Mismatch repair G0 (resting)
genes code for enzymes that
correct replication errors Suppressor genes Modifier genes
influence cell function
inhibit cell cycle and
promote apoptosis
(programmed cell death)

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 KEY CONCEPTS
●● Accurate DNA replication is important for two main reasons:
– heredity: the genetic material of a cell must be transmitted accurately, from:
∎ one cell to another during mitosis, allowing growth, repair and maintenance of an organism
∎ one generation to another during meiosis (e.g. when gametes are formed for sexual
reproduction)
– gene expression: the genetic material of a cell must be transmitted accurately to give the
correct instructions to a cell to ensure the correct structure, functioning and behaviour of an
organism, essential for its survival.

CHECK YOUR
1 Explain the importance of having inbuilt mechanisms for correcting mistakes in DNA replication.
UNDERSTANDING
2 Distinguish between a spontaneous mutation and a mutagenic mutation.
3 List three mutagens. 3.4
4 How is an error in replication, such as a mismatch repair, corrected?
5 How could an error in DNA replication end up affecting a whole organism?

3.5 The continuity of species

All living organisms arise from other living organisms. The continuity of species refers to the ongoing
survival of species as a result of characteristics being passed from parents to offspring in a continuous
lineage. This inheritance of characteristics from ancestors to currently living organisms relies on
the passing on of consistently accurate genetic information (genetic stability) and the occasional
introduction of variation of some genetic information, so that species can adapt and survive in a
changing environment.
Accurate DNA replication brings about genetic stability, whereas mutation results in genetic
Genetic
variation. Although variation is important for evolution of species, genetic stability is important continuity
for the survival of the individual. Both genetic stability and variation play a role in ensuring the
continuity of the species.

Genetic continuity

Dreamstime.com/Vbaleha
Genetic continuity is a way of preserving genetic information across generations
and is dependent on two things:
◗◗ when a cell divides by mitosis, the resulting two daughter cells must have the
same number and type of genes as the original cell.
◗◗ when two sexually reproducing organisms breed, the resulting offspring must
have the same number of genes as the parent organisms and variations in these
genes must not be extremely detrimental or lethal.
Genetic continuity ensures continuation of a species, because it ensures that
new cells or organisms have all the genes they need, in working order, to survive. A
lack of genetic continuity results in disease and sometimes in death and extinction. FIGURE 3.20 Genetic continuity across
generations

Ensuring the continuity of species – genetic stability

In asexual reproduction, offspring inherit identical characteristics from one parent if replication is
consistent. In sexually reproducing organisms, offspring inherit a mix of characteristics from two
parents.

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 In both cases, the correct number of chromosomes and

Shuttestock.com/Volodymyr Burdiak
characteristics are passed on in family lineages. At a genetic level,
stability arises when chromosomes are replicated accurately to
give rise to identical daughter chromosomes.
For continuity at the species level, successful desirable traits
must be passed on, along with some random errors (chance
errors may be introduced by mutation). This allows a species
to evolve if an environmental change occurs. Natural selection
acts so that individuals in a population that are best suited to
FIGURE 3.21 Genetic continuity at the species level involves
conservation of diversity of traits. the environment survive and reproduce, passing on their genes
to their offspring. This mixing of parent genes during sexual
reproduction, including some that have arisen due to mutation, increases genetic diversity and helps
maintain continuity of the species.
The mechanisms that have evolved to ensure genetic continuity (passing on of genetic traits) and the
survival and continuity of species include:
◗◗ consistent replication prior to cell division (mitosis and meiosis)
◗◗ an orderly distribution of chromosomes when cells divide and when gametes form
◗◗ fertilisation methods that ensure that individuals of the same species breed successfully
◗ methods to ensure embryo survival, such as production of large numbers or protection
and nourishment of developing embryos and parental care
◗ natural selection so that the fittest survive to reproductive age and pass on their genes.
 echanisms that result in genetic variation in species include:
M
mutation – changes in DNA due to mutation may be spontaneous or mutagen-induced
◗ 

mixing of parental genes during sexual reproduction (brought about by crossing over
◗ 

and independent assortment during meiosis, and random fertilisation of gametes).

Genetic errors that threaten the continuity of species

You have learned that many variations that arise due to spontaneous changes in DNA are temporary and
are immediately corrected by enzymes that bring about DNA repair. The number of DNA repair enzymes
present in cells is an indication of how important accurate replication and DNA repair are for survival.
In humans, a range of diseases have been linked to the reduced ability of cells to repair DNA during or
after replication. Research shows that people who have a reduced ability to repair DNA tend to be more
susceptible to some cancers. A decrease in the ability of cells to repair DNA during replication is also
thought to be responsible for accelerated ageing and may give rise to neurodegeneration. There is a great
deal of current research in this area.
When a mutation is present in a DNA repair gene, the gene may be expressed in an altered form
or not expressed at all. For example, it has been found that people who have the disease Xeroderma
pigmentosum (XP) are unable to repair DNA and so they are more vulnerable to DNA damage from
ultraviolet rays and, as a result, to skin cancer. Recent research shows a link between germline mutations
in DNA repair genes and lethal forms of prostate cancer. Research in animal embryology shows that if
one of the genes for DNA repair (the base excision repair gene) is missing, this results in the death of the
embryo. Mutations such as these are termed lethal mutations.
Genetic information can only be stored in a stable form and passed on consistently if DNA repair
enzymes continuously scan the DNA for errors in replication and replace incorrect or damaged
nucleotides. Natural selection is a mechanism that ensures individuals carrying damaged genes are
removed from populations so that the continuity of species is not at risk.

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 INVESTIGATION 3.3

Literature review of the effect of cell replication processes on the Literacy

continuity of species
Information and
TASKS communication
technology
A Assess the effect of cell replication processes on the continuity of species. capability

B Present your findings in the form of a literature review.

C Acknowledge and evaluate your sources.
You will complete secondary-source research to review the effect of accurate replication as opposed to Threats to
inaccuracies in replication on the continuity of species. genetic
continuity
PART A: RESEARCH
Focus question: In what ways do cell replication processes support or threaten the continuity of species?
1 Draw up a chart in which you list the following in relation to the focus question:
– What you know: list any facts relevant to the topic that you have learned during this section of the course. What happens
when mitosis
– What you think you know: list anything based on prior knowledge, or that you think you understand. goes wrong?
– What you need to find out: outline some research questions (key concepts) and write some key words.
2 Read the articles listed in the weblinks provided and find your own articles, and use these to gather valid
and reliable information relevant to your research questions.
3 Interpret and analyse your search results. (You may need to modify your search strategy as you go.)
Cancer-specific
4 Make a judgement and use your findings to support your concluding judgement in answering the focus question. defects in DNA
repair pathways
PART B: PRESENT YOUR FINDINGS

5 Write a literature review of approximately 400 words (page 9), with:

WS
– an introduction, where you define the topic
– a body, where you group the literature and your findings according to common themes Evaluating
Homework
resources
– a conclusion, where you explain the link between your research question and the literature you have
reviewed, creating an evidence-based argument.
Evaluate your
PART C: ACKNOWLEDGE SOURCES sources using the
CRAAP technique.
6 Acknowledge and evaluate your sources using an accepted referencing style. (See page 10.)
KEY CONCEPTS

●● Genetic continuity relies on:

– consistent replication of genetic information that is passed from a parent cell to daughter
cells, resulting in continuity in the traits being passed from parents to offspring
– the effect of natural selection and evolution on the gene pool as a result of:
∎ introduction of variation during sexual reproduction
∎ random errors arising by mutation, being replicated and passed on to offspring.
●● Random variations that confer an advantage may be selected over those that confer no
advantage or are harmful.

1 Explain the role of DNA replication in:

CHECK YOUR
UNDERSTANDING
a maintaining genetic continuity in a species b introducing genetic variation in a species.
2 Explain how natural selection can remove harmful variations from a species. 3.5

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 3 CHAPTER SUMMARY
Cell replication: How important is it for genetic material to be replicated exactly?
PROCESSES INVOLVED IN CELL REPLICATION: MITOSIS AND MEOISIS
Roles in life cycle Importance of mitosis and meiosis

Mitosis
Fusion to
form zygote Zygote (2n)
• Embryonic development: zygote blastula embryo
(n to 2n) •• Growth of multicellular organisms – meristem (plants); stem
cells (animals)
Egg (n) Sperm (n) •• Tissue maintenance and repair
Juvenile (2n)
•• Asexual reproduction and genetic stability in populations
Meiosis
(2n to n)

Meiosis Mitosis
(2n to n) (differentiation Meiosis
and growth)
•• Production of gametes
•• Halving the chromosome number
Adults (2n)
Haploid stages •• Introduction of genetic variation
Diploid stages

G0
Cell cycle stops
The cell cycle

G1 – cell growth
INT

Cellular contents,
ERP

excluding the
S – synthesis
HASE

chromosomes,
Each of the 46
are duplicated
chromosomes is
duplicated by the cell

Cytokinesis

Cytoplasm
divides
M – mitosis
Nucleus G2 – 9proof reading9
divides The cell checks the
duplicated chromosomes
for errors, making
any needed repairs
IT
M

O SI S

Stages of mitosis

Interphase – DNA replicates

Prophase – chromosomes appear and split into chromatids

Metaphase – chromosomes align on equator

Anaphase & telophase – daughter chromosomes segregate

and move to poles

Cytokinesis – cytoplasm divides

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 DNA STRUCTURE AND REPLICATION: THE WATSON CRICK DNA MODEL

Nucleotide composition, complementary base pairing and bonding

Nucleotide = phosphate − sugar − nitrogenous base

Nitrogenous base pairing:

Sugar-phosphate ‘backbone’
adenine + thymine; cytosine + guanine
Hydrogen bonds between

Bonds = weak hydrogen

O
P
nitrogenous bases
O
P
G
O C
P
C O

O G P

P A Complementary Original
O
T
O
chains of DNA DNA helix
P
P G

O
O
C
P
OH T
3 end

A
O
Replication
P
fork
Phosphodiester
O
bond
Pool of
nucleotides
P

5 end

Building
units
T A T A
DNA DNA
Semi-conservative DNA replication polymerase
G C G C
polymerase

•• DNA unzips and a replication fork can

be seen. Okazaki fragments
•• Each strand acts as a template. Template joined on this
strand by ligase
•• DNA polymerase adds nucleotides
from 3’ to 5’ end.
•• New molecules consist of one old New
chain
and one new strand.

Leading Lagging strand

strand
3' end 5' end
Original
complementary strands

Ensuring exact replication (enzyme regulation) Accurate DNA replication ensures the
survival of the species.

Oncogenes
G1 promote cell growth
(cell growth)
M (mitosis)
Helicase unwinds Binding proteins stabilise Primase adds a short

Helicase unwinds Binding proteins

parental double helix.
Primase adds a short
stabiliseprimer to template strand.
the strands.

parental double helix. the strands. primer to template strand.

G2

DNA polymerase binds Exonuclease removes Ligase joins Okazaki

nucleotides to form
new strands.
RNA primer and inserts
correct bases.
fragments and seals nicks in
sugar-phosphate backbone.
S (synthesis)
Mismatch repair G0 (resting)
genes code for enzymes that
correct replication errors Suppressor genes Modifier genes
influence cell function
inhibit cell cycle and
DNA polymerase binds Exonuclease removes Ligase joins Okazaki promote apoptosis
nucleotides to form RNA primer and inserts fragments and seals nicks in (programmed cell death)
new strands. correct bases. sugar-phosphate backbone.

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 3 CHAPTER REVIEW QUESTIONS Qz

Review quiz

1 Name the three types of cell division found in all living c Draw a sequence of diagrams of a dividing cell (to
organisms and the purpose of each. scale) to show how the cell changes in cell volume
during the process of mitosis and use data from the
2 Identify the components of chromatin.
graph to justify your explanation.
3 Explain why chromosomes are only visible during cell division. d Describe the relationship between the cell volume and
4 Name the phases of the cell cycle and outline what the amount of DNA in the cell.
happens in each phase. e The cell volume graph is divided into ten units of time.
At what time does cytokinesis occur? (Express this in
5 Give three reasons why it is important for the cell cycle to
numbers of time units.) Give a reason for your answer.
be regulated.
7 Distinguish between mitosis and cytokinesis.
6 The graphs in Figure 3.22 show cell volume (the size of a
cell) and the amount of DNA in a cell at various stages of 8 Explain why it is important for DNA to replicate before cell
the cell cycle. division.
a How many cell cycles (cell divisions) has the cell shown 9 Draw and label a diagram of an RNA nucleotide.
in the graphs completed? Justify your answer.
10 State the rule of base pairing of nitrogenous bases in a
b Explain the change in the amount of DNA during the
DNA molecule.
cell cycle, as shown in the lower graph. Use correct
terminology for the phases of the cell cycle and the
stages of mitosis.
Cell volume

Time
Amount of DNA

G1 S G2 M 1 cell cycle

FIGURE 3.22

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 11 Based on the timeline in Figure 3.23, create a table to summarise the contributions made by scientists in the discovery of the
hereditary material in cells.

Left to right: Alamy Stock Photo/Pictorial Press Ltd; Walter S. Sutton Collection, University of Kansas Medical Center
Archives, Kansas City, KS; Getty Images/Hulton Archive/Stringer; Alamy Stock Photo/PF-(bygone1); Getty Images/
Bettmann Science Photo Library; Bottom: Alamy Stock Photo/Paul Fearn
1944
Avery,
1930s McLeod
Hammerling and 1952
1902 shows that McCarty Hershey
1865 Sutton 1927 hereditary 1931 show that and Chase
Mendel and Boveri Muller information McClintock DNA is the use radioactive 1990s
documents propose shows that is contained demonstrates ‘transforming labelling to Genome
patterns of chromosome X-rays in the nuclei genetic principle’ prove that DNA sequencing
heredity in theory of induce of eukaryotic recombination responsible is responsible projects
pea plants heredity mutations cells in corn for heredity for heredity begin

1869 1915 1928 1941 1950 1953 1961

Miescher Morgan Griffith’s Beadle Chargaff Watson Jacob
first identifies and his ‘Fly Room’ ‘transformation and Tatum discovers and Crick and
DNA (‘nuclein’) colleagues confirm experiments’ describe the that A 5 T propose Monod
the chromosome transform ‘one gene–one and C 5 G the double propose
theory of heredity non-pathogenic enzyme’ (Chargaff’s helix the
bacteria strains hypothesis rules) structure existence
to pathogenic of DNA of mRNA

FIGURE 3.23 Timeline of contributions by scientists to our understanding of genetics and inheritance

12 List two hypotheses that Watson and Crick tested when 18 Explain the following features of the Watson and Crick
creating the models listed below, and explain how their model of DNA, supplementing your explanation with
findings supported their hypotheses: simple diagrams where necessary:
a DNA structure a nucleotide composition
b DNA replication. b nucleotide pairing
13 Explain how nucleotides are added to both strands during c nucleotide bonding.
DNA replication. Use a diagram to illustrate your answer. 19 Assess the effect of the following processes on the
14 Predict the DNA sequence for a complementary strand continuity of species:
of DNA made from the following sequence of bases: a DNA replication
AATTGGCTGACGAATCAT. b action of polymerase enzymes during DNA replication
15 Outline the role of four named enzymes in cell replication. c separation of daughter chromatids during mitosis
16 Explain why DNA replication is referred to as ‘semi- d formation of identical cells in multicellular organisms
conservative’ replication. e cell replication in unicellular organisms
17 Explain, giving examples, the importance of exact f replication of cell organelles during G1 phase.
replication of DNA in cell division and explain the role 20 Answer the inquiry question at the start of this chapter:
that enzymes play in ‘proof reading’ the molecule after How important is it for genetic material to be replicated
replication. exactly? Use information from this chapter to support your
answer.

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