Module in

Genetics
Wilfredo B. Barrera, Jr., LPT, M.Sc.
Department of Mathematics and Natural Sciences
Southern Luzon State University

REVIEWERS
Su Latt Phyu, Ph.D.
Department of Plant Breeding, Physiology and Ecology
Yezin Agricultural University
Sonali Bej, Ph.D. Jed Frank Marqueses, M.A.
Crop Improvement Division
Department of Mathematics and Natural Sciences
Indian Institute of Rice Research Southern Luzon State University
 Module 2
Lesson 5: Replication and Transcription 1
The Structure of Nucleic Acid
The Structure of DNA
DNA as the Genetic Material
Level of Organization of DNA
Replication of the Genetic Material
RNA as the Genetic Material
Transcription: DNA to RNA

Lesson 6: Gene Expression 10

Translation: Protein synthesis
Gene Regulation in Prokaryote
Gene-Enzyme Relationship
Gene Regulation in Eukaryote

Lesson 7: Developmental Genetics 18

The Genetic Basis of Development
Embryonic Development of D. melanogaster
Developmental Programs in Plants and Animals
Differential Gene Action and Epigenetic Control
Neoplasmic Interactions

This module is strictly for SLSU use only.

No part of this module may be reproduced or transmitted in any form or by any means, electronic or
mechanical, including printing, photocopying, recording or any information storage or retrieval system without
permission from the compiler. Some of the illustrations used in this module are subject to copyright.
Cover layout: Rod Anthony A. Sopeña
Module 2 - Molecular Genetics Lesson 5 - Replication & Transcription

5 Replication &
organic base (Figure 5.1). In our example, the
phosphate residue is bonded to the carbon in

Transcription position number 5’ of the monosaccharide

ring. Because this nucleotide is formed from
one phosphate residue, a deoxyribose residue,
In this lesson, you are going to learn about the and an adenine nitrogenous base, we write its
chemical basis of heredity. At the end of the name as deoxyadenosine 5’-monophosphate.
lesson you should be able to:
1. Describe the chemical structure of
polynucleotides (DNA and RNA) and
identify.
2. Discuss the basic mechanisms of DNA
replication and identify and describe
the requirements of DNA replication.
3. Explain the process of transcription
and compare and contrast gene
transcription and DNA replication.
4. Describe the characteristics and the phosphoester
role of mRNA and explain the bonding pattern
mechanisms and outcomes of mRNA
modifications in eukaryotes.
N-glycosidic bond
The structure of nucleic acid
Genes contain genetic information and are
a nucleotide
passed from parents to offspring and the next
generation. Genes could be regarded as a Figure 5.1 The formation of phosphoester and
chemical compound or molecule. A nucleic glycosidic bonds in a nucleotide.
acid is a term used for the class of biological
polymers consisting of deoxyribonucleic acid It is common to use abbreviations
(DNA) and ribonucleic acid (RNA). when naming nucleotides. In this case, we
Nucleoside contains a nitrogenous would abbreviate the name as 5’-dAMP where
base and pentose sugar. It is composed of the lowercase d indicates deoxy. Figure 5.2
purine or pyrimidine base and ribose or shows the different compositions of
deoxyribose sugar. A nucleotide is a nucleotides. There are two major types of
nucleoside with phosphate group added. The nitrogenous bases such as purine which
phosphoester bonding pattern occurs when a includes adenine and guanine, and pyrimidine
monosaccharide bonds with the phosphate. which includes cytosine, thymine and uracil.
An N-glycosidic bond is made when the Thymine is present only in DNA and uracil is
monosaccharide bonds to nitrogen in the present only in RNA.

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Module 2 - Molecular Genetics Lesson 5 - Replication & Transcription

Figure 5.2 The different compositions of nucleotides.

The nucleotides that make up the DNA

For this reason, the alternating
contain a deoxyribose residue while the
monosaccharide and phosphate residues in a
nucleotides that make up the RNA contain a
polynucleotide strand are referred to as the
ribose residue. A polynucleotide is a polymer
sugar-phosphate backbone.
made from nucleotide residues. A chain of
covalently bonded nucleotides is referred to
as a strand. Nucleotides are covalently
bonded to each other by a phosphodiester
bonding pattern between a 3’ carbon on one
monosaccharide and 5’ carbon on the other
monosaccharide. We call this bonding pattern
a 3’  5’ phosphodiester because the
phosphate group is between the 3’ and 5’
carbons (Figure 5.3).
The nucleotides are connected to each
other by covalent bonds between the
monosaccharide residues and the phosphate Figure 5.3 Phosphodiester bonding in a dinucleotide.
residues.

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Module 2 - Molecular Genetics Lesson 5 - Replication & Transcription

One end of the strand is terminated with a together by especially strong hydrogen
phosphate group that is attached to a 5’ bonding between specific pairs of nitrogenous
carbon; we call that end of the strand the 5’ bases, 10 base pairs per helix turn and
terminus. The other end of the strand is nitrogenous bases stacked with above the
terminated with a hydroxyl group (OH) that is other. Adenine (A) forms two hydrogen bonds
attached to a 5’ carbon; we call that end of with thymine (T) bonds while guanine (G)
the strand the 3’ terminus. bonds with cytosine (C) by three hydrogen
bonds. The reason for the strong hydrogen
The structure of DNA bonding between these organic base pairs is
that they have complementary shapes (Figure
Genetic information, the information used to
5.5). The hydrogen bonding between these
make the various proteins and thereby
pairs of organic bases is referred to as base
enabling life, is contained in the sequence of
pairing or complementary base pairing.
nucleotides in DNA. The sequence of
nucleotides in DNA is referred to as the
primary structure of DNA.
DNA is composed of a combination of
deoxyribonucleotides that contain adenine,
guanine, thymine, or cytosine nitrogenous
bases. Nucleotides are given a one-letter
abbreviation based on the first letter in the
name of their nitrogenous base. In order to
describe the primary structure of a DNA
strand, it is customary to list the one-letter
code abbreviation of nucleotides in the order
that the nucleotide residues appear in the
strand e.g. 5’TGCA3’.
X-ray diffraction studies by Rosalind
Franklin (1950–1953) showed that DNA has
3.4-angstrom periodicity; a characteristic of
the helical structure. It was proposed by
Watson and Crick in 1953 that DNA exists as a
double helix, which is made from two Figure 5.4 Double helical structure of the DNA and its
polynucleotide strands (Figure 5.4). The two complemenrary base pairing.
DNA strands of a DNA double helix have their
5’ terminuses and 3’ terminuses with opposite In a double helix DNA, base composition
orientations. We call this opposing orientation studies quantitatively demonstrated that the
of the DNA strands as an antiparallel amount of adenine is equal to the amount of
arrangement. The two DNA strands are held thymine. Similarly, the amount of guanine is

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Module 2 - Molecular Genetics
equal to the amount of cytosine. This known
as Chargaff’s rule thus, the sum of purines (A +
G) is equal to the sum of pyrimidines (C + T).
However, the percentage of (G + C) is not
necessarily equal to the percentage of (A + T).
Figure 5.5 Complementary base pairing in DNA.
DNA as the genetic material
Proteins and nucleic acids are major
candidates for the genetic material. Before,
many geneticists favored proteins as the
genetic material due to its diversity and
abundance in cells. DNA is thought to be too
simple to be genetic material because there
are only four types of nucleotides, whereas
4
Lesson 5 - Replication & Transcription
proteins have 20 different amino acids.
Recently, two additional amino acids such as
selenocysteine and pyrrolysine were
discovered bringing the amino acids to 22.
The major characteristics needed for a
molecule to serve as genetic material include
storage of tremendous genetic information,
accurate duplication and transfer of
information, stable (resistant to frequent
changes), capable of mutation, and dynamic
and flexible enough to respond to other
molecules or changing environment.
In eukaryotes, indirect and direct
evidence supports the concept that DNA is the
genetic material. There is a close correlation
between gametes and diploids in the amount
of DNA and the number of chromosome sets
while this is not observed in proteins.
Molecule serving as genetic material is
expected to absorb at a mutagenic
wavelength (260 nm). DNA absorbs UV at 260
nm while protein absorbs UV at 280 nm.
Recombinant technology provides direct
evidence that DNA is the genetic material.
Level of organization of DNA
DNA is a very large molecule that needs
packing in order to fit inside the cell. In
bacteria, which do not have nuclei,
chromosomes exist as circular units, called
circular chromosomes which can twist or fold
into more compact shapes by negative
supercoiling (Figure 5.6). The bacterial
chromosome is largely devoid of associated
proteins, much smaller than eukaryotic
chromosomes and contains less genetic
information.
Module 2 - Molecular Genetics
In eukaryotic organisms such as plants,
animals, and fungi, the DNA double helix coils
into a more compact structure (Figure 5.7).
Figure 5.6 Level of organization of DNA in bacteria.
DNA is organized into nucleosome
which is composed of an octamer, two of each
type of histone proteins such as H2A, H2B, H3
and H4. The nucleosome further condenses
into a fiber that is called chromatin.
Figure 5.7 Level of organization of DNA in
eukaryotes.
Lesson 5 - Replication & Transcription
Depending on the cell life cycle, chromatin can
undergo further coiling. For example, before a
cell divides chromatin is coiled into its tightest,
most compact shape, which is called a
chromosome.
Figure 5.8 The flow of the genetic
information.
Replication of the genetic
material
The flow of the genetic information Figure 5.8
is known as the central dogma of molecular
biology. A process in which a duplicate copy of
a DNA double helix is made is called DNA
replication. It is an essential function of
genetic material. It must be precise for genetic
continuity between cells following cell division.
There are three modes of DNA
replication such as (1) semiconservative - each
replicated DNA molecule consists of one old
and one new strand; (2) conservative - two
newly synthesized strands come together and
the original helix is conserved; and (3)
dispersive - parental strands are dispersed
into two new double helices. Scientists have
proven that DNA replication is
semiconservative in prokaryotes and in
eukaryotes.
DNA replication is complex, involves
several enzymes and basically divided into
three major steps such as initiation,
elongation and termination. Figure 5.9
summarizes the process of DNA replication.
Each of the two original DNA strands called
5
Module 2 - Molecular Genetics Lesson 5 - Replication & Transcription

the parent strands is used as a template for
the formation of a new daughter strand.
Replication starts at the ORI site, proteins
called helicases unwind and then open
sections of the double helices by disrupting
the hydrogen bonds between base pairs thus
creating a replication fork.
strands of the double helix are antiparallel and
DNA polymerase III only synthesizes 5ʹ to 3ʹ
direction. As a result, one strand such as the
leading strand is continuously synthesized
while the other strand such as the lagging
strand is discontinuously synthesized.

Figure 5.9 Overview of the DNA replication. Source: Klug et al. (2017).

Unwinding the double helix creates a

tension that is relieved by DNA gyrase by
making single- or double-stranded cuts.
Single-stranded binding proteins (SSBPs)
stabilize the open conformation of the helix by
binding specifically to single strands of DNA.
The primase recruited to replication fork by
helicase synthesizes RNA primer which
5’ end attaches to
provides free 3'-OH required by DNA 3’ end
+
polymerase III for elongation.
Chain elongation by the DNA +
polymerase III occurs in 5' to 3' direction by
adding one nucleotide at a time to 3'-OH. The
addition of nucleotide allowed two terminal
phosphates to be cleaved off providing new
exposed 3'-OH. The 3'-OH can participate in
addition to another nucleotide as DNA
synthesis proceeds (Figure 5.10). The two Figure 5.10 Chemical bonding in DNA replication.
Source: Klug et al. (2017).

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Module 2 - Molecular Genetics
The lagging strand is synthesized as Okazaki
fragments each with RNA primer. DNA
polymerase I removes primers on the lagging
strand and DNA ligase catalyzes the formation
of phosphodiester bonds to seals nicks and
joins fragments. Both DNA strands are
synthesized concurrently at a single
replication fork. Lagging strand looped and
inverts physically but not biochemical
direction. DNA clamp prevents core enzyme
dissociation from the template.
Proofreading and error correction is an
integral part of DNA replication. Occasionally,
the synthesis of non-complementary base
pairs is inserted. DNA polymerase proofreads
its work to see if the correct
deoxyribonucleotide residue has been added.
The 3’5’ exonuclease activity of the enzyme
allows the excision of nucleotides if a mistake
has been made.
RNA as the genetic material
Some viruses have RNA core instead of
DNA. For example, the tobacco mosaic virus
demonstrated RNA serves as genetic material.
The replication of viral RNA is dependent on
RNA replicase. Retroviruses have unusual
replication method. RNA serves as a template
for DNA synthesis. Complementary DNA is
synthesized by RNA-dependent DNA
polymerase reverse transcriptase.
The discovery of the reverse
transcriptase led to the development of the
quantitative real-time polymerase chain
reaction (qPCR). This technology has many
applications in different areas such as virus
detection and gene expression analysis of
agricultural crops.
Lesson 5 - Replication & Transcription
Transcription: DNA to RNA
A gene is generally defined as a region of DNA
that carries the information needed to
produce a protein. In the transcription process,
the information such as the sequence of
nucleotides in a gene is used to create a
specific sequence of ribonucleotides in a
single-stranded messenger RNA (mRNA). The
transcription process is similar to DNA
replication, with three main differences: (1)
RNA is produced in transcription, whereas
DNA is produced in DNA replication; (2) in
transcription, only one of the DNA double
helix strands is used as a template to produce
one mRNA strand, whereas in DNA replication,
each of the two DNA strands in a double helix
produce a daughter strand; and (3) different
enzymes are used.
Transcription (Figure 5.11) involves
three general steps which include (1) initiation
- RNA polymerase catalyzes initiation by
binding to promoter and insertion of first 5'
ribonucleotide triphosphate; (2) elongation –
continuous addition of the ribonucleotide
triphosphate; and (3) termination – end of
transcription once the termination nucleotide
sequence is encountered.
Transcription begins with template
binding by RNA polymerase at the promoter.
The promoter is a specific DNA sequences in 5'
region upstream of the initial transcription
point. The sigma (σ) factor subunit is
responsible for promoter recognition
(initiation of transcription). When RNA
polymerase binds to the promoter site, the
hydrogen bonding between base pairs within
the DNA double helix is disrupted. This
unwinds and opens a section of the double
7
Module 2 - Molecular Genetics
helix of the gene. Next, RNA polymerase
moves along one strand of the gene while
catalyzing the addition of free ribonucleotides
that are complementary to the DNA template,
to the 3’ end of a growing mRNA strand.
When the RNA polymerase reaches a
nucleotide sequence in the gene, called the
termination site, the mRNA strand is released.
In bacteria, termination transcribed into RNA
causes newly formed transcript to fold back
on itself e.g. hairpin. At times, termination
depends on the rho (ρ) termination factor.
free ribonucleotides
promoter
Figure 5.11 Overview of transcription.
In bacteria during transcription, large
polycistronic mRNA is produced which
encodes more than one protein. Genes in
bacteria called cistrons. On the other hand,
eukaryotes have monocistronic mRNAs.
Transcription in eukaryotes differs from
prokaryotic transcription in several ways. It
occurs within the nucleus unlike in
prokaryotes which happens in the cytoplasm.
After transcription, mRNA must leave the
nucleus for translation.
8
Lesson 5 - Replication & Transcription
Eukaryotic mRNAs require processing
or transcriptional modifications to produce
mature mRNAs. This includes the addition of
5' cap or capping, the addition of 3' tail or
tailing and excision of introns or splicing. The
5' cap protects the mRNA from nuclease
attack and may be involved in transport of
transcript across the nucleus while the 3' poly-
A tail aids in the transport to cytoplasm.
Basically, the mechanism of splicing involves
the cutting of intron through its end by the
endonuclease.
The terminal ends of the adjacent exons
joined by ligase to form the mature mRNA.
This results in a mature mRNA smaller than
initial RNA. Alternative splicing produces
related protein from a single gene thus,
increasing the number of products derived
from a single genome.
Module 2 - Molecular Genetics Lesson 5 - Replication & Transcription

Learning Check! 5. The linear chromosome of phage T2 is 52

µm long. The chromosome consists of
1. If the GC content of a dsDNA molecule is double-stranded DNA, with 0.34 nm
56 percent, what are the percentages of between each base pair. How many base
the four bases (A, T, G, and C) in this pairs does a chromosome of T2 contain?
molecule?
6. The following table lists the relative
2. Calculate the approximate number of base percentages of bases of nucleic acids
pairs in ten helical turns of the DNA based isolated from different species:
on the values from early X-ray diffraction
structures. Species A G T C U
1 21 29 21 29 0
2 29 21 29 21 0
3. Which enzyme (DNA polymerase, RNA
3 21 21 29 29 0
polymerase, or reverse transcriptase) is 4 21 29 0 29 21
involved in each of the processes below: 5 21 29 0 21 29
a. Transcription
b. Replication For each species, what type of nucleic acid is
c. Using a DNA template to make RNA involved? Is it double or single-stranded?
d. Using an RNA template to make DNA Explain your answer.

4. The following diagram represents the

structure of a gene in Drosophila
melanogaster; blue segments are exons,

and yellow segments are introns.

a. Which segments of the gene will be
represented in the initial RNA
transcript?
b. Which segments of the gene will be
removed by RNA splicing?
c. Which segments would most likely
bind proteins that interact with RNA
polymerase?

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Module 2 - Molecular Genetics
6 Gene
Expression
In this lesson, you are going to learn about the
chemical basis of heredity. At the end of the
lesson you should be able to:
1. Understand and explain the
mechanism of translation.
2. Explain what the genetic code is and
identify its major characteristics.
3. Use the genetic code table to predict
the sequence of amino acids in the
polypeptide that would be produced in
translation.
4. Define the term gene regulation and
compare and contrast the basic
principles of gene regulation in
eukaryotes and prokaryotes.
Translation: protein synthesis
The information contained in the sequence of
DNA nucleotides is used to generate proteins.
This process is central to the existence of all
known life forms because proteins are critical
in directing and controlling cell growth and
function, and in regulating the metabolism of
an organism.
The process of making proteins from
the information in DNA is called protein
synthesis or gene expression. The mRNA
produced in transcription contains the
information from a gene that is required to
produce a protein. Therefore, there is
colinearity between the gene and the protein
that will be produced. This led to one-gene:
10
Lesson 6 - Gene Expression
one-enzyme hypothesis which was changed to
one-gene: one-polypeptide chain hypothesis.
Two factors modified the hypothesis because
(1) nearly all enzymes are proteins but not all
proteins are enzymes; and (2) proteins have a
subunit structure with two or more
polypeptide chains. However, there should be
a functional protein for a gene to be
expressed, therefore, one-gene: one-
functional protein is more appropriate.
In translation, the sequence of
nucleotides in mRNA is converted (translated)
to a sequence of amino acid residues in a
polypeptide. The genetic code is based on
three-nucleotide sequences called codons. A
codon directs the addition of a specific amino
acid residue to a polypeptide that is being
formed.
The genetic code is degenerate—a
given amino acid can be specified by more
than one triplet codon. The start and stop
signals are triplets that initiate and terminate
translation. Once translation begins, codons
are read with no break i.e. commaless. Any
single ribonucleotide within mRNA is part of
one triplet thus, nonoverlapping.
Many amino acids are specified by
more than one codon. Figure 6.1 shows the
codons that specify the 20 amino acids. Only
tryptophan and methionine are encoded by a
single codon. The genetic code shows order—
chemically similar amino acids share one or
two middle bases in triplets encoding them.
Wobble hypothesis states that codons that
specify the same amino acid differ at third
base position. Thus, the initial two
ribonucleotides of triplet codes are often
more critical than the third.
Module 2 - Molecular Genetics
Figure 6.1 The genetic code specifying the 20 amino
acids and the stop codons. Source: Klug et
al. (2017).
The codon AUG is the initiator which
codes for methionine. In bacteria, methionine
is modified in a form of N-formylmethionine
(fmet) but appears internally in mRNA
unformylated. There are three termination
codons such as UAG, UAA, and UGA. They do
not code for any amino acid and are not
recognized by tRNA. Translation terminates
when these codons are encountered.
The mechanism of translation involves
three steps (Figure 6.2) such as initiation,
elongation and termination. Once the
messenger RNA leaves the nucleus and enters
the cytoplasm, a ribosome attaches to its start
codon (AUG). Ribosomes are relatively large
particles that contain protein and very long
RNA strands called ribosomal RNA (rRNA).
Next, a transfer RNA with a start anticodon
(UAC) binds to the mRNA start codon to form
a ribosome-mRNA-tRNA complex. The
Lesson 6 - Gene Expression
formation of this complex is called initiation.
Elongation begins when a second tRNA binds
to the initiation complex. The second tRNA
delivers the amino acid that corresponds to
the codon that follows the start codon. A
polypeptide chain is made by the formation of
peptide bonds between the amino acid
residues that are specified by the genetic code
in mRNA. A part of the ribosome, called the
transferase center, catalyzes the formation of
peptide bonds between the amino acid
residues. The ribosome moves a distance of
three bases (a codon) along the mRNA strand
in the 5’ to 3’ direction. The tRNA that was
attached to the first amino acid detaches from
the mRNA and diffuses throughout the
cytoplasm where it will encounter an enzyme
that catalyzes its reattachment to an amino
acid that matches its anticodon. Termination
occurs when the ribosome reaches a stop
codon. When this happens, the polypeptide,
mRNA, tRNA, and ribosome become
separated.
In some cases, the polypeptide that is
produced in translation is a fully functional
protein. In other cases, the polypeptides
undergo further folding to acquire the
secondary and tertiary structures that they
need in order to function. In many cases,
multiple polypeptide subunits must assemble
into a quaternary structure in order to form a
fully-functional protein. Some proteins require
covalent bonding modifications, called post-
translational modifications after they are
transcribed. An example of post-translational
modification is the addition of heme groups to
heme-containing proteins, such as
hemoglobin and myoglobin.
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Module 2 - Molecular Genetics Lesson 6 - Gene Expression

ribosome-mRNA-tRNA complex (a)

first amino acid

messenger RNA
(mRNA)

peptide bond (b)

several cycles of
amino acid addition

(c)

Figure 6.2 Overview of translation (a) initiation; (b) elongation; and (c) termination.

Gene regulation in prokaryote Sometimes, several genes are

transcribed from the same promoter site. A
Gene expression is another term used for
section of DNA made up of genes that are
protein synthesis. Another way to control
transcribed from the same promoter site is
metabolic pathways, or other conditions that
called an operon. Figure 6.3 is an example of a
involve proteins, is to regulate the rate of
lactose operon in bacteria. A gene is under
production of a protein or enzyme. Increasing
negative control unless a repressor protein
the rate of gene expression is called
blocks it. If a gene is under positive control, a
upregulation while decreasing the rate of
regulatory protein is necessary to promote
gene expression is called downregulation. The
transcription. Because the repressor is always
major point of control of gene regulation is
produced, the lac operon is inducible.
transcription.

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Module 2 - Molecular Genetics
DNA
promoter (P site)
Figure 6.3 The lac operon showing the regulatory gene, promoter, operator and structural genes.
The lac operon consists of three
protein-coding genes plus associated control
regions that code for proteins involved in the
metabolism or transport of lactose. The three
genes of the lac operon are called lacZ, lacY,
and lacA. The lac operon is preceded by a lac
regulator, lacI gene. The lacI gene produces an
active repressor protein that binds to a
segment of DNA called an operator site.
When the repressor protein is bound
to the operator site, RNA polymerase is
blocked from moving along the lac operon to
transcribe the three structural genes
therefore, the operon is off (Figure 6.4a). In
the presence of glucose, lactose is not needed
as an energy source and almost zero lactose is
allowed into the bacterial cell, therefore it is
not necessary for the genes of the lac operon
to be expressed. Let us consider the scenario
in which lactose is present in the absence of
glucose. Whenever lactose is present, some of
it is converted to a compound called
allolactose which binds to and thereby
inactivate the repressor protein. The inactive
repressor protein cannot bind to the operator
site thus, allowing RNA polymerase to move
along the lac operon and transcribe the lac
operon genes (Figure 6.4b).
Lesson 6 - Gene Expression
lac operon
The mRNAs from the lac operon genes are
translated to the three proteins: β-
galactosidase, β-galactoside permease, and β-
galactoside transacetylase. β-galactosidase is
an enzyme involved in lactose metabolism. β-
galactoside permease is a transmembrane
protein that transports lactose into cells. β-
galactoside transacetylase is an enzyme
thought to be involved in the breakdown of
some nonlactose species that are transported
into cells by permease. The presence of
lactose and the subsequent presence of
allolactose upregulate the genes of the lac
operon. Whenever lactose is no longer
available, the concentration of allolactose will
decrease and then active regulator proteins
become available again to downregulate the
lac operon.
Another regulatory mechanism of the
lac operon is called the catabolite
repression—an example of positive regulation.
RNA polymerase does not bind efficiently to
the promoter unless the catabolite activating
protein (CAP) is present. CAP binds to the 5’
end of the promoter at the CAP binding site.
When the CAP is bound, the binding of RNA
polymerase is enhanced. The presence of
glucose determines whether cAMP levels are
high or low. If glucose is present, the cell does
13
Module 2 - Molecular Genetics Lesson 6 - Gene Expression

not need to metabolize lactose for energy. operon, the operator and promoter are cis-
The presence of glucose results in low levels acting because they function just upstream of
of cAMP. Without cAMP, the transcription of the lac structural genes.
the lac operon is inefficient.

RNA polymerase (a)

operator (O site)

(b)

RNA polymerase

Transcription

mRNA mRNA mRNA

Translation

β-galactosidase permease transacetylase

Figure 6.4 The lac operon: (a) no transcription of the structural genes; and (b) there is transcription of
structural genes.

DNA sequences can affect other
sequences either next to them on the same
chromosome e.g. at a cis-acting site, or at a
relatively distant site or sites on different
chromosome (trans-acting site). In the lac
In contrast, the lacI gene can act as trans
producing a diffusible repressor that can
travel to a sequence a great distance away.
Several mutants were discovered and
developed in the laboratory to understand

14
Module 2 - Molecular Genetics Lesson 6 - Gene Expression

how genes are regulated in the lac operon. In The repressor is by a regulatory gene, trpR
I- repressor mutants, the lacI gene does not and is inactive by default. Therefore, the five
produce a functional repressor protein and structural genes are transcribed. These
RNA polymerase is never inhibited therefore, structural genes codes for polypeptides
the lac operon is always transcribed. Similarly, subunits that make up enzymes for the
the operon is always on in I-d mutant repressor synthesis of tryptophan. When tryptophan is
gene because it cannot bind to the operator present, it binds to the trp repressor protein,
sequence. In Is super-repressed mutation, the which turns the operon off (Figure 6.6). The
mutant repressor gene cannot bind inducer. repressor is active only in the presence of its
Thus, the repressor protein binds very tightly corepressor tryptophan thus, the trp operon is
to the operator and does not bind to lactose. turned off (repressed) if tryptophan levels are
Therefore, the lac operon is always repressed, high.
even when lactose is present. For the
operator mutants Oc the gene to which it is promoter
attached is affected. The mutants are said to
act in cis or to be cis-dominant. The operator
X
DNA
is mutated so that the repressor can no longer
no mRNA
bind to it. Therefore, the structural genes are mRNA made
RNA polymerase
always transcribed. Conversely, in cis-acting Os blocked
mutants, the operator binds so tightly to the protein
repressor that it never gets released active
repressor
Therefore, the operon is always repressed.
tryptophan
An example of repressible operon is (corepressor)
the tryptophan (trp) operon. It is usually on
Figure 6.6 The tryptophan operon showing no
(Figure 6.5) and the binding of a repressor to transcription of the structural genes.
the operator shuts off transcription. Modified from Reece et al. (2011).

promoter
promoter

trpE trpD trpC trpB trpA

DNA operator

mRNA RNA polymerase

protein E D C B A

inactive repressor
Figure 6.5 The tryptophan operon showing the transcription of the structural genes. Modified from
Reece et al. (2011).

15
Module 2 - Molecular Genetics
Gene-enzyme relationship
There is a specific connection between genes
and enzymes. For example, the ultimate
product of the metabolic process is affected
by a stepwise succession of enzymes. Beadle
and Tatum used mutants to work out the
details of biochemical pathways e.g.
biosynthesis of niacin (vitamin B3) in pink
bread mold.
Table 6.1 Growth of mutants in response to
supplements.
Mutant
Supplement
A B C
1 + + +
2 - + +
3 - - +
Let us use the data in Table 6.1 to
determine the biochemical pathway. The later
in the pathway a mutant is blocked, the fewer
nutritional supplements must be added to
allow growth. In the data given, the nutritional
supplements are listed in the order in which
they appear in the pathway. Thus, initially, the
pathway is:
Compound A  Compound B  Compound C
Now, we need to use the mutants to
help define the biochemical pathway. Mutant
1 will grow on all compounds. Therefore, the
defective enzyme in mutant 1 must act to
convert a precursor molecule to compound A.
Mutant 2 will on all compounds except
compound A. Thus, the defective enzyme
must act on the conversion of compound A to
compound B. Lastly, mutant 3 will grow only in
16
Lesson 6 - Gene Expression
the presence of compound C. Thus the
defective enzyme produced by the mutant
gene in mutant 3 must act before the
formation compound C and after the
formation of compounds A and B from the
precursor molecule. Thus, the final deduced
pathway, and the positions of the mutant
blocks, are as follows:
Mutant 1 Mutant 2 Mutant 3
Precursor  A  B  C
Gene regulation in eukaryote
Gene regulation in eukaryotes is mainly for
the regulation of genetic program that
underlies embryological development and
tissue differentiation and response to abiotic
and biotic stress.
Specific regulation of expression can
be facilitated by hormones or can be
explained by the Davidson-Britten model. The
model explains that the eukaryotic genome
does not contain simple operons like
prokaryotes. Instead, there is individual
regulation of related genes in the
chromosomes since chromatin contains too
few proteins to function as a repressor.
Another example of gene regulation in
eukaryotes is by hormones e.g. steroid pass
freely through the cell membrane. The
hormone binds to receptors in the cytoplasm
forming the hormone-receptor (H-R’) complex
which can cross the nuclear membrane. This
complex binds to a short stretch of DNA called
a response element (regulatory sequence)
near a gene. The binding modifies the
expression of the gene.
Module 2 - Molecular Genetics Lesson 6 - Gene Expression

Learning check! 5. The following table gives the growth

patterns for single-gene mutant strains
1. Write the three-letter abbreviations for with either growth (+) or no growth (0).
the amino acid residues, in order from N-
terminus to C-terminus, of the polypeptide Mutant Supplements
strains A B C D E
that would be produced in the translation
1 + 0 + 0 0
of the mRNA shown below.
2 + + + + 0
3 + 0 + + 0
4 0 0 + 0 0

Diagram a biochemical pathway that is

2. Determine whether each of the following
conditions would result in the genes of the
lac operon being upregulated or
downregulated.
a. RNA polymerase is blocked from
moving along the lac operon.
b. Allolactose binds to the repressor
protein.
c. Lactose concentration within a cell is
decreased.
d. Allolactose concentration within a cell
is increased.
e. Active repressor proteins are present
in the absence of allolactose.
f. Glucose is available.
consistent with the data, indicating where in
the pathway each mutant strain is blocked.
6. Given several diploids, draw the operon
and determine whether Z and Y gene
products are made in the presence or
absence of an inducer.
a. I- P- Oc Z+ Y+/I+P+ O+ Z- Y-
b. I+ P- O+ Z+ Y+/I- P+ O+ Z+ Y-
c. I+ P+ Oc Z- Y+/I+ P- O+ Z+ Y-
d. Is P+ O+ Z+ Y-/I- P+ Oc Z- Y+

3. The human β-globin polypeptide is 146

amino acids long. How long is the coding
portion of the human -globin mRNA?

4. Write the three-letter abbreviations for

the amino acid residues, in order from N-
terminus to C-terminus, of the polypeptide
that would be produced by the
transcription and translation of DNA with a
sequence of 3’-TACGGGGTACACACT-5’.

17
Module 2 - Molecular Genetics Lesson 7 - Developmental Genetics

Developmental
regulation and cell differentiation which deals

7
with the coordinated expression of genes

Genetics from fertilization to adult formation.

The variable gene activity hypothesis
states that differentiation is accomplished by
activating and inactivating genes at different
In this lesson, you are going to learn about the
times in different cell types. It assumes that
genetic control of development. At the end of
each cell contains an entire genome and that
the lesson you should be able to:
differential transcription of selected genes
1. Discuss the principal cellular
controls the development and differentiation
mechanisms of development and
of each cell.
identify the genetic and molecular
Genetic analysis shows that the size
elements that are involved.
and shape of all animal bodies are controlled
2. Compare and contrast the genetic
by a common set of genes and developmental
developmental control in plants and
mechanisms. Homeotic genes (Hox genes) are
animals.
a single set of genes with different patterns of
3. Define epigenetics and explain the role
expression. Homeotic genes from a wide
of chromatin structure in controlling
range of organisms have a common ancestry.
cellular development and homeostasis.
The evolutionary mechanisms for differential
development of organisms include mutation,
The genetic basis of development gene duplication and divergence, assignment
of new functions to old genes, and
Multicellular organisms share many genes,
recruitment of genes to new developmental
genetic pathways, and molecular signaling
pathways. The integration of evolutionary
mechanisms that control developmental
biology with developmental biology is called
events from zygote to adult.
Evo-Devo.
At the cellular level, development is
A developmental geneticist studies the
marked by three important events such as
regulation and spatio-temporal expression of
specification - first cues confer spatially
genes and the consequences of defective
distinct identity; determination - specific
genes in development. To answer these
developmental fate for cell becomes fixed;
questions, they often use model organisms
and differentiation - cell achieves final adult
such as Drosophila melanogaster (fruit fly),
form and function. Development is a
Caenorhabditis elegans (nematode) and
programmed series of phenotypic changes
Arabidopsis thaliana (flowering plant). The
under temporal, spatial, and quantitative
analysis of developmental mechanisms
control which involves a series of complex
involves an understanding of the general
biochemical pathways that are under genetic
processes of development like how an adult
control. Therefore, developmental genetics is
body plan is laid down in an embryo, program
the study of the relationship between gene

18
Module 2 - Molecular Genetics Lesson 7 - Developmental Genetics

of gene expression turns undifferentiated cells
into differentiated cells and role of cell to cell
communication in development.
Embryonic development of D.
melanogaster
The development of Drosophila has a ten-day
cycle and divided into five distinct phases
before the adult stage such as embryo, three
larval stages, pupal stage, and adult stage.
The early development of Drosophila
starts when the egg is fertilized. After the
tenth division, the nuclei migrate to the
periphery of the egg and the germ cells form.
Nuclei are enclosed in a membrane called
blastoderm and segmentation pattern is
established (Figure 7.1).
There are two different sets of genes
that control the embryonic development of
Drosophila such as maternal-effect genes and
zygotic genes. The mRNA and proteins
produced by the maternal effect genes are
deposited in the egg cytoplasm. The products
are distributed in gradient or concentrated in
specific regions of the cell. Maternal effect
genes also encode transcription factors and
proteins that regulate gene expression by
activation or repression of the zygotic genome.
Gradients of maternal-effect gene products
along the anterior-posterior axis of embryo
regulate expression of segmentation genes
(gap, pair-rule, and segment polarity genes)
and homeotic selector (Hox) genes, which
specify the fate of each segment.

Diploid zygote nucleus is produced by Nuclei become enclosed in membranes,

fusion of parental gamete nuclei forming a single layer of cells over
embryo surface

Nine rounds of nuclear divisions

produce multinucleated syncytium

Approximately four further divisions

take place at the cell periphery

Figure 7.1 The early embryonic developmental stages of D. melanogaster. Source: Klug et al. (2017).
19
Module 2 - Molecular Genetics
Figure 7.2 The early embryonic developmental
stages of D. melanogaster. Source: Klug et
al. (2017).
Maternal-effect gene products are
present in the egg during oogenesis and then
activated immediately after fertilization. These
genes establish the anterior-posterior axis of
the embryo (Figure 7.2a) and encode
transcription factors that activate
transcription of the gap gene. The expression
of the gap gene divides the embryo into the
head, thorax, and abdominal regions of adult
(Figure 7.2b). Gap proteins act as transcription
factors and activate pair-rule genes, whose
products divide embryo into smaller regions
two segments wide (Figure 7.2c). The action
of maternal and segmentation genes is
20
Lesson 7 - Developmental Genetics
defined by the action of homeotic (Hox) genes
(Figure 7.2e). Zygotic genes program the
segment formation of Drosophila. There are
several segmentation genes identified and
classified based on the mutant phenotype
(Figure 7.3). Gap gene mutations delete a
group of adjacent segments and cause gaps in
normal body plan. Pair-rule gene mutations
affect every other segment and eliminate a
specific part of then segment. Segment
polarity gene mutations cause defects in
segment portions.
Homeotic genes (selector genes) are
activated as targets of zygotic genes. Selector
genes determine which adult structures will
be formed by each body segment—antennae,
mouthparts, legs, wings, thorax, and abdomen.
Drosophila genome contains two clusters of
homeotic genes on chromosome 3 which
include antennapedia complex and bithorax
complex.
The antennapedia (ANT-C) complex
contains five genes that specify structures in
the head and the first two thoracic segments.
On the other hand, the bithorax (BX-C)
complex contains three genes that specify
structures in the second thoracic segment, the
entire third thoracic segment and the
abdominal segment (Figure 7.4). Hox genes
contain a 180-bp domain known as the
homeobox. It encodes a DNA-binding
sequence of 60 amino acids called
homeodomain. Expression of Hox genes is
collinear with anterior to the posterior
organization in the embryo. In summary, the
developmental program in Drosophila involves
a cascade of events, one affecting the other.
Module 2 - Molecular Genetics Lesson 7 - Developmental Genetics

Normal larva with affected parts

Gene (shaded blue) Effect of mutation Time of expression

Gap (Krüppel)

Pair-rule (even-skipped)

Segment polarity gene

(gooseberry)

Figure 7.3 Example of some mutants for the segmentation genes. Shown here are the effect of
mutation relative to the normal phenotype and the corresponding time of expression.
Source: Russell (2010).

Figure 7.4 The antennapedia and bithorax
complexes in Drosophila and their
corresponding genes and segment
position. Source: Klug et al. (2017).
Developmental programs in
plants and animals
The developmental patterns evolved
independently in plants and animals.
Multicellular development in animals and
plants follows a body plan or pattern. The
term pattern refers to the spatial arrangement
of different regions of the body. At the cellular
level, the body pattern is due to the
arrangement of cells and their specialization.
Plant pattern formation is studied
using a model organism Arabidopsis thaliana
while there are several model organisms to
study development in animals like D.
melanogaster, Danio rerio, and C. elegans. In
animals, development involves movement of
cells and tissues and the growth is limited to
embryonic and juvenile periods. On the other
hand, in plants, there is a presence of
perpetual embryonic tissues called apical
meristems that are continuously growing.

21
Module 2 - Molecular Genetics Lesson 7 - Developmental Genetics

Therefore, growth is not limited to embryonic
and juvenile periods. The difference in
developmental programs among cells in
multicellular organisms is due to different
patterns of gene expression and not from the
differences in the genome of the cells. All cells
in an individual have the same genes (genomic
equivalence). In plants, differentiated somatic
cells can give rise to whole new plant (Figure
7.5) while animal cells, will not always divide
in culture unless cell division is induced. In
other words, many plant cells are totipotent
i.e. have the ability to differentiate into every
cell type and to produce an entire individual.
Nuclear transplantation experiments
showed that animal cells if grown in culture,
do not have the capacity to develop into a
whole new individual. However, once
differentiated, it can be induced to
dedifferentiate into other types of cells.
Figure 7.6 shows the nuclear transplantation
experiment using two different types of
animal cells from embryo and frog intestine.
The cells from the frog embryo are less
differentiated while the intestinal cells are
fully differentiated. The nucleus from each
type of cell was transplanted to enucleated
frog egg cell and observed if tadpoles will
develop. As expected, most tadpoles
developed on the enucleated cell where the
source of the nucleus is the frog embryo.
In animals, specifically the bilaterians,
researchers have discovered that
development generally proceeds in four
overlapping stages such as (1) formation of
body axes - the first phase of pattern
development; (2) segmentation of the body -
the body is subdivided into segments.

Root of
carrot plant

Transverse
section of
root
“Embryoid”
(somatic embryo)
2-mg develops from
fragments cultured cells

Free cells in suspension

begin to divide Adult
plant

Fragments cultured
in nutrient medium
Plantlets cultured on agar medium
and later transferred to soil

Figure 7.5 Tissue culture technology to show the genomic equivalence in plants. Any type of
plants can be grown to a whole new plant provided that suitable growing conditions
are provided. Source: Reece et al. (2011).

22
Module 2 - Molecular Genetics
In invertebrates, the segments may remain
morphologically distinct, whereas, in
vertebrates, distinct segments are obvious
only in the early stages of development; (3)
determination of structures within the
segments - the segments form, groups of
cells become destined to develop into
particular structures and cell types. This
process, occurs before the structures and cell
types have changed their morphologies; and
(4) cell differentiation - towards the end of
development, cells differentiate into particular
cell types. This results in tissues and organs
with specific morphologies. The formation of a
bilaterian body with a specific pattern is the
result of these four overlapping phases. The
phases themselves are controlled by sets of
genes.
Frog embryo Frog egg cell
Less differentiated
cell
Donor nucleus
transplanted Enucleated egg cell
Most develop
into tadpoles
Figure 7.6 Nuclear transplantation experiment to show the genomic equivalence in animals. Source:
Reece et al. (2011).
Lesson 7 - Developmental Genetics
The morphological patterns of growth
are markedly different between animals and
plants. As described previously, animal
embryos become organized along
anteroposterior, dorsoventral, and left-right
axes, and then they subdivide into segments.
By comparison, the form of plants has two key
features. The first is the root-shoot axis. Most
plant growth occurs via cell division near the
tips of the shoots and the bottoms of the
roots. Second, this growth occurs in a well-
defined radial pattern. Overall, the radial
pattern in which a plant shoot gives off the
buds that produce leaves, flowers, and
branches is an important mechanism that
determines much of the general morphology
of the plant.
Frog tadpole
Fully differentiated
(intestinal cell)
Donor nucleus
transplanted
<2% develop
into tadpoles
23
Module 2 - Molecular Genetics Lesson 7 - Developmental Genetics

Differential gene action and In terms of transcriptional activity,

epigenetic control chromatin can be classified as euchromatin
Regulation of gene expression during which is uncoiled and active and appears
development can occur at any stage in the unstained during interphase, and
DNA-mRNA-protein pathway (Figure 7.7). To heterochromatin which is condensed,
accommodate DNA-protein interactions, genetically inactive and appears stained
replication, and gene expression, chromatin during interphase.
structure must change. By chromatin
remodeling, the compact structure is relaxed
and the regions of DNA are exposed to
regulatory proteins. Chromatin modification such
as histone acetylation and
The twists and turns of DNA superhelix DNA demethylation
encircle histones. Unstructured histone tails
Gene available for
are not packed into folded histone domains transcription
within the nucleosome. Tails are devoid of Transcriptional control
secondary structure and protrude through the
minor groove. Histone tails provide potential Primary transcript

targets along with chromatin fiber for

RNA processing control
chemical modifications such as acetylation,
methylation and phosphorylation.
The enzyme histone acetyltransferase RNA transport control
(HAT) facilitates the addition of the acetyl
group to a positively charged amino group on mRNA in the cytoplasm
the side chain (lysine) which changes net
Degradation of Translational control
charge of the protein by neutralizing positive
mRNA control
charge. This loosens chromatin structure,
thereby promoting the initiation of Polypeptide
transcription Post-translational
The enzyme methyltransferase adds modification control

methyl groups to arginine and lysine residues

in histones while the enzyme kinase adds Degradation of
protein
phosphate groups to hydroxyl groups of Protein activity control
amino acids serine and histidine. The addition
of methyl groups (methylation) can condense Active protein
chromatin while the addition of phosphate
groups (phosphorylation) next to a
Figure 7.7 The major point of controls in gene
methylated amino acid can loosen chromatin.
expression starting from DNA up to
protein. Modified from: Reece et al.
(2011).
24
Module 2 - Molecular Genetics Lesson 7 - Developmental Genetics

The inheritance of traits transmitted Neoplasmic interactions

by mechanisms not directly involving the
Cytoplasmic determinants regulate the
nucleotide sequence is called epigenetic
expression of genes that affect the
inheritance. The epigenome (Figure 7.8)
developmental fate of the cell (Figure 7.9).
includes all the information, other than the
Cells have cytoplasmic memory where
DNA sequence itself, which is heritable during
components encoded by active genes are in
cell division. DNA sequences remain
the cytoplasm or extracellular environment.
essentially unchanged throughout the lifetime
They act back on the genome directly or
of an organism, while the epigenome changes
indirectly. The quantity of cytoplasmic
immensely in response to internal or external
determinants during cell division may vary
environmental cues. Epigenetic “tags” present
between daughter cells. For example, some
on parents’ epigenomes are passed down to
receive more while others receive less. Other
offspring. Evidence is growing for a
types of cytoplasmic determinants can
transgenerational epigenetic inheritance,
provide signals to the neighboring cells and
though mechanisms are not necessarily
affect gene expression via a signal
straightforward.
transduction pathway.

Figure 7.8 The epigenome and the biological processes it regulates.

25
Module 2 - Molecular Genetics
Unfertilized egg
Sperm Nucleus
Fertilization Molecules of two
different cytoplasmic
determinants
Zygote
(fertilized egg)
Mitotic cell division
Two-celled embryo
Figure 7.9 Cytolasmic determinants in egg.
Source: Reece et al. (2011).
As the zygote divides by mitosis, cells
contain different cytoplasmic determinants,
which lead to different gene expression. The
other important source of developmental
information is the environment around the
cell, especially signals from nearby embryonic
cells. In the process called induction, signal
molecules from embryonic cells cause
transcriptional changes in nearby target cells.
Thus, interactions between cells induce
differentiation of specialized cell type.
Genomic imprinting
For most genes, we inherit two functional
copies each from the maternal and paternal
origin. Imprinted genes are different—
depending on the gene, either the maternal
copy or paternal copy is epigenetically
silenced which occurs during egg and sperm
formation.
The environment has a stronger effect
on imprinted genes—only one active copy, no
26
Lesson 7 - Developmental Genetics
backup, so any epigenetic change has a
greater impact on gene expression. Improper
imprinting leads to developmental defects
such as cancer or syndromes e.g. Angelman
syndrome (1 in 10,000) - abnormally silenced
gene on Chromosome 15 from the mother.
Other organisms like insects, mammals,
flowering plants demonstrated imprinting. In
animals, the formation of hybrids with
different appearance has been attributed to
genomic imprinting. For example, lions and
tigers can produce hybrid offspring in captivity
and the offspring is different depending on
whether the tiger or the lion is the mother or
father. A cross between male lion x female
tiger produces a liger (largest of big cats) while
a cross between male tiger x female lion
produces a tigon (about the same size as
parents). Another example is a cross between
a horse x donkey which can produce either a
mule or a hinny.
Learning Check!
1. Genetic mechanisms underlying gene
silencing involve DNA sequence alterations,
while epigenetic mechanisms do not.
What are three epigenetic mechanisms
that can lead to gene silencing? What
features do they share in common, and
how are they different?
2. DNA, histones, promoter-binding proteins,
and enhancer-binding proteins are mixed
together in the following orders:
a. afirst histones and DNA, then
promoter-binding proteins
Module 2 - Molecular Genetics Lesson 7 - Developmental Genetics

b. first histones and promoter-binding the host, it will form eye, brain, and other
proteins, then DNA material characteristic of the head region.
c. first DNA and promoter-binding If the tissue is transplanted to other
proteins, then histones regions of the host, it will form organs and
d. first histones, promoter-binding tissues characteristic of those regions in
proteins, and enhancer-binding normal development. In contrast, in Figure
proteins, then DNA b, if tissue destined to be an eye is excised
from a neurula and transplanted into an
For each case, state whether transcription can
older embryo host to exactly the same
occur. Explain your answers.
places as used for the blastula or gastrula
3. There are several distinct genes that code transplants, in every case the transplanted
for - and -like globin polypeptides. These tissue differentiates into an eye. Explain
α-, β-, γ-, δ-, ε-, ζ–globin genes are these results.
transcriptionally active at specific stages of
development, resulting in the synthesis of
polypeptides that are assembled in
specific combinations to form different
types of hemoglobin. Fill in the following
table, indicating whether the globin gene
in question is sensitive (S) or resistant (R)
to DNase I digestion at each of the
developmental stages listed.

Tissue
Globin
Embryonic Fetal Adult bone
gene
yolk sac spleen marrow
α
β
γ
δ
ε
ζ

4. A piece of tissue is excised from a region

of the late blastula or early gastrula that
would later develop into an eye and is
transplanted to three different regions of
an older embryo host. In Figure a, if the
tissue is transplanted to the head region of

27
Module 2 - Molecular Genetics Lesson 7 - Developmental Genetics

References
Brooker, Robert J. (2012). Genetics : Analysis
& Principles 4th edition. The McGraw-Hill
Companies, Inc.,

Brown, Terry (2012). Introduction to Genetics:

A Molecular Approach. Garland Science,
Taylor & Francis Group, LLC

Griffiths, A. J.F., Wessler, S.R., Caroll, S.B.,

Doebley, J.,. (2012). Introduction to Genetic
Analysis 10th Edition. W. H. Freeman and
Company

Hartwell, L.H., Goldberg, M.L., Fischer, J.A.,

and Hood, L., (2015). Genetics from Genes to
Genomes 5th edition. Mc Graw Hill Companies.

Klug, W.S., Cummings, M.R., Spencer, C.A.,

and Palladino, M.A., (2017). Essentials of
Genetics 9th edition. Pearson Education, Inc.

Pierce, B.A., (2013) Genetics Essentials:

Concepts and Connections. 2nd edition, W.H.
Freeman & Company

Reece JB, Urry, LA, Cain, ML, Wasserman, SA,

Minorsky, PV, Jackson, RB. 2011. Campbell
Biology, 9thedition. Pearson Education, Inc.

Russell, Peter J. (2010). iGenetics: A Molecular

approach 3rd edition. Pearson Education, Inc.,

Snustad, D.P., and Simmons, M.J., (2012).

Principles of Genetics, 6th edition, John Wiley
& Sons, Inc.

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