Unit 3 Transcription in Eukaryotes-I

UNIT 3

TRANSCRIPTION IN
EUKARYOTES – I

Structure
3.1 Introduction 3.3 Inhibitors of Eukaryotic
Transcription and their
Expected Learning
Applications
Outcomes
3.4 Fidelity of Transcription
3.2 Transcription by RNA
and Replication
Polymerase I and III
3.5 Summary
Eukaryotic RNA
Polymerases and its Types 3.6 Terminal Questions
Transcription Cycle 3.7 Answers
RNA Polymerase I Mediated
Transcription

RNA Polymerase III

Mediated Transcription

3.1 INTRODUCTION
Transcription is the first step in gene expression, in which information from a
gene is used to synthesize RNA molecule. The goal of transcription is to make
RNA using DNA (the genetic material) sequence. DNA safely and stably
stores genetic information in the nuclei of cells as a reference book in a library.
mRNA is comparable to a copy from a reference source, because it carries the
same information as present in DNA, which can move to cytosol and be
available for directing protein synthesis. For a protein-coding gene, the mRNA
copy, or transcript, carries the information needed for synthesis of a
polypeptide. Eukaryotic transcripts also need to undergo some processing
steps before translation into proteins. Proteins are the key molecules that give
cells structure and keep them functioning. Not all genes are transcribed all the
time. Instead, transcription is controlled individually for each gene. Cells
carefully regulate transcription, transcribing just the genes the products of
which are physiologically required at a particular moment. 31
 Block 1 Transcription

In the first two units of this block you have learnt about prokaryotic
transcription, and in unit 3 and 4 you will explore eukaryotic transcription. The
first half of this unit is dedicated to eukaryotic RNA polymerases and their role
in transcription. The second half of the unit will explain inhibitors of eukaryotic
transcription. At the end, you will explore the uniqueness of transcription and
its fidelity. Before you proceed further in this unit, watch the YouTube video
link provided: https://youtu.be/Yrwvrrvraq4

Expected Learning Outcomes

After studying this unit, you shall be able to:

❖ list the steps in eukaryotic transcription;

❖ discuss the role of RNA polymerases in transcription;

❖ compare and contrast the three RNA polymerases; and

❖ explain the significance of transcription factors.

3.2 TRANSCRIPTION BY RNA

POLYMERASE I AND III
Transcription uses one of the two DNA strands as a template; this strand is
Transcription is a vital
biological process occurs
called the template strand. The RNA product is complementary to the template
in our everyday lives, and strand, and is identical in sequence to the other DNA strand, called the non-
it's also something our template (or coding) strand, except for the presence of uracil (U) in place of
cells must do, in a more thymine (T). The site on the DNA from which the first RNA nucleotide is
specialized and narrowly transcribed is called the +1site, or the initiation site. Nucleotides present
defined way. In before the initiation site are assigned negative numbers, and said to be
biology, transcription is
present upstream of the transcription start site. Nucleotides that come after the
the process of copying
initiation site are assigned positive numbers and said to be present
out the DNA sequence of
a gene in the similar downstream. If the gene that is transcribed encodes a protein, the RNA
alphabet of RNA. molecule will called mRNA, and will be read to make a protein in a process
called translation. In eukaryotes the initial product of transcription is called a
pre-mRNA, which undergoes extensive processing including splicing
(discussed in Block-II) to produce mature mRNA that can be translated by
ribosomes.

3.2.1 Eukaryotic RNA Polymerases and its Types

In this section you will know the role of eukaryotic RNA polymerases and their
types.

RNA polymerases (RNAP) are crucial enzymes that transcribe DNA into RNA.
Using a DNA template, RNA polymerase always synthesizes a new RNA
strand in the 5' to 3' direction, adding new nucleotide to the 3' end of the
strand through phosphodiester bond if it is correctly base paired with the DNA
template. For instance, if there is a G in DNA template, RNA polymerase will
add a C to the new, growing RNA strand. It can only add the ribonucleotides
(A, U, C, or G) to the 3' end of the strand. To know more about these
polymerases watch this video available at the link given below:
32 https://www.youtube.com/watch?v=QmTf4M4qrbI
Unit 3 Transcription in Eukaryotes-I

Types of RNA Polymerases

RNA polymerase enzymes are essential to life and are found in all organisms
and many viruses (Table 3.1). In bacteria, a single type of RNAP catalyses the
synthesis of both messenger RNA and other noncoding RNAs. In eukaryotes,
there are several types of RNAP that synthesise different types of RNAs. For
example, mRNAs and microRNAs are transcribed by RNAP II, tRNAs are
transcribed by RNAP III, while most of the rRNAs are transcribed by RNAP I.
The archaeal RNAP is
The features of eukaryotic mRNA synthesis are more complex than those of closely related to
prokaryotes. Instead of a single polymerase comprising four subunits, the eukaryotic RNAPII in
eukaryotes have three types of RNA polymerases. Each RNAP is made up of terms of subunit
composition and
10 or more subunits. Each eukaryotic RNA polymerase also requires a distinct
architecture, promoter
set of transcription factors for initiation of transcription. elements and basal
transcription factors
RNA polymerase I is located in the nucleolus, a specialised nuclear required for the initiation
substructure where rRNA is transcribed. RNA polymerase I consists of 14 and elongation phase of
protein subunits, and twelve of its subunits have identical or related transcription. RNAPs of
this class are large and
counterparts with RNA polymerase II (Pol II) and RNA polymerase III (Pol III). sophisticated enzymes
The other two subunits are related to Pol II initiation factors and have that interact in a complex
structural homology with Pol III. The rRNA molecules are considered structural manner with DNA/RNA
scaffolds, substrates
RNAs because they are not translated into protein. The rRNAs are
NTPs and a plethora of
components of the ribosome and are essential for the process of translation. transcription factors –
RNA polymerase I synthesises all the rRNAs except for the 5S rRNA interactions that often
result in an allosteric
molecule.
regulation of RNAP
RNA polymerase II is located in the nucleus and synthesises all protein- activity. The 12 subunits
of RNAP play distinct
coding nuclear pre-mRNAs. Eukaryotic pre-mRNAs undergo extensive roles including RNAP
processing after transcription, but before translation. RNA polymerase II is assembly and stability,
responsible for transcribing the overwhelming majority of eukaryotic genes, catalysis and functional
contacts with exogenous
including all of the protein-encoding genes which ultimately are translated into
factors.
proteins and genes for several types of regulatory RNAs, including microRNAs
(miRNAs) and long-coding RNAs (lncRNAs).

RNA polymerase III is also located in the nucleus. This polymerase

transcribes a variety of structural RNAs that includes the 5S pre-rRNA, pre-
transfer RNAs (pre-tRNAs), and pre-small nuclear RNAs. The tRNAs have a
critical role in translation; they serve as the adaptor molecules between the
mRNA template and the growing polypeptide chain. Small nuclear RNAs have
a variety of functions including splicing of pre-mRNAs. Not all miRNAs are
transcribed by RNA Polymerase II, RNA Polymerase III transcribes some of
them.

RNA polymerase IV synthesises siRNA in plants.

RNA polymerase V synthesises RNAs involved in siRNA-directed

heterochromatin formation in plants.
33
Block 1 Transcription
Table 3.1: Eukaryotic RNA polymerases

Type Present in Transcribes

RNA polymerase I All Large rRNAs

eukaryote
s

RNA polymerase II All Pre-mRNA, some snRNAs,

eukaryote snoRNAs, some miRNAs
s

RNA polymerase III All tRNA, small rRNAs, some

eukaryote snRNAs, some miRNAs
s

RNA polymerase IV Plants Some siRNAs

RNA polymerase V Plants RNA molecules taking part in

heterochromatin formation

SAQ 1
i) The enzyme required for transcription is

a) RNAase

b) DNA polymerase

c) RNA polymerase

d) Restriction enzymes

ii) Which of the following is TRUE for the RNA polymerase activity?

a) DNA dependent DNA synthesis

b) Direct repair

c) DNA dependent RNA synthesis

d) RNA dependent RNA synthesis

Up to now we have studied about different types of RNAP and their roles in
eukaryotic transcription. Now in this section you’ll learn about the actual
working mechanism of eukaryotic RNA polymerases.

3.2.2 Transcription Cycle

Let’s begin this section by understanding the basics of the transcription cycle.

In the process of transcription (by any RNA polymerase), there are three main
34 stages (Fig. 3.1).
Unit 3 Transcription in Eukaryotes-I

Initiation: the assembly of transcription apparatus at the promoter of a gene.

This is achieved with the help of transcription factors, and RNA polymerase is
an important component of the transcription apparatus.

Elongation: Sequential addition of ribonucleotides to the nascent RNA with

phosphodiester bonds as directed by the DNA template.

Termination: The cessation of polymerization, the disassembly of the

transcription apparatus, and release of transcript (the newly synthesized
RNA).

Fig. 3.1: Schematic representation of transcription cycle.

Transcription proceeds in the following steps

1. RNA polymerase, together with one or more general transcription

factors, binds DNA at the promoter region of a gene.

2. RNA polymerase creates a transcription bubble by local DNA strand

separation. This is done by breaking the hydrogen bonds of the double
stranded DNA.

3. RNA polymerase adds ribonucleotides (which are complementary to the

nucleotides of template DNA strand) at the growing end of RNA.

4. Hydrogen bonds of the RNA–DNA helix break, releasing the newly

synthesised RNA strand.

5. RNA may undergo further processing that may include cleavage and
polyadenylation, capping, splicing, base modification etc.

6. The RNA may remain in the nucleus or exit to the cytoplasm through the
nuclear pore complex.
35
Block 1 Transcription

Transcription factors for RNA pol I

a) Core promoter covers the start site of transcription, plus an upstream
control element located about 70 bp upstream.
b) The upstream binding transcription factor (UBF1) binds to a GC-rich
sequence in both the upstream control element and in the core
promoter.
c) A multi-subunit complex called selective factor (SL1) binds to the UBF1-
DNA complex, again at both the upstream and core elements.
d) One of the subunits of SL1 is Tata box Binding Protein (TBP). RNA
polymerase I then binds to this complex of DNA+UBF1+SL1 to initiate
transcription at the correct nucleotide and elongate to make pre‑rRNA.

Transcription factors for RNA Pol III

a) TFIIIA: binds to the internal control region of genes that encode 5S RNA
(type 1 internal promoter)

b) TFIIIC: binds to internal control regions of genes for 5S RNA (alongside

TFIIIA) and for tRNAs (type 2 internal promoters)

c) TFIIIB: The binding of TFIIIC directs TFIIIB to bind to sequences (-40 to

+11) that overlap the start site for transcription. One subunit of TFIIIB is
transcription binding protein (TBP), even though no TATA box is
required for transcription.

d) TFIIIA and TFIIIC can be removed without affecting the ability of RNA
polymerase III to initiate transcription. Thus TFIIIA and TFIIIC are
assembly factors, and TFIIIB is the initiation factor.

3.2.3 RNA Polymerase I Mediated Transcription

Eukaryotic RNA polymerase I (Pol I) is specialised in higher eukaryotes, that
only transcribes ribosomal RNA (rRNA). It is important to note that rRNA is the
most abundant cellular RNA. High level rRNA production relies on efficient
binding of initiation factors to the rRNA gene promoter and recruitment of Pol I
complexes containing initiation factor Rrn3. In mammals, the 47S pre-rRNA
produced by Pol I is processed into mature 18S, 5.8S and 28S rRNAs, which
are essential structural and catalytic components of ribosomes. Ribosomes
constitute the core of the protein synthesis machinery; consequently, ribosome
content is a critical determinant of protein accumulation and hence cell growth
and division. The abundance of ribosomes within a cell depends upon the
availability of rRNA, therefore, given the high metabolic burden of rRNA
production and its direct influence on protein synthesis capacity, a tightly
regulated transcription system has evolved, which ensures that rRNA
synthesis is closely coupled with cellular growth demands. You are advised to
watch the video link provided
36 https://www.youtube.com/watch?v=KqKO8DzL9Fk
Unit 3 Transcription in Eukaryotes-I
Initiation:

RNA polymerase I requires two ancillary factors for initiation:

 The factor, UBF, binds in a sequence-specific manner to GC rich

sequences in the core promoter and UPE

 The second factor, SL1 which contains TBP and 3 other proteins, does
not, by itself, have the specificity for the promoter, but once UBF has
bound, SL1 can bind cooperatively to extend the region of DNA that is
covered.

 Once both these factors are bound, RNA polymerase can bind to the
core promoter to initiate transcription.

 UBF is a single protein that binds specifically to the GC rich element in

the core promoter and UPE.

 UBF and RNA pol I from mouse can recognize the human genes, i.e.
UBF and RNA pol I can act on heterologous templates.

 The factor SL1 from mouse cannot initiate transcription on human

templates.

 It means that SL1 confers species specificity on the initiation reaction.

 SL1 consists for 4 proteins

 One of them is TBP, which is also required for initiation by RNA pol II
and III.

Elongation

As Pol I moves from the promoter region, UBF and SL1 remain-bound to
promoter and ready to recruit another Pol I. While elongation proceeds
unimpeded in vitro, it is unclear at this point whether this process happens as
smoothly in a cell where nucleosome structures are present. Pol I do seem to
transcribe through nucleosomes, either bypassing or disrupting them, perhaps
assisted by chromatin-remodelling activities. In addition, UBF might also act as
positive feedback, enhancing Pol I elongation through an anti-repressor
function. An additional factor, TIF-IC, can also stimulate the overall rate of
transcription by suppressing pausing of Pol I. As Pol I continues elongation,
the status of DNA supercoiling is changed in the region of DNA that is being
transcribed. Positive supercoils are accumulated (overwinding of DNA double
strand) ahead of the transcription bubble, and accumulation of negative
supercoils (underwiniding of double stranded DNA) behind the transcription
bubble takes place, which is negotiated by topoisomerases.

Termination

In higher eukaryotes, transcription termination factor (TTF-I) binds and bends

the termination site at the 3' end of the transcribed region. This will force Pol I
to pause. TTF-I, with the help of transcript-release factor (PTRF) and a T-rich
region, will induce Pol I into terminating transcription and dissociating from the
DNA and the new transcript. Evidence suggests that termination might be rate-
limiting in cases of high rRNA production. TTF-I and PTRF will then indirectly
stimulate the reinitiation of transcription by Pol I at the same ribosomal DNA
gene (Fig. 3.2).
37
Block 1 Transcription

Fig. 3.2: Transcription by RNA Polymerase I-rRNA synthesis.

3.2.4 RNA Polymerase III Mediated Transcription

In eukaryotic cells, RNA polymerase III (also called Pol III) transcribes DNA to
synthesise ribosomal 5S RNA, tRNA and other small RNAs. The genes
transcribed by RNA Pol III fall in the category of "housekeeping genes”
which are expressed in all cell types and under most environmental conditions.
Therefore, the regulation of Pol III transcription is closely linked to the
regulation of cell growth and the cell cycle, thus requiring fewer regulatory
proteins than RNA polymerase II.

Initiation: Assembly of the transcription apparatus at promoter. Pol III is

unusual (compared to Pol II) as it requires no control sequences upstream of
the gene, instead it normally relies on internal control sequences - sequences
within the transcribed section of the gene (although upstream sequences are
occasionally seen, e.g. snRNA gene which also has an upstream TATA box as
seen in Pol II Promoters).

There are three classes of Pol III initiation, corresponding to 5S rRNA, tRNA,
and snRNA initiation. In all cases, the process starts with transcription factors
binding to control sequences, and ends with TFIIIB (Transcription Factor for
polymerase III B) being recruited to the complex and assembling Pol III. TFIIIB
consists of three subunits: TATA binding protein (TBP), a TFIIB-related factor
(BRF1, or BRF2 for transcription of a subset of Pol III-transcribed genes in
vertebrates), and a B-double-prime (BDP1) unit. The overall architecture bears
similarities to that of Pol II.

Class I

Typical stages in 5S RNA (also termed class I) gene initiation:

TFIIIA (Transcription Factor for polymerase III A) binds to the intragenic (lying
within the transcribed DNA sequence) 5S RNA control sequence, the C Block
(also termed box C).
38
Unit 3 Transcription in Eukaryotes-I

TFIIIA serves as a platform that replaces the A and B Blocks for positioning
TFIIIC in an orientation with respect to the start site of transcription that is
equivalent to what is observed for tRNA genes.

Once TFIIIC is bound to the TFIIIA-DNA complex, the assembly of TFIIIB

proceeds as described for tRNA transcription.

Class II

Typical stages in a tRNA (also termed class II) gene initiation:

TFIIIC (Transcription Factor for polymerase III C) binds to two internal (lying
within the transcribed DNA sequence) control sequences, the A and B Blocks
(also termed box A and box B).

TFIIIC acts as an assembly factor that positions TFIIIB to bind to DNA at a site
centered approximately 26 base pairs upstream of the start site of
transcription.

TFIIIB is the transcription factor that assembles Pol III at the start site of
transcription. Once TFIIIB is bound to DNA, TFIIIC is no longer required.
TFIIIB also plays an essential role in DNA strand separation

Class III

Typical stages in a snRNA (also termed class III) gene initiation (documented
in vertebrates only):

SNAPc (snRNA Activating Protein complex; subunits: 1, 2, 3, 4, 5) (also

termed PBP and PTF) binds to the PSE (Proximal Sequence Element)
centered approximately 55 base pairs upstream of the start site of
transcription. This assembly is greatly stimulated by the Pol II transcription
factors Oct1 and STAF that bind to an enhancer-like DSE (Distal Sequence
Element) at least 200 base pairs upstream of the start site of transcription.
These factors and promoter elements are shared between Pol II and Pol III
transcription of snRNA genes.

SNAPc acts to assemble TFIIIB at a TATA box centered 26 base pairs

upstream of the start site of transcription. It is the presence of a TATA box that
specifies that the snRNA gene is transcribed by Pol III rather than Pol II.

The TFIIIB for snRNA transcription contains a smaller Brf1 paralogue, Brf2.

TFIIIB is the transcription factor that assembles Pol III at the start site of
transcription. Sequence conservation predicts that TFIIIB containing Brf2 also
plays a role in promoter opening (Fig. 3.3).

Elongation

TFIIIB remains bound to DNA following initiation of transcription by Pol III. This
leads to a high rate of transcriptional reinitiation of Pol III-transcribed genes.

Termination

Polymerase III terminates transcription at small polyTs stretch (5-6). In

eukaryotes, a hairpin loop is not required, as it is in prokaryotes. 39
Block 1 Transcription

Fig. 3.3: Transcription by RNA Polymerase III-tRNA, 5SrRNA and stable RNAs.

SAQ 2
RNA Polymerase-I is involved in the transcription of .....................

a) t RNA

b) r RNA

c) m RNA

d) ALL types of RNAs

3.3 INHIBITORS OF EUKARYOTIC

TRANSCRIPTION AND THEIR
APPLICATIONS
So far you have studied the mechanism of action of Pol-I, II and III enzymes.
Now in this section you’ll explore various eukaryotic transcription inhibitors and
their importance.

Many anticancer drugs inhibit transcription, and most transcription inhibitors

have useful pharmacological properties. Many laboratory experiments also
utilize transcription inhibitors. Among the various drugs available to inhibit
transcription, each has its advantages and drawbacks. Selectivity, efficiency,
rapidity of action and reversibility are the key issues.

The classical inhibitors of transcription have advantages and drawbacks.

Amanitin is highly selective for RNAP II and RNAP III but slow, actinomycin
40
Unit 3 Transcription in Eukaryotes-I

D is fast but its selectivity is poor, CDK9 inhibitors are fast and reversible. New
inhibitors such as triptolide are fast, selective and completely arrest
ranscription as they trigger rapid degradation of RNAP II.

Amanitin actinomycin D

Triptolide

SAQ 3
Mark the one, which is NOT the transcription inhibitor in eukaryotes.

a) Rifampicin
b) Acridine dye
c) Actinomycin D
d) Rho factor

3.4 FIDELITY OF TRANSCRIPTION AND

REPLICATION
The nucleic acid is present in the nucleus of a cell in the form of either DNA or
RNA (RNA in case of the only retrovirus) which is genetic material in living
organisms. A polymerase is the enzyme that synthesises nucleic acids. By
now you are aware that DNA polymerase is an enzyme that synthesises DNA
(replication), while the RNA polymerase is an enzyme that synthesises the
RNA (transcription). DNA polymerase synthesises double-stranded DNA while
the RNA polymerase synthesises a single-stranded RNA. DNA polymerase
always requires a short-single stranded DNA/RNA molecule- called primer for 41
Block 1 Transcription

initiating the synthesis, which is not required for RNA polymerase. The DNA
polymerase adds dATP, dGTP, dCTP and dTTP to the growing DNA strand,
while the RNA polymerase inserts to the growing RNA strand.

Though the function of both polymerases is to synthesise nucleic acid but they
function differently. The DNA polymerase has polymerization as well as
proofreading activity while the RNA polymerase only has the polymerization
activity. DNA polymerase inserts nucleotides and also repairs the mismatched
pairing by its proof-reading activity.

The proofreading is achieved by exonuclease activity of DNA polymerase

which excises the wrong nucleotides so that the correct ones could be
incorporated in DNA. The exonuclease domain of DNA polymerase removes
the mismatch and the polymerization domain inserts new nucleotides in place
of mismatch. On the other hand, the RNA polymerase does not have the
exonuclease activity, thus it cannot repair the mismatch. Because of this
reason, the error rate of the DNA polymerase is much lower than the
RNA polymerase.

3.5 SUMMARY
● Transcription in eukaryotes involves one of three types of polymerases,
depending on the gene being transcribed. RNA polymerase II
transcribes all of the protein-coding genes, whereas RNA polymerase I
transcribes rRNA genes, and RNA polymerase III transcribes rRNA,
tRNA, and small nuclear RNA genes.

● The initiation of transcription in eukaryotes involves the binding of

several transcription factors to complex promoter sequences that are
usually located upstream of the gene being transcribed.

● All RNA molecules are synthesised in 5' to 3' direction.

● The mechanism of termination of different types of eukaryotic RNA

polymerases are different.

● The termination of eukaryotic RNA polymerases usually takes place

further downstream of the 3’ end of mature RNAs. The extra RNA
sequence present at the 3’ end of primary transcript is removed during
RNA processing steps. The details of processing reactions for mRNA,
rRNA and tRNA molecules are different.

3.6 TERMINAL QUESTIONS

1. List the steps in eukaryotic transcription.

2. Define and discuss the role of RNA polymerases in transcription.

3. Compare and contrast the three eukaryotic RNA polymerases

4. Explain the significance of transcription factors

42
Unit 3 Transcription in Eukaryotes-I
5. Transcription is the transfer of genetic information from

a) DNA to mRNA

b) RNA to protein

c) mRNA to tRNA

d) tRNA to mRNA

6. A DNA sequence is transcribed by an RNA polymerase to produce

complementary RNA strand known as

a) Negative transcript

b) Secondary transcript

c) Primary transcript

d) Tertiary transcript

7. In eukaryotes, in order to initiate transcription

a) RNA strand must be present

b) RNA polymerase must be present

c) Core promoter sequence must be present

d) None of these

3.7 ANSWERS
Self Assessment Questions
1. i) c)

ii) c)

2. b)

3. d)

Terminal Questions
1. Eukaryotic transcription is carried out in the nucleus of the cell by one of
three RNA polymerases, depending on the RNA being transcribed, and
proceeds in three sequential stages:

a) Initiation

b) Elongation

c) Termination. Refer section 3.2.2

2. RNA polymerase is an enzyme that is responsible for copying a DNA

sequence into an RNA sequence, during the process of transcription. As
complex molecule composed of protein subunits, RNA polymerase
controls the process of transcription, during which the information stored
in a molecule of DNA is copied into a new molecule of messenger 43
Block 1 Transcription

RNA.RNA polymerases have been found in all species, but the number
and composition of these proteins vary across taxa. For instance,
bacteria contain a single type of RNA polymerase, while eukaryotes
(multicellular organisms and yeasts) contain three distinct types. Despite
these differences, there are striking similarities among transcriptional
mechanisms. For example, all species require a mechanism by which
transcription can be regulated in order to achieve spatial and temporal
changes in gene expression. Refer section 3.2

3.
Type Present in Transcribes

RNA polymerase I All Large rRNAs

eukaryotes

RNA polymerase II All Pre-mRNA, some snRNAs,

eukaryotes snoRNAs, some miRNAs

RNA polymerase III All tRNA, small rRNAs, some

eukaryotes snRNAs, some miRNAs

RNA polymerase IV Plants Some siRNAs

RNA polymerase V Plants RNA molecules taking part

in heterochromatin
formation
Refer section 3.2.1 and Table 3.1
4. Transcription factors are proteins involved in the process of converting,
or transcribing, DNA into RNA. Transcription factors include a wide
number of proteins, excluding RNA polymerase, that initiate and regulate
the transcription of genes. One distinct feature of transcription factors is
that they have DNA-binding domains that give them the ability to bind to
specific sequences of DNA called enhancer or promoter sequences.
Some transcription factors bind to a DNA promoter sequence near the
transcription start site and help form the transcription initiation complex.
Other transcription factors bind to regulatory sequences, such as
enhancer sequences, and can either stimulate or repress transcription of
the related gene. These regulatory sequences can be thousands of base
pairs upstream or downstream from the gene being transcribed.
Regulation of transcription is the most common form of gene control.
The action of transcription factors allows for unique expression of each
gene in different cell types and during development. Refer the boxes
provided in section 3.2.2

5. a)

6. c)

7. b) & c)

44