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Prokaryotic transcriptionFrom Wikipedia, the free encyclopedia
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bechallengedandremoved.(October 2011)
Prokaryotic transcription is the process in whichmessenger RNAtranscripts of genetic material
inprokaryotesare produced, to be translated for the production ofproteins. Prokaryotic transcription occurs
in thecytoplasmalongsidetranslation. Unlike ineukaryotes, prokaryotic transcription and translation can
occur simultaneously. This is impossible in eukaryotes, where transcription occurs in a membrane-bound
nucleus while translation occurs outside the nucleus in the cytoplasm. In prokaryotes genetic material is not
enclosed in a membrane-enclosed nucleus and has access toribosomesin the cytoplasm.[1]
Transcription is known to be controlled by a variety of regulators in prokaryotes. Many of these transcription
factors are homodimers containing helix-turn-helix DNA-binding motifs.[2]
Contents
[hide]
1 Initiation
2 Elongation
3 Termination
4 References
5 External links
[edit]Initiation
The following steps occur, in order, for transcription initiation:
RNA polymerase (RNAP) binds to one of several specificity factors,, to form aholoenzyme. In this
form, it can recognize and bind to specificpromoterregions in the DNA. The -35 region and the -10
("Pribnow box") region comprise the basic prokaryoticpromoter, and |T| stands for theterminator. The
DNA on the template strand between the +1 site and the terminator is transcribed into RNA, which is
thentranslatedintoprotein.At this stage, the DNA is double-stranded ("closed"). This
holoenzyme/wound-DNA structure is referred to as the closed complex.
The DNA is unwound and becomes single-stranded ("open") in the vicinity of the initiation site (defined
as +1). This holoenzyme/unwound-DNA structure is called the open complex.
The RNA polymerase transcribes the DNA (the beta subunit initiates the synthesis), but produces
about 10 abortive (short, non-productive) transcripts which are unable to leave the RNA polymerase
because the exit channel is blocked by the -factor. The -factor eventually dissociates from the holoenzyme, and elongation proceeds.
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[edit]Elongation
Promoters can differ in "strength"; that is, how actively they promote transcription of their adjacent DNA
sequence. Promoter strength is in many (but not all) cases, a matter of how tightly RNA polymerase and its
associated accessory proteins bind to their respective DNA sequences. The more similar the sequences
are to aconsensus sequence, the stronger the binding is. Additional transcription regulation comes
fromtranscription factorsthat can affect the stability of the holoenzyme structure at initiation.
Most transcripts originate using adenosine-5'-triphosphate (ATP) and, to a lesser extent, guanosine-5'-
triphosphate (GTP)(purinenucleoside triphosphates) at the +1 site. Uridine-5'-triphosphate (UTP) and
cytidine-5'-triphosphate (CTP)(pyrimidinenucleoside triphosphates) are disfavoured at the initiation site.
[edit]Termination
Two termination mechanisms are well known:
Intrinsic termination (also calledRho-independent transcription termination) involves terminator
sequences within the RNA that signal the RNA polymerase to stop. The terminator sequence is usually
apalindromicsequence that forms a stem-loophairpinstructure that leads to the dissociation of the
RNAP from the DNA template.
Rho-dependent termination uses atermination factorcalled factor(rho factor) which is a protein to
stop RNA synthesis at specific sites. This protein binds at a rho utilisation site on the nascent RNA
strand and runs along the mRNA towards the RNAP. A stem loop structure upstream of the terminator
region pauses the RNAP, when -factor reaches the RNAP, it causes RNAP to dissociate from the
DNA, terminating transcription.
[edit]References
o fully understand transcription, students need to appreciate the dynamic nature of this process. Transcription"works" because nucleotides diffuse into position from the cytoplasm, the DNA helix unwinds and then rewinds,
and the RNA polymerase travels rapidly along a DNA template strand for hundreds or thousands of bases toproduce a very long RNA molecule in its wake. This simple, 40-s animation is presented at three levels ofmagnification to emphasize the movements of all the players.
At the beginning of the animation, an RNA polymerase (RNAP) is shown binding to the promoter region of adouble helix of DNA. The sigma subunit of RNAP helps to locate the promoter, after which it is no longerneeded.
Then, as we look "inside" the RNAP, we can see that the double helix begins to unwind. A growing strand ofRNA (in red) begins to form along one of the DNA strands. As the strand lengthens, the RNAP continues totravel along the DNA, unwinding it to expose more DNA bases. Behind the RNAP the DNA helix reforms,displacing the RNA strand.
Finally, we zoom in further to the active site of the RNAP molecule. Here, we can see how an RNA nucleotidetriphosphate (in this case, an "A," for a ribose triphosphate with an adenine base) diffuses into position andbinds to its complement a "T" on the DNA strand (note that the blue deoxyriboses on the DNA strand have
the opposite orientation to the red ribose residues on the the RNA strand). Once hydrogen bonding haspositioned the "A" and the "T" together, the RNAP is able to catalyze covalent bonding of the "A" to the 3-OHon the growing RNA strand. This bond formation is powered by the cleavage of the diphosphate from the RNA
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nucleotide. The process is quickly repeated as another RNA nucleotide triphosphate, "C," is added to the RNAchain. Moments later, a DNA nucleotidea "T"diffuses into position. While this nucleotide can hydrogen bondwith the "A" on the DNA template strand, the RNAP discriminates between RNA and DNA nucleotides, and noreaction is catalyzed. Ultimately, the deoxyribonucleotide diffuses away, allowing RNA strand synthesis tocontinue when another ribonucleotide triphosphatea "U"hydrogen bonds with the A on the DNA templatestrand.
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Concept 2: Transcription and Translation in CellsIn a prokaryotic cell, transcription and translation are coupled; that is, translation
begins while the mRNA is still being synthesized. In a eukaryotic cell, transcription
occurs in the nucleus, and translation occurs in the cytoplasm.Prokaryotic Cell
Because there is no nucleus to separate the processes of transcription and translation, when
bacterial genes are transcribed, their transcripts can immediately be translated.Eukaryotic Cell
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Transcription and translation are spatially and temporally separated in eukaryotic cells; that is,transcription occurs in the nucleus to produce a pre-mRNA molecule.
The pre-mRNA is typically processed to produce the mature mRNA, which exits the nucleus and istranslated in the cytoplasm.
Concept 3: Different Genes for Different RNAsThere are 4 types of RNA, each encoded by its own type of gene.The genomic DNA contains all the information for the structure and function of an organism.
In any cell, only some of the genes are expressed, that is, transcribed into RNA.
There are 4 types of RNA, each encoded by its own type of gene: mRNA - Messenger RNA: Encodes amino acid sequence of a polypeptide.
tRNA - Transfer RNA: Brings amino acids to ribosomes during translation.
rRNA - Ribosomal RNA: With ribosomal proteins, makes up the ribosomes, the organelles thattranslate the mRNA.
snRNA - Small nuclear RNA: With proteins, forms complexes that are used in RNA processing ineukaryotes. (Not found in prokaryotes.)
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Concept 4: Basic Structure of a Protein-Coding Gene
A protein-coding gene consists of a promoter followed by the coding sequence for the
protein and then a terminator.
The promoter is a base-pair sequence that specifies where transcription begins.
The coding sequence is a base-pair sequence that includes coding information for the polypeptidechain specified by the gene.
The terminator is a sequence that specifies the end of the mRNA transcript.
Concept 6: The Transcription ProcessRNA synthesis involves separation of the DNA strands and synthesis of an RNA moleculein the 5' to 3' direction by RNA polymerase, using one of the DNA strands as a template. In complementary base pairing, A, T, G, and C on the template DNA strand specify U, A, C, and G,respectively, on the RNA strand being synthesized.
Concept 7:
Complete Transcription of an RNA Molecule
Transcription begins at the promoter, proceeds through the coding region, and ends at
the terminator.
Concept 8: mRNA in Prokaryotes
The sequence of a prokaryotic protein-coding gene is colinear with the translated mRNA;
that is, the transcript of the gene is the molecule that is translated into the polypeptide.
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mRNA in Eukaryotes
The sequence of a eukaryotic protein-coding gene is typically not colinear with the
translated mRNA; that is, the transcript of the gene is a molecule that must be
processed to remove extra sequences (introns) before it is translated into thepolypeptide.
Most eukaryotic protein-coding genes contain segments called introns, which break up the aminoacid coding sequence into segments called exons.
The transcript of these genes is the pre-mRNA (precursor-mRNA).
The pre-mRNA is processed in the nucleus to remove the introns and splice the exons together
into a translatable mRNA. That mRNA exits the nucleus and is translated in the cytoplasm.
Pre-mRNA Processing (Splicing)
Eukaryotic pre-mRNAs typically include introns. Introns are removed by RNA processing
in which the intron is looped out and cut away from the exons by snRNPs, and the exons
are spliced together to produce the translatable mRNA.
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The steps of pre-mRNA splicing (intron removal) are as follows: The intron loops out as snRNPs (small nuclear ribonucleoprotein particles, complexes of snRNAsand proteins) bind to form the spliceosome.
The intron is excised, and the exons are then spliced together.
The resulting mature mRNA may then exit the nucleus and be translated in the cytoplasm.
Transcription in ProkaryotesThe most detailed molecular information about the transcription cycle is available in
bacterial systems. The synthesis of RNA is initiated at the promoter sequence by the
enzyme RNA polymerase. A single RNA polymerase type is responsible for the
synthesis of messenger, transfer, and ribosomal RNAs.
When isolated from bacteria, prokaryotic RNA polymerase has two forms:
The coreenzyme and the holoenzyme. The core enzyme is a tetramer whose
composition is given as 2 (two alpha subunits, one beta subunit, and one beta-
prime subunit). Core RNA polymerase is capable of faithfully copying DNA into RNA
but does not initiate at the correct site in a gene. That is, it does not recognize the
promoter specifically. Correct promoter recognition is the function of the holoenzyme
form of RNA polymerase.
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Figure 1
The RNA polymerase holoenzyme contains another subunit, s( sigma), in addition to
the subunits found in the core enzyme. Holoenzyme, 2, is capable of correct
initiation at the promoter region of a gene. Sigma thus must be involved in promoter
recognition. Sigma subunits are related but distinct in different forms of RNA
polymerase holoenzyme. These specialized subunits direct RNA polymerase to
promoter sequences for different classes of genes. For example, bacteria exposed to
high temperatures synthesize a set of protective proteins called heat-shock
proteins. The genes for the heat-shock proteins have special promoter sequences
that are recognized by an RNA polymerase holoenzyme with a specific subunit.
The discussed here is the major of the common bacteriumE. coli, about which
most is known.
Promoter recognitionRNA polymerase holoenzyme starts by recognizing the promoter of a gene. The
promoter isn't copied into RNA, but it is, nonetheless, an important piece of genetic
information. The information in a promoter was determined by lining up a large
number of promoters and counting how many times a particular base appeared at a
given position in the various promoter sequences. The consensus sequence is
given by the statistically most probable base at each pointthe bases that appear
most often in the promoter collection. Very few, if any, naturally occurring
promoters match the consensus sequence exactly, but the strength of a promoter
(how actively RNA polymerase initiates at it) correlates well with the degree of
consensus match. For example, the promoters of genes for ribosomal RNA match
the consensus well, while the promoters for the mRNA encoding some regulatory
proteins match the consensus poorly. This correlates with the relative amounts of
each gene product that are needed at any one time: many ribosomes, and only a
few regulatory proteins.
The consensus sequence for an E. colipromoter has two conserved regions near
positions -35 and -10 relative to the transcription start site. That is, the template-
directed synthesis of RNA begins 35 base pairs downstream of the first consensus
region and ten base pairs downstream of the second. The -35 consensus is:
TTG ACA.
The -10 consensus is:
TATAA T.
A couple of important points exist about the consensus. First, not all bases in the
consensus are conserved to the same amount. The bases marked with bold type and
underlined are more conserved than the others, and the -10 region is more
conserved overall than is the -35 region. Secondly, the promoter sequence is
asymmetrical; that is, it reads differently in one direction than in the other.
(Compare this to the recognition sequence for the restriction enzyme BamHI,
GGATCC.) This asymmetry means that RNA polymerase gets directional
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Figure 3
Elongation is the function of the RNA polymerase core enzyme. RNA polymerase
moves along the template, locally unzipping the DNA double helix. This allows a
transient base pairing between the incoming nucleotide and newly-synthesized RNA
and the DNA template strand. As it is made, the RNA transcript forms secondary
structure through intra-strand base pairing. The average speed of transcription is
about 40 nucleotides per second, much slower than DNA polymerase. Other protein
factors may bind to polymerase and alter the rate of transcription and some specific
sequences are transcribed more slowly than others are. Eventually, RNA polymerasemust come to the end of the region to be transcribed.
Termination of transcription in vitro is classified as to its dependence on the protein
factor, rho (). Rho-independent terminators have a characteristic structure, which
features (a) A strong G-C rich stem and loop, (b) a sequence of 46 U residues in
the RNA, which are transcribed from a corresponding stretch of As in the template.
Rho-factor-dependent terminators are less well defined, as shown in Figure4.
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Ribosomal RNA
Ribosomal RNA is essential for protein synthesis. In fact, RNA is thought to be the
catalytically active part of the very large complex of proteins and RNAs that
synthesize proteins. Ribosomes and ribosomal RNAs are heterogeneous, withdifferent sized rRNAs found in the small and large subunits of the ribosome.
Ribosomes can be separated into two subunits. Each subunit contains both protein
and RNA. Although they vary widely in size, ribosomal RNAs have common
secondary structures. The larger size of the eukaryotic RNAs is due to their having
extra structural domains inserted into the midst of the smaller ones, rather than by
a totally new folding pattern.
Antibiotics are natural products, usually from soil bacteria and molds, which interfere
with the growth of other bacteria. Often these antibiotics act on ribosomal RNA
targets. For example, streptomycin, which has been used to treat tuberculosis, bindsto a single region of bacterial 16S RNA, interfering with protein synthesis. The drug
doesn't disrupt protein synthesis in humans, which allows for streptomycin's
relatively high therapeutic indexthe ratio of harmful to helpful doses of the drug.
Conversely, bacteria can become resistant to antibiotics by changes in their rRNA,
either by a change in the nucleotide sequence of the ribosomal RNA or by
methylation of the rRNA.
Transcription (genetics)From Wikipedia, the free encyclopedia
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Transcription is the process of creating acomplementaryRNAcopy of a sequence ofDNA.[1]
Both RNA
and DNA arenucleic acids, which usebase pairsofnucleotidesas acomplementarylanguage that can be
converted back and forth from DNA to RNA by the action of the correctenzymes. During transcription, a
DNA sequence is read by anRNA polymerase, which produces a complementary,antiparallelRNA strand.
As opposed toDNA replication, transcription results in an RNA complement that includesuracil(U) in all
instances wherethymine(T) would have occurred in a DNA complement. Also unlikeDNA
replicationwhere DNA is synthesised, transcription does not involve anRNA primerto initiate RNA
synthesis.
Transcription is explained easily in 4 or 5 steps, each moving like a wave along the DNA.
1. RNA polymerase moves the transcription bubble, a stretch of unpaired nucleotides, by breaking
the hydrogen bonds between complementary nucleotides.
2. RNA polymerase adds matching RNA nucleotides that are paired with complementary DNA bases.
3. RNA sugar-phosphate backbone forms with assistance from RNA polymerase.
4. Hydrogen bonds of the untwisted RNA + DNA helix break, freeing the newly synthesized RNA
strand.
5. If the cell has anucleus, the RNA is further processed (addition of a 3' poly-A tail and a 5' cap) and
exits through to the cytoplasm through thenuclear porecomplex.
Transcription is the first step leading togene expression. The stretch of DNA transcribed into an RNA
molecule is called a transcription unitand encodes at least onegene. If the gene transcribed encodes
aprotein, the result of transcription ismessenger RNA(mRNA), which will then be used to create that
protein via the process oftranslation. Alternatively, the transcribed gene may encode for eithernon-coding
RNAgenes (such asmicroRNA,lincRNA, etc.) orribosomal RNA(rRNA) ortransfer RNA(tRNA), other
components of the protein-assembly process, or otherribozymes.[2]
A DNA transcription unit encoding for a protein contains not only the sequence that will eventually be
directly translated into the protein (the coding sequence) but also regulatory sequencesthat direct and
regulate the synthesis of that protein. The regulatory sequence before (upstreamfrom) the coding
sequence is called thefive prime untranslated region(5'UTR), and the sequence following
(downstreamfrom) the coding sequence is called thethree prime untranslated region(3'UTR).[2]
Transcription has some proofreading mechanisms, but they are fewer and less effective than the controls
for copying DNA; therefore, transcription has a lower copying fidelity than DNA replication .[3]
As in DNA replication, DNA is read from 3' 5' during transcription. Meanwhile, the complementary RNA
is created from the 5' 3' direction. This means its 5' end is created first in base pairing. Although DNA is
arranged as two antiparallel strands in adouble helix, only one of the two DNA strands, called the template
strand, is used for transcription. This is because RNA is only single-stranded, as opposed to double-
stranded DNA. The other DNA strand is called the coding (lagging) strand, because its sequence is the
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n.wikipedia.org/wiki/Transcription_(genetics)#cite_note-Stryer_2006-2http://en.wikipedia.org/wiki/Double_helixhttp://en.wikipedia.org/wiki/Double_helixhttp://en.wikipedia.org/wiki/Double_helixhttp://en.wikipedia.org/wiki/Double_helixhttp://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-Stryer_2006-2http://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-Biology-1http://en.wikipedia.org/wiki/Three_prime_untranslated_regionhttp://en.wikipedia.org/wiki/Upstream_and_downstream_(DNA)http://en.wikipedia.org/wiki/Five_prime_untranslated_regionhttp://en.wikipedia.org/wiki/Upstream_and_downstream_(DNA)http://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-Biology-1http://en.wikipedia.org/wiki/Ribozymehttp://en.wikipedia.org/wiki/Transfer_RNAhttp://en.wikipedia.org/wiki/Ribosomal_RNAhttp://en.wikipedia.org/wiki/LincRNAhttp://en.wikipedia.org/wiki/MicroRNAhttp://en.wikipedia.org/wiki/Non-coding_RNAhttp://en.wikipedia.org/wiki/Non-coding_RNAhttp://en.wikipedia.org/wiki/Translation_(genetics)http://en.wikipedia.org/wiki/Messenger_RNAhttp://en.wikipedia.org/wiki/Proteinhttp://en.wikipedia.org/wiki/Genehttp://en.wikipedia.org/wiki/Gene_expressionhttp://en.wikipedia.org/wiki/Nuclear_porehttp://en.wikipedia.org/wiki/Cell_nucleushttp://en.wikipedia.org/wiki/Primer_(molecular_biology)http://en.wikipedia.org/wiki/DNA_replicationhttp://en.wikipedia.org/wiki/DNA_replicationhttp://en.wikipedia.org/wiki/Thyminehttp://en.wikipedia.org/wiki/Uracilhttp://en.wikipedia.org/wiki/DNA_replicationhttp://en.wikipedia.org/wiki/Antiparallel_(biochemistry)http://en.wikipedia.org/wiki/RNA_polymerasehttp://en.wikipedia.org/wiki/Enzymehttp://en.wikipedia.org/wiki/Complementarity_(molecular_biology)http://en.wikipedia.org/wiki/Nucleotidehttp://en.wikipedia.org/wiki/Base_pairhttp://en.wikipedia.org/wiki/Nucleic_acidhttp://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-0http://en.wikipedia.org/wiki/DNAhttp://en.wikipedia.org/wiki/RNAhttp://en.wikipedia.org/wiki/Complementarity_(molecular_biology) 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same as the newly created RNA transcript (except for the substitution of uracil for thymine). The use of only
the 3' 5' strand eliminates the need for theOkazaki fragmentsseen in DNA replication.[2]
Transcription is divided into 5 stages: pre-initiation, initiation, promoter
clearance, elongationand termination.
[2]
Contents
[hide]
1 Major steps
o 1.1 Pre-initiation
o 1.2 Initiation
o 1.3 Promoter clearance
o 1.4 Elongation
o 1.5 Termination
2 Measuring and detecting transcription
3 Transcription factories
4 History
5 Reverse transcription
6 Inhibitors
7 See also
8 References
9 External links
[edit]Major steps
[edit]Pre-initiation
In eukaryotes,RNA polymerase, and therefore the initiation of transcription, requires the presence of a
corepromotersequence in the DNA. Promoters are regions of DNA that promote transcription and, in
eukaryotes, are found at -30, -75, and -90 base pairs upstream from the transcription start site(abbreviated
to TSS). Core promoters are sequences within the promoter that are essential for transcription initiation.
RNA polymerase is able to bind to core promoters in the presence of various specific transcription
factors.[citation needed]
The most characterized type of core promoter in eukaryotes is a short DNA sequence known as aTATA
box, found 25-30 base pairs upstream from the TSS.[citation needed]
The TATA box, as a core promoter, is the
binding site for a transcription factor known asTATA-binding protein(TBP), which is itself a subunit of
another transcription factor, calledTranscription Factor II D(TFIID). After TFIID binds to the TATA box via
the TBP, five more transcription factors and RNA polymerase combine around the TATA box in a series of
stages to form apreinitiation complex. One transcription factor,Transcription factor II H, has two
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iki/Transcription_preinitiation_complexhttp://en.wikipedia.org/wiki/Transcription_Factor_II_Dhttp://en.wikipedia.org/wiki/TATA-binding_proteinhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/TATA_boxhttp://en.wikipedia.org/wiki/TATA_boxhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Transcription_factorhttp://en.wikipedia.org/wiki/Transcription_factorhttp://en.wikipedia.org/wiki/Transcription_start_sitehttp://en.wikipedia.org/wiki/Promoter_(biology)http://en.wikipedia.org/wiki/RNA_polymerasehttp://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=2http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=1http://en.wikipedia.org/wiki/Transcription_(genetics)#External_linkshttp://en.wikipedia.org/wiki/Transcription_(genetics)#Referenceshttp://en.wikipedia.org/wiki/Transcription_(genetics)#See_alsohttp://en.wikipedia.org/wiki/Transcription_(genetics)#Inhibitorshttp://en.wikipedia.org/wiki/Transcription_(genetics)#Reverse_transcriptionhttp://en.wikipedia.org/wiki/Transcription_(genetics)#Historyhttp://en.wikipedia.org/wiki/Transcription_(genetics)#Transcription_factorieshttp://en.wikipedia.org/wiki/Transcription_(genetics)#Measuring_and_detecting_transcriptionhttp://en.wikipedia.org/wiki/Transcription_(genetics)#Terminationhttp://en.wikipedia.org/wiki/Transcription_(genetics)#Elongationhttp://en.wikipedia.org/wiki/Transcription_(genetics)#Promoter_clearancehttp://en.wikipedia.org/wiki/Transcription_(genetics)#Initiationhttp://en.wikipedia.org/wiki/Transcription_(genetics)#Pre-initiationhttp://en.wikipedia.org/wiki/Transcription_(genetics)#Major_stepshttp://en.wikipedia.org/wiki/Transcription_(genetics)http://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-Biology-1http://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-Biology-1http://en.wikipedia.org/wiki/Okazaki_fragment 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components withhelicaseactivity and so is involved in the separating of opposing strands of double-
stranded DNA to form the initial transcription bubble. However, only a low, or basal, rate of transcription is
driven by the preinitiation complex alone. Other proteins known asactivatorsandrepressors, along with
any associatedcoactivatorsorcorepressors, are responsible for modulating transcription rate.[citation needed]
Thus, preinitiation complex contains:[citation needed]
1. Core Promoter Sequence 2. Transcription Factors 3.
RNA Polymerase 4. Activators and Repressors. The transcription preinitiation inarchaeais, in essence,
homologous to that of eukaryotes, but is much less complex.[4]
The archaeal preinitiation complex
assembles at a TATA-box binding site; however, in archaea, this complex is composed of only RNA
polymerase II, TBP, and TFB (the archaeal homologue of eukaryotictranscription factor II B(TFIIB)).[5][6]
[edit]Initiation
Simple diagram of transcription initiation. RNAP = RNA polymerase
Inbacteria, transcription begins with the binding of RNA polymerase to the promoter in DNA. RNA
polymerase is acore enzymeconsisting of five subunits: 2 subunits, 1 subunit, 1 ' subunit, and 1
subunit. At the start of initiation, the core enzyme is associated with asigma factorthat aids in finding the
appropriate -35 and -10 base pairs downstream ofpromotersequences.[7]
When the sigma factor and RNA
polymerase combine, they form a holoenzyme.
Transcription initiation is more complex in eukaryotes. Eukaryotic RNA polymerase does not directly
recognize the core promoter sequences. Instead, a collection of proteins calledtranscription
factorsmediate the binding of RNA polymerase and the initiation of transcription. Only after certain
transcription factors are attached to the promoter does the RNA polymerase bind to it. The completed
assembly of transcription factors and RNA polymerase bind to the promoter, forming a transcription
initiation complex. Transcription in the archaea domain is similar to transcription in eukaryotes.[8]
[edit]Promoter clearance
After the first bond is synthesized, the RNA polymerase must clear the promoter. During this time there is a
tendency to release the RNA transcript and produce truncated transcripts. This is called abortive
initiationand is common for both eukaryotes and prokaryotes.[9]
Abortive initiation continues to occur until
the factor rearranges, resulting in the transcription elongation complex (which gives a 35 bp moving
footprint). The factor is released before 80 nucleotides of mRNA are synthesized .[10]
Once the transcript
reaches approximately 23 nucleotides, it no longer slips and elongation can occur. This, like most of the
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remainder of transcription, is anenergy-dependent process, consumingadenosine
triphosphate(ATP).[citation needed]
Promoter clearance coincides with phosphorylation of serine 5 on the carboxy terminal domain of RNAP II
in eukaryotes, which is phosphorylated by TFIIH.
[citation needed]
[edit]Elongation
Simple diagram of transcription elongation
One strand of the DNA, the template strand(or noncoding strand), is used as a template for RNA
synthesis. As transcription proceeds, RNA polymerase traverses the template strand and uses base pairing
complementarity with the DNA template to create an RNA copy. Although RNA polymerase traverses the
template strand from 3' 5', the coding (non-template) strand and newly-formed RNA can also be used as
reference points, so transcription can be described as occurring 5' 3'. This produces an RNA molecule
from 5' 3', an exact copy of the coding strand (except thatthyminesare replaced withuracils, and the
nucleotides are composed of a ribose (5-carbon) sugar where DNA has deoxyribose (one less oxygen
atom) in its sugar-phosphate backbone).[citation needed]
Unlike DNA replication, mRNA transcription can involve multiple RNA polymerases on a single DNA
template and multiple rounds of transcription (amplification of particular mRNA), so many mRNA molecules
can be rapidly produced from a single copy of a gene.[citation needed]
Elongation also involves a proofreading mechanism that can replace incorrectly incorporated bases. In
eukaryotes, this may correspond with short pauses during transcription that allow appropriate RNA editing
factors to bind. These pauses may be intrinsic to the RNA polymerase or due to chromatin structure.[citation
needed]
[edit]Termination
Bacteria use two different strategies for transcription termination.1.Rho-independent transcription 2.Rho-
dependent transcription. InRho-independent transcription termination,also called intrinsic termination, RNA
transcription stops when the newly synthesized RNA molecule forms a G-C-richhairpin loopfollowed by a
run of Us. When the hairpin forms, the mechanical stress breaks the weak rU-dA bonds, now filling the
DNA-RNA hybrid. This pulls the poly-U transcript out of the active site of the RNA polymerase, in effect,
terminating transcription. In the "Rho-dependent" type of termination, a protein factor called "Rho"
destabilizes the interaction between the template and the mRNA, thus releasing the newly synthesized
mRNA from the elongation complex.[11]
http://en.wikipedia.org/wiki/Energyhttp://en.wikipedia.org/wiki/Energyhttp://en.wikipedia.org/wiki/Energyhttp://en.wikipedia.org/wiki/Adenosine_triphosphatehttp://en.wikipedia.org/wiki/Adenosine_triphosphatehttp://en.wikipedia.org/wiki/Adenosine_triphosphatehttp://en.wikipedia.org/wiki/Adenosine_triphosphatehttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=5http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=5http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=5http://en.wikipedia.org/wiki/Thyminehttp://en.wikipedia.org/wiki/Thyminehttp://en.wikipedia.org/wiki/Thyminehttp://en.wikipedia.org/wiki/Uracilhttp://en.wikipedia.org/wiki/Uracilhttp://en.wikipedia.org/wiki/Uracilhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=6http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=6http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=6http://en.wikipedia.org/wiki/Rho-independent_transcription_terminationhttp://en.wikipedia.org/wiki/Rho-independent_transcription_terminationhttp://en.wikipedia.org/wiki/Rho-independent_transcription_terminationhttp://en.wikipedia.org/wiki/Hairpin_loophttp://en.wikipedia.org/wiki/Hairpin_loophttp://en.wikipedia.org/wiki/Hairpin_loophttp://en.wikipedia.org/wiki/Rho_factorhttp://en.wikipedia.org/wiki/Rho_factorhttp://en.wikipedia.org/wiki/Rho_factorhttp://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-10http://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-10http://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-10http://en.wikipedia.org/wiki/File:Simple_transcription_elongation1.svghttp://en.wikipedia.org/w/index.php?title=File:Simple_transcription_elongation1.svg&page=1http://en.wikipedia.org/wiki/File:Simple_transcription_elongation1.svghttp://en.wikipedia.org/w/index.php?title=File:Simple_transcription_elongation1.svg&page=1http://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-10http://en.wikipedia.org/wiki/Rho_factorhttp://en.wikipedia.org/wiki/Hairpin_loophttp://en.wikipedia.org/wiki/Rho-independent_transcription_terminationhttp://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=6http://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Uracilhttp://en.wikipedia.org/wiki/Thyminehttp://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=5http://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Adenosine_triphosphatehttp://en.wikipedia.org/wiki/Adenosine_triphosphatehttp://en.wikipedia.org/wiki/Energy -
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Transcription termination in eukaryotes is less understood but involves cleavage of the new transcript
followed by template-independent addition ofAs at its new 3' end, in a process calledpolyadenylation.[12]
[edit]Measuring and detecting transcription
Electron micrographof the ribosomal transcription process. The formingmRNAstrands are visible as branches from the
main DNA strand.[citation needed]
Transcription can be measured and detected in a variety of ways:[citation needed]
Nuclear Run-on assay: measures the relative abundance of newly formed transcripts
RNase protection assayandChIP-ChipofRNAP: detect active transcription sites
RT-PCR: measures the absolute abundance of total or nuclear RNA levels, which may however differ
from transcription rates
DNA microarrays: measures the relative abundance of the global total or nuclear RNA levels; however,
these may differ from transcription rates
In situ hybridization: detects the presence of a transcript
MS2 tagging: by incorporating RNAstem loops, such as MS2, into a gene, these become incorporated
into newly synthesized RNA. The stem loops can then be detected using a fusion of GFP and the MS2
coat protein, which has a high affinity, sequence-specific interaction with the MS2 stem loops. The
recruitment of GFP to the site of transcription is visualised as a single fluorescent spot. This
remarkable new approach has revealed that transcription occurs in discontinuous bursts, or pulses
(seeTranscriptional bursting). With the notable exception of in situ techniques, most other methods
provide cell population averages, and are not capable of detecting this fundamental property of
genes.[13]
Northern blot: the traditional method, and until the advent ofRNA-Seq, the most quantitative
http://en.wikipedia.org/wiki/Polyadenylationhttp://en.wikipedia.org/wiki/Polyadenylationhttp://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-11http://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-11http://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-11http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=7http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=7http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=7http://en.wikipedia.org/wiki/Electron_micrographhttp://en.wikipedia.org/wiki/Electron_micrographhttp://en.wikipedia.org/wiki/MRNAhttp://en.wikipedia.org/wiki/MRNAhttp://en.wikipedia.org/wiki/MRNAhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Nuclear_run-onhttp://en.wikipedia.org/wiki/Nuclear_run-onhttp://en.wikipedia.org/wiki/RNase_protection_assayhttp://en.wikipedia.org/wiki/RNase_protection_assayhttp://en.wikipedia.org/wiki/ChIP-Chiphttp://en.wikipedia.org/wiki/ChIP-Chiphttp://en.wikipedia.org/wiki/ChIP-Chiphttp://en.wikipedia.org/wiki/RNAPhttp://en.wikipedia.org/wiki/RNAPhttp://en.wikipedia.org/wiki/RNAPhttp://en.wikipedia.org/wiki/RT-PCRhttp://en.wikipedia.org/wiki/RT-PCRhttp://en.wikipedia.org/wiki/DNA_microarrayshttp://en.wikipedia.org/wiki/DNA_microarrayshttp://en.wikipedia.org/wiki/In_situ_hybridizationhttp://en.wikipedia.org/wiki/In_situ_hybridizationhttp://en.wikipedia.org/wiki/MS2_tagginghttp://en.wikipedia.org/wiki/MS2_tagginghttp://en.wikipedia.org/wiki/Stem-loophttp://en.wikipedia.org/wiki/Stem-loophttp://en.wikipedia.org/wiki/Stem-loophttp://en.wikipedia.org/wiki/Transcriptional_burstinghttp://en.wikipedia.org/wiki/Transcriptional_burstinghttp://en.wikipedia.org/wiki/Transcriptional_burstinghttp://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-12http://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-12http://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-12http://en.wikipedia.org/wiki/Northern_blothttp://en.wikipedia.org/wiki/Northern_blothttp://en.wikipedia.org/wiki/RNA-Seqhttp://en.wikipedia.org/wiki/RNA-Seqhttp://en.wikipedia.org/wiki/RNA-Seqhttp://en.wikipedia.org/wiki/File:Transcription_label_en.jpghttp://en.wikipedia.org/wiki/File:Transcription_label_en.jpghttp://en.wikipedia.org/wiki/File:Transcription_label_en.jpghttp://en.wikipedia.org/wiki/File:Transcription_label_en.jpghttp://en.wikipedia.org/wiki/RNA-Seqhttp://en.wikipedia.org/wiki/Northern_blothttp://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-12http://en.wikipedia.org/wiki/Transcriptional_burstinghttp://en.wikipedia.org/wiki/Stem-loophttp://en.wikipedia.org/wiki/MS2_tagginghttp://en.wikipedia.org/wiki/In_situ_hybridizationhttp://en.wikipedia.org/wiki/DNA_microarrayshttp://en.wikipedia.org/wiki/RT-PCRhttp://en.wikipedia.org/wiki/RNAPhttp://en.wikipedia.org/wiki/ChIP-Chiphttp://en.wikipedia.org/wiki/RNase_protection_assayhttp://en.wikipedia.org/wiki/Nuclear_run-onhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/MRNAhttp://en.wikipedia.org/wiki/Electron_micrographhttp://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=7http://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-11http://en.wikipedia.org/wiki/Polyadenylation -
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RNA-Seq: applies next-generation sequencing techniques to sequence whole transcriptomes, which
allows the measurement of relative abundance of RNA, as well as the detection of additional variations
such as fusion genes, post-translational edits and novel splice sites
[edit]Transcription factories
Main article:Transcription factories
Active transcription units are clustered in the nucleus, in discrete sites calledtranscription
factoriesoreuchromatin. Such sites can be visualized by allowing engaged polymerases to extend their
transcripts in tagged precursors (Br-UTP or Br-U) and immuno-labeling the tagged nascent RNA.
Transcription factories can also be localized using fluorescence in situ hybridization or marked by
antibodies directed against polymerases. There are ~10,000 factories in the nucleoplasm of a HeLa cell,
among which are ~8,000 polymerase II factories and ~2,000 polymerase III factories. Each polymerase II
factory contains ~8 polymerases. As most active transcription units are associated with only one
polymerase, each factory usually contains ~8 different transcription units. These units might be associated
through promoters and/or enhancers, with loops forming a cloud around the factor.[citation needed]
[edit]History
A molecule that allows the genetic material to be realized as a protein was first hypothesized byFranois
JacobandJacques Monod.Severo Ochoawon aNobel Prize in Physiology or Medicinefor developing a
process of RNA synthesis in 1959. RNA synthesis by RNA polymerase was established in vitroby several
laboratories by 1965; however, the RNA synthesized by these enzymes had properties that suggested the
existence of an additional factor needed to terminate transcription correctly.[citation needed]
In 1972, Walter Fiers became the first person to actually prove the existence of the terminating enzyme.
Roger D. Kornbergwon the 2006Nobel Prize in Chemistry"for his studies of the molecular basis of
eukaryotic transcription".[14]
[edit]Reverse transcription
http://en.wikipedia.org/wiki/RNA-Seqhttp://en.wikipedia.org/wiki/RNA-Seqhttp://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=8http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=8http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=8http://en.wikipedia.org/wiki/Transcription_factorieshttp://en.wikipedia.org/wiki/Transcription_factorieshttp://en.wikipedia.org/wiki/Transcription_factorieshttp://en.wikipedia.org/wiki/Transcription_factorieshttp://en.wikipedia.org/wiki/Transcription_factorieshttp://en.wikipedia.org/wiki/Transcription_factorieshttp://en.wikipedia.org/wiki/Transcription_factorieshttp://en.wikipedia.org/wiki/Euchromatinhttp://en.wikipedia.org/wiki/Euchromatinhttp://en.wikipedia.org/wiki/Euchromatinhttp://en.wikipedia.org/wiki/HeLa_cellhttp://en.wikipedia.org/wiki/HeLa_cellhttp://en.wikipedia.org/wiki/HeLa_cellhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=9http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=9http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=9http://en.wikipedia.org/wiki/Fran%C3%A7ois_Jacobhttp://en.wikipedia.org/wiki/Fran%C3%A7ois_Jacobhttp://en.wikipedia.org/wiki/Fran%C3%A7ois_Jacobhttp://en.wikipedia.org/wiki/Fran%C3%A7ois_Jacobhttp://en.wikipedia.org/wiki/Jacques_Monodhttp://en.wikipedia.org/wiki/Jacques_Monodhttp://en.wikipedia.org/wiki/Jacques_Monodhttp://en.wikipedia.org/wiki/Severo_Ochoahttp://en.wikipedia.org/wiki/Severo_Ochoahttp://en.wikipedia.org/wiki/Severo_Ochoahttp://en.wikipedia.org/wiki/Nobel_Prize_in_Physiology_or_Medicinehttp://en.wikipedia.org/wiki/Nobel_Prize_in_Physiology_or_Medicinehttp://en.wikipedia.org/wiki/Nobel_Prize_in_Physiology_or_Medicinehttp://en.wikipedia.org/wiki/In_vitrohttp://en.wikipedia.org/wiki/In_vitrohttp://en.wikipedia.org/wiki/In_vitrohttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Roger_D._Kornberghttp://en.wikipedia.org/wiki/Roger_D._Kornberghttp://en.wikipedia.org/wiki/Nobel_Prize_in_Chemistryhttp://en.wikipedia.org/wiki/Nobel_Prize_in_Chemistryhttp://en.wikipedia.org/wiki/Nobel_Prize_in_Chemistryhttp://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-13http://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-13http://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-13http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=10http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=10http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=10http://en.wikipedia.org/wiki/File:RetroTranscription.jpghttp://en.wikipedia.org/wiki/File:RetroTranscription.jpghttp://en.wikipedia.org/wiki/File:RetroTranscription.jpghttp://en.wikipedia.org/wiki/File:RetroTranscription.jpghttp://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=10http://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-13http://en.wikipedia.org/wiki/Nobel_Prize_in_Chemistryhttp://en.wikipedia.org/wiki/Roger_D._Kornberghttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/In_vitrohttp://en.wikipedia.org/wiki/Nobel_Prize_in_Physiology_or_Medicinehttp://en.wikipedia.org/wiki/Severo_Ochoahttp://en.wikipedia.org/wiki/Jacques_Monodhttp://en.wikipedia.org/wiki/Fran%C3%A7ois_Jacobhttp://en.wikipedia.org/wiki/Fran%C3%A7ois_Jacobhttp://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=9http://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/HeLa_cellhttp://en.wikipedia.org/wiki/Euchromatinhttp://en.wikipedia.org/wiki/Transcription_factorieshttp://en.wikipedia.org/wiki/Transcription_factorieshttp://en.wikipedia.org/wiki/Transcription_factorieshttp://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=8http://en.wikipedia.org/wiki/RNA-Seq -
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Scheme of reverse transcription
Someviruses(such asHIV, the cause ofAIDS), have the ability to transcribe RNA into DNA. HIV has an
RNA genome that is duplicated into DNA. The resulting DNA can be merged with the DNA genome of the
host cell. The main enzyme responsible for synthesis of DNA from an RNA template is called reverse
transcriptase. In the case of HIV, reverse transcriptase is responsible for synthesizing a complementary
DNAstrand (cDNA) to the viral RNA genome. An associated enzyme, ribonuclease H, digests the RNA
strand, and reverse transcriptase synthesises a complementary strand of DNA to form a double helix DNA
structure. This cDNA is integrated into the host cell's genome via another enzyme (integrase) causing the
host cell to generate viral proteins that reassemble into new viral particles. In HIV, subsequent to this, the
host cell undergoes programmed cell death,apoptosisofT cells.[15]
However, in other retroviruses, the host
cell remains intact as the virus buds out of the cell.
Some eukaryotic cells contain an enzyme with reverse transcription activity called telomerase. Telomerase
is a reverse transcriptase that lengthens the ends of linear chromosomes. Telomerase carries an RNA
template from which it synthesizes DNA repeating sequence, or "junk" DNA. This repeated sequence of
DNA is important because, every time a linear chromosome is duplicated, it is shortened in length. With
"junk" DNA at the ends of chromosomes, the shortening eliminates some of the non-essential, repeated
sequence rather than the protein-encoding DNA sequence farther away from the chromosome end.
Telomerase is often activated in cancer cells to enable cancer cells to duplicate their genomes indefinitely
without losing important protein-coding DNA sequence. Activation of telomerase could be part of the
process that allows cancer cells to become immortal. However, the truein vivosignificance of telomerasehas still not beenempiricallyproven.
[citation needed]
[edit]Inhibitors
Transcription inhibitors can be used asantibioticsagainst, for example,pathogenic bacteria(antibacterials)
andfungi(antifungals). An example of such an antibacterial isrifampicin, which inhibitsprokaryotic DNA
transcriptioninto mRNA by inhibiting DNA-dependentRNA polymeraseby binding its beta-subunit.8-
Hydroxyquinolineis an antifungal transcription inhibitor.[16]
[edit]
See also
http://en.wikipedia.org/wiki/Viruseshttp://en.wikipedia.org/wiki/Viruseshttp://en.wikipedia.org/wiki/Viruseshttp://en.wikipedia.org/wiki/HIVhttp://en.wikipedia.org/wiki/HIVhttp://en.wikipedia.org/wiki/HIVhttp://en.wikipedia.org/wiki/AIDShttp://en.wikipedia.org/wiki/AIDShttp://en.wikipedia.org/wiki/AIDShttp://en.wikipedia.org/wiki/Reverse_transcriptasehttp://en.wikipedia.org/wiki/Reverse_transcriptasehttp://en.wikipedia.org/wiki/Reverse_transcriptasehttp://en.wikipedia.org/wiki/Reverse_transcriptasehttp://en.wikipedia.org/wiki/Complementary_DNAhttp://en.wikipedia.org/wiki/Complementary_DNAhttp://en.wikipedia.org/wiki/Complementary_DNAhttp://en.wikipedia.org/wiki/Integrasehttp://en.wikipedia.org/wiki/Integrasehttp://en.wikipedia.org/wiki/Integrasehttp://en.wikipedia.org/wiki/Apoptosishttp://en.wikipedia.org/wiki/Apoptosishttp://en.wikipedia.org/wiki/Apoptosishttp://en.wikipedia.org/wiki/T_cellhttp://en.wikipedia.org/wiki/T_cellhttp://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-14http://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-14http://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-14http://en.wikipedia.org/wiki/Telomerasehttp://en.wikipedia.org/wiki/Telomerasehttp://en.wikipedia.org/wiki/Telomerasehttp://en.wikipedia.org/wiki/In_vivohttp://en.wikipedia.org/wiki/In_vivohttp://en.wikipedia.org/wiki/In_vivohttp://en.wikipedia.org/wiki/Empiricalhttp://en.wikipedia.org/wiki/Empiricalhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=11http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=11http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=11http://en.wikipedia.org/wiki/Antibiotichttp://en.wikipedia.org/wiki/Antibiotichttp://en.wikipedia.org/wiki/Antibiotichttp://en.wikipedia.org/wiki/Pathogenic_bacteriahttp://en.wikipedia.org/wiki/Pathogenic_bacteriahttp://en.wikipedia.org/wiki/Pathogenic_bacteriahttp://en.wikipedia.org/wiki/Antibacterialhttp://en.wikipedia.org/wiki/Antibacterialhttp://en.wikipedia.org/wiki/Antibacterialhttp://en.wikipedia.org/wiki/Fungushttp://en.wikipedia.org/wiki/Fungushttp://en.wikipedia.org/wiki/Fungushttp://en.wikipedia.org/wiki/Antifungal_medicationhttp://en.wikipedia.org/wiki/Antifungal_medicationhttp://en.wikipedia.org/wiki/Antifungal_medicationhttp://en.wikipedia.org/wiki/Rifampicinhttp://en.wikipedia.org/wiki/Rifampicinhttp://en.wikipedia.org/wiki/Rifampicinhttp://en.wikipedia.org/wiki/Prokaryotic_transcriptionhttp://en.wikipedia.org/wiki/Prokaryotic_transcriptionhttp://en.wikipedia.org/wiki/Prokaryotic_transcriptionhttp://en.wikipedia.org/wiki/RNA_polymerasehttp://en.wikipedia.org/wiki/RNA_polymerasehttp://en.wikipedia.org/wiki/RNA_polymerasehttp://en.wikipedia.org/wiki/8-Hydroxyquinolinehttp://en.wikipedia.org/wiki/8-Hydroxyquinolinehttp://en.wikipedia.org/wiki/8-Hydroxyquinolinehttp://en.wikipedia.org/wiki/8-Hydroxyquinolinehttp://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-15http://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-15http://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-15http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=12http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=12http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=12http://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=12http://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-15http://en.wikipedia.org/wiki/8-Hydroxyquinolinehttp://en.wikipedia.org/wiki/8-Hydroxyquinolinehttp://en.wikipedia.org/wiki/RNA_polymerasehttp://en.wikipedia.org/wiki/Prokaryotic_transcriptionhttp://en.wikipedia.org/wiki/Prokaryotic_transcriptionhttp://en.wikipedia.org/wiki/Rifampicinhttp://en.wikipedia.org/wiki/Antifungal_medicationhttp://en.wikipedia.org/wiki/Fungushttp://en.wikipedia.org/wiki/Antibacterialhttp://en.wikipedia.org/wiki/Pathogenic_bacteriahttp://en.wikipedia.org/wiki/Antibiotichttp://en.wikipedia.org/w/index.php?title=Transcription_(genetics)&action=edit§ion=11http://en.wikipedia.org/wiki/Wikipedia:Citation_neededhttp://en.wikipedia.org/wiki/Empiricalhttp://en.wikipedia.org/wiki/In_vivohttp://en.wikipedia.org/wiki/Telomerasehttp://en.wikipedia.org/wiki/Transcription_(genetics)#cite_note-14http://en.wikipedia.org/wiki/T_cellhttp://en.wikipedia.org/wiki/Apoptosishttp://en.wikipedia.org/wiki/Integrasehttp://en.wikipedia.org/wiki/Complementary_DNAhttp://en.wikipedia.org/wiki/Complementary_DNAhttp://en.wikipedia.org/wiki/Reverse_transcriptasehttp://en.wikipedia.org/wiki/Reverse_transcriptasehttp://en.wikipedia.org/wiki/AIDShttp://en.wikipedia.org/wiki/HIVhttp://en.wikipedia.org/wiki/Viruses