RIBONUCLEIC ACID (RNA) is a polymeric molecule essential in various
biological roles in coding , decoding , regulation , and expression of
DNAare nucleic acids , and, along with lipids ,
proteins and carbohydrates , constitute the four major macromolecules
essential for all known forms of life . Like DNA,
RNAis assembled as
a chain of nucleotides , but unlike
DNAit is more often found in
nature as a single-strand folded onto itself, rather than a paired
double-strand. Cellular organisms use messenger
RNA(_MRNA_) to convey
genetic information (using the letters G, U, A, and C to denote the
nitrogenous bases guanine , uracil , adenine , and cytosine ) that
directs synthesis of specific proteins. Many viruses encode their
genetic information using an
RNAmolecules play an active role within cells by catalyzing
biological reactions, controlling gene expression , or sensing and
communicating responses to cellular signals. One of these active
processes is protein synthesis , a universal function where RNA
molecules direct the assembly of proteins on ribosomes . This process
RNA(_TRNA_) molecules to deliver amino acids to the
ribosome, where ribosomal
RNA(_RRNA_) then links amino acids together
to form proteins.
* 1 Comparison with
* 2 Structure
* 3 Synthesis
* 4 Types of
* 4.1 Overview
* 4.2 In length
* 4.3 In translation
* 4.4 Regulatory RNAs
* 4.5 In
* 4.7 In reverse transcription
* 4.8 Double-stranded
* 5 Key discoveries in
* 6 Evolution
* 7 See also
* 8 References
* 9 External links
COMPARISON WITH DNA
Bases in an
representation of the
Ribosomal RNAis in
ochre, proteins in blue. The active site is a small segment of rRNA,
indicated in red.
The chemical structure of
RNAis very similar to that of
differs in three main ways:
* Unlike double-stranded DNA,
RNAis a single-stranded molecule in
many of its biological roles and has a much shorter chain of
RNAcan, by complementary base pairing, form
intrastrand (i.e., single-strand) double helixes, as in tRNA.
DNAcontains _deoxyribose _,
RNAcontains _ribose _ (in
deoxyribose there is no hydroxyl group attached to the pentose ring in
the 2\' position). These hydroxyl groups make
RNAless stable than DNA
because it is more prone to hydrolysis .
* The complementary base to adenine in
DNAis thymine , whereas in
RNA, it is uracil , which is an unmethylated form of thymine.
Like DNA, most biologically active RNAs, including m
, snRNAs , and other non-coding RNAs , contain self-complementary
sequences that allow parts of the
RNAto fold and pair with itself to
form double helices. Analysis of these RNAs has revealed that they are
highly structured. Unlike DNA, their structures do not consist of long
double helices, but rather collections of short helices packed
together into structures akin to proteins. In this fashion, RNAs can
achieve chemical catalysis (like enzymes). For instance,
determination of the structure of the ribosome—an enzyme that
catalyzes peptide bond formation—revealed that its active site is
composed entirely of RNA.
Nucleic acidstructure Watson-Crick base pairs in
RNA(hydrogen atoms are not shown)
Each nucleotide in
RNAcontains a ribose sugar, with carbons numbered
1' through 5'. A base is attached to the 1' position, in general,
adenine (A), cytosine (C), guanine (G), or uracil (U).
guanine are purines , cytosine and uracil are pyrimidines . A
phosphate group is attached to the 3' position of one ribose and the
5' position of the next. The phosphate groups have a negative charge
RNAa charged molecule (polyanion). The bases form
hydrogen bonds between cytosine and guanine, between adenine and
uracil and between guanine and uracil. However, other interactions
are possible, such as a group of adenine bases binding to each other
in a bulge, or the GNRA tetraloop that has a guanine–adenine
base-pair. Chemical structure of
An important structural feature of
RNAthat distinguishes it from DNA
is the presence of a hydroxyl group at the 2' position of the ribose
sugar. The presence of this functional group causes the helix to
mostly adopt the A-form geometry , although in single strand
RNAcan rarely also adopt the B-form most
commonly observed in DNA. The A-form geometry results in a very deep
and narrow major groove and a shallow and wide minor groove. A second
consequence of the presence of the 2'-hydroxyl group is that in
conformationally flexible regions of an
RNAmolecule (that is, not
involved in formation of a double helix), it can chemically attack the
adjacent phosphodiester bond to cleave the backbone. Secondary
structure of a telomerase
RNAis transcribed with only four bases (adenine, cytosine, guanine
and uracil), but these bases and attached sugars can be modified in
numerous ways as the RNAs mature.
Pseudouridine(Ψ), in which the
linkage between uracil and ribose is changed from a C–N bond to a
C–C bond, and ribothymidine (T) are found in various places (the
most notable ones being in the TΨC loop of t
RNA). Another notable
modified base is hypoxanthine, a deaminated adenine base whose
nucleoside is called inosine (I).
Inosineplays a key role in the
wobble hypothesis of the genetic code .
There are more than 100 other naturally occurring modified
nucleosides, The greatest structural diversity of modifications can
be found in t
RNA, while pseudouridine and nucleosides with
2\'-O-methylribose often present in r
RNAare the most common. The
specific roles of many of these modifications in
RNAare not fully
understood. However, it is notable that, in ribosomal RNA, many of the
post-transcriptional modifications occur in highly functional regions,
such as the peptidyl transferase center and the subunit interface,
implying that they are important for normal function.
The functional form of single-stranded
RNAmolecules, just like
proteins, frequently requires a specific tertiary structure . The
scaffold for this structure is provided by secondary structural
elements that are hydrogen bonds within the molecule. This leads to
several recognizable "domains" of secondary structure like hairpin
loops , bulges, and internal loops . Since
RNAis charged, metal ions
such as Mg2+ are needed to stabilise many secondary and tertiary
The naturally occurring enantiomer of
D-ribonucleotides. All chirality centers are located in the D-ribose.
By the use of L-ribose or rather L-ribonucleotides, L-
RNAis much more stable against degradation by RNase .
Like other structured biopolymers such as proteins, one can define
topology of a folded
RNAmolecule. This is often done based on
arrangement of intra-chain contacts within a folded RNA, termed as
circuit topology .
RNAis usually catalyzed by an enzyme—
DNAas a template, a process known as transcription .
Initiation of transcription begins with the binding of the enzyme to a
promoter sequence in the
DNA(usually found "upstream" of a gene). The
DNAdouble helix is unwound by the helicase activity of the enzyme.
The enzyme then progresses along the template strand in the 3’ to
5’ direction, synthesizing a complementary
elongation occurring in the 5’ to 3’ direction. The
also dictates where termination of
RNAsynthesis will occur.
Primary transcriptRNAs are often modified by enzymes after
transcription. For example, a poly(A) tail and a 5\' cap are added to
RNAand introns are removed by the spliceosome .
There are also a number of RNA-dependent
RNApolymerases that use RNA
as their template for synthesis of a new strand of RNA. For instance,
a number of
RNAviruses (such as poliovirus) use this type of enzyme
to replicate their genetic material. Also, RNA-dependent RNA
polymerase is part of the
RNA interferencepathway in many organisms.
TYPES OF RNA
List of RNAs
Structure of a hammerhead ribozyme , a ribozyme that cuts
Messenger RNA(mRNA) is the
RNAthat carries information from
the ribosome , the sites of protein synthesis (translation ) in the
cell. The coding sequence of the m
RNAdetermines the amino acid
sequence in the protein that is produced. However, many RNAs do not
code for protein (about 97% of the transcriptional output is
non-protein-coding in eukaryotes ).
These so-called non-coding RNAs ("ncRNA") can be encoded by their own
RNAgenes), but can also derive from m
RNAintrons . The most
prominent examples of non-coding RNAs are transfer
RNA(rRNA), both of which are involved in the process of
translation. There are also non-coding RNAs involved in gene
RNA processingand other roles. Certain RNAs are able to
catalyse chemical reactions such as cutting and ligating other RNA
molecules, and the catalysis of peptide bond formation in the
ribosome ; these are known as ribozymes .
According to the length of
RNA. Usually, small RNAs are shorter than 200 nt in length, and long
RNAs are greater than 200 nt long. Long RNAs, also called large RNAs,
mainly include long non-coding
RNA(lncRNA) and m
RNA. Small RNAs
mainly include 5.8S ribosomal
RNA(rRNA), 5S rRNA, transfer RNA
RNA(miRNA), small interfering
Piwi-interacting RNA(piRNA), tRNA-derived
RNA(tsRNA) and small rDNA-derived
Messenger RNA(mRNA) carries information about a protein sequence to
the ribosomes , the protein synthesis factories in the cell. It is
coded so that every three nucleotides (a codon ) corresponds to one
amino acid. In eukaryotic cells, once precursor m
been transcribed from DNA, it is processed to mature mRNA. This
removes its introns —non-coding sections of the pre-mRNA. The mRNA
is then exported from the nucleus to the cytoplasm, where it is bound
to ribosomes and translated into its corresponding protein form with
the help of t
RNA. In prokaryotic cells, which do not have nucleus and
cytoplasm compartments, m
RNAcan bind to ribosomes while it is being
transcribed from DNA. After a certain amount of time the message
degrades into its component nucleotides with the assistance of
Transfer RNA(tRNA) is a small
RNAchain of about 80 nucleotides that
transfers a specific amino acid to a growing polypeptide chain at the
ribosomal site of protein synthesis during translation. It has sites
for amino acid attachment and an anticodon region for codon
recognition that binds to a specific sequence on the messenger RNA
chain through hydrogen bonding.
Ribosomal RNA(rRNA) is the catalytic component of the ribosomes.
Eukaryoticribosomes contain four different r
RNAmolecules: 18S, 5.8S,
28S and 5S rRNA. Three of the r
RNAmolecules are synthesized in the
nucleolus , and one is synthesized elsewhere. In the cytoplasm,
RNAand protein combine to form a nucleoprotein called a
ribosome. The ribosome binds m
RNAand carries out protein synthesis.
Several ribosomes may be attached to a single m
RNAat any time.
Nearly all the
RNAfound in a typical eukaryotic cell is rRNA.
Transfer-messenger RNA(tmRNA) is found in many bacteria and plastids
. It tags proteins encoded by mRNAs that lack stop codons for
degradation and prevents the ribosome from stalling.
Several types of
RNAcan downregulate gene expression by being
complementary to a part of an m
RNAor a gene's DNA. MicroRNAs
(miRNA; 21-22 nt ) are found in eukaryotes and act through RNA
interference (RNAi), where an effector complex of mi
can cleave complementary mRNA, block the m
RNAfrom being translated,
or accelerate its degradation.
While small interfering RNAs (siRNA; 20-25 nt) are often produced by
breakdown of viral RNA, there are also endogenous sources of siRNAs.
siRNAs act through
RNA interferencein a fashion similar to miRNAs.
Some miRNAs and siRNAs can cause genes they target to be methylated ,
thereby decreasing or increasing transcription of those genes.
Animals have Piwi-interacting RNAs (piRNA; 29-30 nt) that are active
in germline cells and are thought to be a defense against transposons
and play a role in gametogenesis .
Many prokaryotes have
CRISPRRNAs, a regulatory system similar to RNA
interference. Antisense RNAs are widespread; most downregulate a
gene, but a few are activators of transcription. One way antisense
RNAcan act is by binding to an mRNA, forming double-stranded
is enzymatically degraded. There are many long noncoding RNAs that
regulate genes in eukaryotes, one such
RNAis Xist , which coats one
X chromosome in female mammals and inactivates it.
RNAmay contain regulatory elements itself, such as riboswitches
, in the 5\' untranslated region or 3\' untranslated region ; these
cis-regulatory elements regulate the activity of that mRNA. The
untranslated regions can also contain elements that regulate other
Uridineto pseudouridine is a common
Many RNAs are involved in modifying other RNAs. Introns are spliced
out of pre-m
RNAby spliceosomes , which contain several small nuclear
RNAs (snRNA), or the introns can be ribozymes that are spliced by
RNAcan also be altered by having its nucleotides
modified to nucleotides other than A , C , G and U . In eukaryotes,
RNAnucleotides are in general directed by small
nucleolar RNAs (snoRNA; 60-300 nt), found in the nucleolus and cajal
bodies . snoRNAs associate with enzymes and guide them to a spot on an
RNAby basepairing to that RNA. These enzymes then perform the
nucleotide modification. rRNAs and tRNAs are extensively modified, but
snRNAs and mRNAs can also be the target of base modification. RNA
can also be methylated.
RNAcan carry genetic information.
RNAviruses have genomes
RNAthat encodes a number of proteins. The viral genome is
replicated by some of those proteins, while other proteins protect the
genome as the virus particle moves to a new host cell. Viroids are
another group of pathogens, but they consist only of RNA, do not
encode any protein and are replicated by a host plant cell's
IN REVERSE TRANSCRIPTION
Reverse transcribing viruses replicate their genomes by reverse
DNAcopies from their RNA; these
DNAcopies are then
transcribed to new RNA. Retrotransposons also spread by copying DNA
RNAfrom one another, and telomerase contains an
RNAthat is used
as template for building the ends of eukaryotic chromosomes.
RNAwith two complementary strands,
similar to the
DNAfound in all cells. ds
RNAforms the genetic
material of some viruses (double-stranded
RNAsuch as viral
RNAcan trigger RNA
interference in eukaryotes , as well as interferon response in
Recently, it was shown that there is a single stranded covalently
closed, _i.e._ circular form of
RNAexpressed throughout the animal
and plant kingdom (see circ
RNA). circRNAs are thought to arise via a
"back-splice" reaction where the spliceosome joins a downstream donor
to an upstream acceptor splice site. So far the function of circRNAs
is largely unknown, although for few examples a micro
activity has been demonstrated.
KEY DISCOVERIES IN
History of RNA biology Robert W. Holley,
left, poses with his research team.
RNAhas led to many important biological discoveries and
numerous Nobel Prizes. Nucleic acids were discovered in 1868 by
Friedrich Miescher, who called the material 'nuclein' since it was
found in the nucleus . It was later discovered that prokaryotic
cells, which do not have a nucleus, also contain nucleic acids. The
RNAin protein synthesis was suspected already in 1939.
Severo Ochoawon the 1959
Nobel Prize in Medicine(shared with Arthur
Kornberg ) after he discovered an enzyme that can synthesize
the laboratory. However, the enzyme discovered by Ochoa
(polynucleotide phosphorylase ) was later shown to be responsible for
RNAsynthesis. In 1956 Alex Rich and David Davies
hybridized two separate strands of
RNAto form the first crystal of
RNAwhose structure could be determined by X-ray crystallography.
The sequence of the 77 nucleotides of a yeast t
RNAwas found by
Robert W. Holley
Robert W. Holleyin 1965, winning Holley the 1968 Nobel Prize in
Medicine (shared with
Har Gobind Khorana
Har Gobind Khoranaand
Marshall Nirenberg). In
Carl Woesehypothesized that
RNAmight be catalytic and
suggested that the earliest forms of life (self-replicating molecules)
could have relied on
RNAboth to carry genetic information and to
catalyze biochemical reactions—an
During the early 1970s, retroviruses and reverse transcriptase were
discovered, showing for the first time that enzymes could copy RNA
DNA(the opposite of the usual route for transmission of genetic
information). For this work,
Howard Teminwere awarded a Nobel Prize in 1975. In 1976, Walter Fiers
and his team determined the first complete nucleotide sequence of an
RNA virusgenome, that of bacteriophage MS2 .
In 1977, introns and
RNA splicingwere discovered in both mammalian
viruses and in cellular genes, resulting in a 1993 Nobel to Philip
Sharp and Richard Roberts . Catalytic
RNAmolecules (ribozymes ) were
discovered in the early 1980s, leading to a 1989 Nobel award to Thomas
Sidney Altman. In 1990, it was found in _
introduced genes can silence similar genes of the plant's own, now
known to be a result of
At about the same time, 22 nt long RNAs, now called microRNAs , were
found to have a role in the development of _C. elegans _. Studies on
RNA interferencegleaned a Nobel Prize for Andrew Fire and Craig Mello
in 2006, and another Nobel was awarded for studies on the
Roger Kornbergin the same year. The discovery
of gene regulatory RNAs has led to attempts to develop drugs made of
RNA, such as si
RNA, to silence genes.
In March 2015, complex
RNAnucleotides , including uracil ,
cytosine and thymine , were reportedly formed in the laboratory under
outer space conditions, using starter chemicals, such as pyrimidine ,
an organic compound commonly found in meteorites . Pyrimidine, like
polycyclic aromatic hydrocarbons (PAHs), is one of the most
carbon-rich compounds found in the
Universeand may have been formed
in red giants or in interstellar dust and gas clouds.
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