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From Wikipedia, the free encyclopedia
This article is about the biological macromolecule. For other uses, see RNA (disambiguation).
A hairpin loop from a pre-mRNA. Highlighted are the nucleobases (green) and the ribose-phosphate
backbone (blue). This is a single strand of RNA that folds back upon itself.
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Ribonucleic acid (RNA) is a polymeric molecule that is essential for most biological functions, either by
performing the function itself (non-coding RNA) or by forming a template for the production of
proteins (messenger RNA). RNA and deoxyribonucleic acid (DNA) are nucleic acids. The nucleic acids
constitute one of the four major macromolecules essential for all known forms of life. RNA is
assembled as a chain of nucleotides. Cellular organisms use messenger RNA (mRNA) to convey genetic
information (using the nitrogenous bases of guanine, uracil, adenine, and cytosine, denoted by the
letters G, U, A, and C) that directs synthesis of specific proteins. Many viruses encode their genetic
information using an RNA genome.
Some RNA molecules 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 in which RNA molecules direct the synthesis of
proteins on ribosomes. This process uses transfer RNA (tRNA) molecules to deliver amino acids to the
ribosome, where ribosomal RNA (rRNA) then links amino acids together to form coded proteins.
It has become widely accepted in science[1] that early in the history of life on Earth, prior to the
evolution of DNA and possibly of protein-based enzymes as well, an "RNA world" existed in which
RNA served as both living organisms' storage method for genetic information—a role fulfilled today by
DNA, except in the case of RNA viruses—and potentially performed catalytic functions in cells—a
function performed today by protein enzymes, with the notable and important exception of the
ribosome, which is a ribozyme.
Chemical structure of RNA
Main article: Nucleic acid structure
Basic chemical composition
Watson-Crick base pairs in a siRNA. Hydrogen atoms are not shown.
Each nucleotide in RNA contains 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). Adenine
and guanine are purines, and 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 each, making RNA a charged molecule (polyanion). The bases form standard hydrogen bonds
between cytosine and guanine and between adenine and uracil, while guanine and uracil can pair
through a non-canonical G–U wobble base pair.[2] However, other interactions are possible, such as a
group of adenine bases binding to each other in a bulge,[3] or the GNRA tetraloop that has a guanine–
adenine base-pair.[2]
Differences between DNA and RNA
Three-dimensional representation of the 50S ribosomal subunit. Ribosomal RNA is in brown, proteins
in blue. The active site is a small segment of rRNA, indicated in red.
The chemical structure of RNA is very similar to that of DNA, but differs in three primary ways:
Unlike double-stranded DNA, RNA is usually a single-stranded molecule (ssRNA)[4] in many of its
biological roles and consists of much shorter chains of nucleotides.[5] However, double-stranded RNA
(dsRNA) can form and (moreover) a single RNA molecule can, by complementary base pairing, form
intrastrand double helixes, as in tRNA.
While the sugar-phosphate "backbone" of DNA contains deoxyribose, RNA contains ribose instead.[6]
Ribose has a hydroxyl group attached to the pentose ring in the 2' position, whereas deoxyribose does
not. The hydroxyl groups in the ribose backbone make RNA more chemically labile than DNA by
lowering the activation energy of hydrolysis.
The complementary base to adenine in DNA is thymine, whereas in RNA, it is uracil, which is an
unmethylated form of thymine.[7]
Like DNA, most biologically active RNAs, including mRNA, tRNA, rRNA, snRNAs, and other non-coding
RNAs, contain self-complementary sequences that allow parts of the RNA to fold[8] 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).[9] For instance, determination of
the structure of the ribosome—an RNA-protein complex that catalyzes the assembly of proteins—
revealed that its active site is composed entirely of RNA.[10]
Structure of a fragment of an RNA, showing a guanosyl subunit
An important structural component of RNA that 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 take the A-form geometry,[11] although in single strand dinucleotide contexts, RNA
can rarely also adopt the B-form most commonly observed in DNA.[12] The A-form geometry results
in a very deep and narrow major groove and a shallow and wide minor groove.[13] A second
consequence of the presence of the 2'-hydroxyl group is that in conformationally flexible regions of an
RNA molecule (that is, not involved in formation of a double helix), it can chemically attack the
adjacent phosphodiester bond to cleave the backbone.[14]
Secondary and tertiary structures
The functional form of single-stranded RNA molecules, just like proteins, frequently requires a specific
spatial 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.[15] In order to create, i.e., design,
RNA for any given secondary structure, two or three bases would not be enough, but four bases are
enough.[16] This is likely why nature has "chosen" a four base alphabet: fewer than four would not
allow the creation of all structures, while more than four bases are not necessary to do so. Since RNA
is charged, metal ions such as Mg2+ are needed to stabilise many secondary and tertiary structures.
[17]
The naturally occurring enantiomer of RNA is D-RNA composed of D-ribonucleotides. All chirality
centers are located in the D-ribose. By the use of L-ribose or rather L-ribonucleotides, L-RNA can be
synthesized. L-RNA is much more stable against degradation by RNase.[18]
