Ribonucleic acid (RNA) is a polymeric molecule essential in various
biological roles in coding, decoding, regulation, and expression of
DNA are nucleic acids, and, along with lipids, proteins
and carbohydrates, constitute the four major macromolecules essential
for all known forms of life. Like DNA,
RNA is assembled as a chain of
nucleotides, but unlike
DNA it 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 nitrogenous bases 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 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 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 DNA
4 Types of RNA
4.2 In length
4.3 In translation
4.4 Regulatory RNAs
4.7 In reverse transcription
4.8 Double-stranded RNA
4.9 Circular RNA
5 Key discoveries in
5.1 Relevance for prebiotic chemistry and abiogenesis
6 See also
8 External links
Comparison with DNA
Bases in an
Three-dimensional representation of the
50S ribosomal subunit.
Ribosomal RNA is in ochre, 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 a single-stranded molecule in
many of its biological roles and consists of a much shorter chain of
RNA can, by complementary base pairing, form
intrastrand (i.e., single-strand) double helixes, as in tRNA.
While the sugar-phosphate "backbone" of
DNA contains deoxyribose, RNA
contains ribose instead.
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 less stable than DNA
because it is more prone to hydrolysis.
The complementary base to adenine in
DNA is thymine, whereas in RNA,
it is uracil, which is an unmethylated form of thymine.
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 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 RNA-protein
complex that catalyzes peptide bond formation—revealed that its
active site is composed entirely of RNA.
Nucleic acid structure
Watson-Crick base pairs in a si
RNA (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).
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 each, making
RNA a 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 RNA
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, although in single strand
RNA can 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
RNA molecule (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 RNA.
RNA is 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 tRNA). Another notable
modified base is hypoxanthine, a deaminated adenine base whose
nucleoside is called inosine (I).
Inosine plays 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 tRNA, while pseudouridine and nucleosides with
2'-O-methylribose often present in r
RNA are the most common. The
specific roles of many of these modifications in
RNA are 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
RNA molecules, 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
RNA is charged, metal
ions such as Mg2+ are needed to stabilise many secondary and tertiary
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
RNA is much more stable against degradation by
Like other structured biopolymers such as proteins, one can define
topology of a folded
RNA molecule. This is often done based on
arrangement of intra-chain contacts within a folded RNA, termed as
RNA is usually catalyzed by an enzyme—RNA
DNA as 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
DNA double 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
with elongation occurring in the 5’ to 3’ direction. The DNA
sequence also dictates where termination of
RNA synthesis will
Primary transcript RNAs are often modified by enzymes after
transcription. For example, a poly(A) tail and a
5' cap are added to
RNA and introns are removed by the spliceosome.
There are also a number of RNA-dependent
RNA polymerases that use RNA
as their template for synthesis of a new strand of RNA. For instance,
a number of
RNA viruses (such as poliovirus) use this type of enzyme
to replicate their genetic material. Also, RNA-dependent RNA
polymerase is part of the
RNA interference pathway in many
Types of RNA
See also: List of RNAs
Structure of a hammerhead ribozyme, a ribozyme that cuts RNA
Messenger RNA (mRNA) is the
RNA that carries information from
the ribosome, the sites of protein synthesis (translation) in the
cell. The coding sequence of the m
RNA determines 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
RNA genes), but can also derive from m
RNA introns. The most
prominent examples of non-coding RNAs are transfer
RNA (tRNA) and
RNA (rRNA), both of which are involved in the process of
translation. There are also non-coding RNAs involved in gene
RNA processing and 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 includes small
RNA and long
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
mRNA. Small RNAs mainly include 5.8S ribosomal
RNA (rRNA), 5S rRNA,
RNA (tRNA), micro
RNA (miRNA), small interfering
Piwi-interacting RNA (piRNA),
RNA (tsRNA) and small rDNA-derived RNA
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
RNA (pre-mRNA) has
been transcribed from DNA, it is processed to mature mRNA. This
removes its introns—non-coding sections of the pre-mRNA. The m
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 tRNA. In prokaryotic cells, which do not have nucleus and
cytoplasm compartments, m
RNA can 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
RNA chain 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.
Eukaryotic ribosomes contain four different r
RNA molecules: 18S, 5.8S,
28S and 5S rRNA. Three of the r
RNA molecules are synthesized in the
nucleolus, and one is synthesized elsewhere. In the cytoplasm,
RNA and protein combine to form a nucleoprotein called a
ribosome. The ribosome binds m
RNA and carries out protein synthesis.
Several ribosomes may be attached to a single m
RNA at any time.
Nearly all the
RNA found 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
RNA can downregulate gene expression by being
complementary to a part of an m
RNA or 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
RNA and enzymes
can cleave complementary mRNA, block the m
RNA from 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 interference in 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
Many prokaryotes have
CRISPR RNAs, 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
RNA can 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
RNA is Xist, which coats
one X chromosome in female mammals and inactivates it.
RNA may 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
Uridine to pseudouridine is a common
Many RNAs are involved in modifying other RNAs. Introns are spliced
out of pre-m
RNA by spliceosomes, which contain several small nuclear
RNAs (snRNA), or the introns can be ribozymes that are spliced by
RNA can also be altered by having its nucleotides
modified to nucleotides other than A, C, G and U. In eukaryotes,
RNA nucleotides 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
RNA by 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
RNA can also be methylated.
RNA can carry genetic information.
RNA viruses have genomes
RNA that 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
DNA copies from their RNA; these
DNA copies are then
transcribed to new RNA. Retrotransposons also spread by copying DNA
RNA from one another, and telomerase contains an
RNA that is
used as template for building the ends of eukaryotic chromosomes.
RNA (dsRNA) is
RNA with two complementary strands,
similar to the
DNA found in all cells. ds
RNA forms the genetic
material of some viruses (double-stranded
RNA such as viral
RNA or si
RNA can trigger RNA
interference in eukaryotes, as well as interferon response in
In the late 1970s, it was shown that there is a single stranded
covalently closed, i.e. circular form of
RNA expressed throughout the
animal and plant kingdom (see circRNA). 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
RNA sponging activity has been demonstrated.
Key discoveries in
Further information: History of
Robert W. Holley, left, poses with his research team.
RNA has 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
RNA in protein synthesis was suspected already in 1939.
Severo Ochoa won the 1959
Nobel Prize in Medicine
Nobel Prize in Medicine (shared with Arthur
Kornberg) after he discovered an enzyme that can synthesize
RNA in the
laboratory. However, the enzyme discovered by Ochoa
(polynucleotide phosphorylase) was later shown to be responsible for
RNA degradation, not
RNA synthesis. In 1956 Alex Rich and David Davies
hybridized two separate strands of
RNA to form the first crystal of
RNA whose structure could be determined by X-ray crystallography.
The sequence of the 77 nucleotides of a yeast t
RNA was found by Robert
W. Holley in 1965, winning Holley the 1968 Nobel Prize in Medicine
Har Gobind Khorana
Har Gobind Khorana and Marshall Nirenberg).
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, David Baltimore,
Renato Dulbecco and
Howard Temin were awarded a Nobel Prize in 1975. In 1976, Walter Fiers
and his team determined the first complete nucleotide sequence of an
RNA virus genome, that of bacteriophage MS2.
In 1977, introns and
RNA splicing were discovered in both mammalian
viruses and in cellular genes, resulting in a 1993 Nobel to Philip
Sharp and Richard Roberts. Catalytic
RNA molecules (ribozymes) were
discovered in the early 1980s, leading to a 1989 Nobel award to Thomas
Cech and 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 interference gleaned a Nobel Prize for Andrew Fire and Craig Mello
in 2006, and another Nobel was awarded for studies on the
Roger Kornberg in the same year. The discovery
of gene regulatory RNAs has led to attempts to develop drugs made of
RNA, such as siRNA, to silence genes.
Relevance for prebiotic chemistry and abiogenesis
Carl Woese hypothesized that
RNA might be catalytic and
suggested that the earliest forms of life (self-replicating molecules)
could have relied on
RNA both to carry genetic information and to
catalyze biochemical reactions—an
In March 2015, complex
RNA nucleotides, 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
Universe and may have been formed
in red giants or in interstellar dust and gas clouds.
^ "RNA: The Versatile Molecule". University of Utah. 2015.
^ "Nucleotides and Nucleic Acids" (PDF). University of California, Los
^ Shukla RN (2014-06-30). Analysis of Chromosomes.
^ a b c Berg JM, Tymoczko JL, Stryer L (2002). Biochemistry (5th ed.).
WH Freeman and Company. pp. 118–19, 781–808.
ISBN 0-7167-4684-0. OCLC 179705944.
^ Tinoco I & Bustamante C (October 1999). "How
RNA folds". Journal
of Molecular Biology. 293 (2): 271–81. doi:10.1006/jmbi.1999.3001.
^ Higgs PG (August 2000). "
RNA secondary structure: physical and
computational aspects". Quarterly Reviews of Biophysics. 33 (3):
199–253. doi:10.1017/S0033583500003620. PMID 11191843.
^ a b Nissen P, Hansen J, Ban N, Moore PB, Steitz TA (August 2000).
"The structural basis of ribosome activity in peptide bond synthesis".
Science. 289 (5481): 920–30. Bibcode:2000Sci...289..920N.
doi:10.1126/science.289.5481.920. PMID 10937990.
^ a b Lee JC, Gutell RR (December 2004). "Diversity of base-pair
conformations and their occurrence in r
RNA structure and RNA
structural motifs". Journal of Molecular Biology. 344 (5): 1225–49.
doi:10.1016/j.jmb.2004.09.072. PMID 15561141.
^ Barciszewski J, Frederic B, Clark C (1999).
RNA biochemistry and
biotechnology. Springer. pp. 73–87. ISBN 0-7923-5862-7.
^ Salazar M, Fedoroff OY, Miller JM, Ribeiro NS, Reid BR (April 1993).
DNA strand in DNA.
RNA hybrid duplexes is neither B-form nor
A-form in solution". Biochemistry. 32 (16): 4207–15.
doi:10.1021/bi00067a007. PMID 7682844.
^ Sedova A, Banavali NK (October 2015). "
RNA approaches the B-form in
stacked single strand dinucleotide contexts". Biopolymers. 105 (2):
65–82. doi:10.1002/bip.22750. PMID 26443416.
^ Hermann T, Patel DJ (March 2000). "
RNA bulges as architectural and
recognition motifs". Structure. 8 (3): R47–54.
doi:10.1016/S0969-2126(00)00110-6. PMID 10745015.
^ Mikkola S, Stenman E, Nurmi K, Yousefi-Salakdeh E, Strömberg R,
Lönnberg H (1999). "The mechanism of the metal ion promoted cleavage
RNA phosphodiester bonds involves a general acid catalysis by the
metal aquo ion on the departure of the leaving group". Perkin
transactions 2 (8): 1619–26. doi:10.1039/a903691a.
^ Jankowski JA, Polak JM (1996). Clinical gene analysis and
manipulation: Tools, techniques and troubleshooting. Cambridge
University Press. p. 14. ISBN 0-521-47896-0.
^ Yu Q, Morrow CD (May 2001). "Identification of critical elements in
RNA acceptor stem and T(Psi)C loop necessary for human
immunodeficiency virus type 1 infectivity". Journal of Virology. 75
(10): 4902–6. doi:10.1128/JVI.75.10.4902-4906.2001.
PMC 114245 . PMID 11312362.
^ Elliott MS, Trewyn RW (February 1984). "
Inosine biosynthesis in
RNA by an enzymatic insertion of hypoxanthine". The Journal
of Biological Chemistry. 259 (4): 2407–10. PMID 6365911.
^ Cantara WA, Crain PF, Rozenski J, McCloskey JA, Harris KA, Zhang X,
Vendeix FA, Fabris D, Agris PF (January 2011). "The
Database, RNAMDB: 2011 update". Nucleic Acids Research. 39 (Database
issue): D195–201. doi:10.1093/nar/gkq1028. PMC 3013656 .
^ Söll D, RajBhandary U (1995). TRNA: Structure, biosynthesis, and
function. ASM Press. p. 165. ISBN 1-55581-073-X.
^ Kiss T (July 2001). "Small nucleolar RNA-guided post-transcriptional
modification of cellular RNAs". The EMBO Journal. 20 (14): 3617–22.
doi:10.1093/emboj/20.14.3617. PMC 125535 .
^ King TH, Liu B, McCully RR, Fournier MJ (February 2003). "Ribosome
structure and activity are altered in cells lacking snoRNPs that form
pseudouridines in the peptidyl transferase center". Molecular Cell. 11
(2): 425–35. doi:10.1016/S1097-2765(03)00040-6.
^ Mathews DH, Disney MD, Childs JL, Schroeder SJ, Zuker M, Turner DH
(May 2004). "Incorporating chemical modification constraints into a
dynamic programming algorithm for prediction of
structure". Proceedings of the National Academy of Sciences of the
United States of America. 101 (19): 7287–92.
PMC 409911 . PMID 15123812.
^ Tan ZJ, Chen SJ (July 2008). "Salt dependence of nucleic acid
hairpin stability". Biophysical Journal. 95 (2): 738–52.
PMC 2440479 . PMID 18424500.
^ Vater A, Klussmann S (January 2015). "Turning mirror-image
oligonucleotides into drugs: the evolution of Spiegelmer(®)
therapeutics". Drug Discovery Today. 20 (1): 147–55.
doi:10.1016/j.drudis.2014.09.004. PMID 25236655.
^ Nudler E, Gottesman ME (August 2002). "Transcription termination and
anti-termination in E. coli". Genes to Cells. 7 (8): 755–68.
doi:10.1046/j.1365-2443.2002.00563.x. PMID 12167155.
^ Hansen JL, Long AM, Schultz SC (August 1997). "Structure of the
RNA polymerase of poliovirus". Structure. 5 (8):
1109–22. doi:10.1016/S0969-2126(97)00261-X. PMID 9309225.
^ Ahlquist P (May 2002). "RNA-dependent
RNA polymerases, viruses, and
RNA silencing". Science. 296 (5571): 1270–3.
^ a b c Cooper GC, Hausman RE (2004). The Cell: A Molecular Approach
(3rd ed.). Sinauer. pp. 261–76, 297, 339–44.
ISBN 0-87893-214-3. OCLC 174924833.
^ Mattick JS, Gagen MJ (September 2001). "The evolution of controlled
multitasked gene networks: the role of introns and other noncoding
RNAs in the development of complex organisms". Molecular Biology and
Evolution. 18 (9): 1611–30.
doi:10.1093/oxfordjournals.molbev.a003951. PMID 11504843.
^ Mattick JS (November 2001). "Non-coding RNAs: the architects of
eukaryotic complexity". EMBO Reports. 2 (11): 986–91.
doi:10.1093/embo-reports/kve230. PMC 1084129 .
^ Mattick JS (October 2003). "Challenging the dogma: the hidden layer
of non-protein-coding RNAs in complex organisms" (PDF). BioEssays. 25
(10): 930–9. doi:10.1002/bies.10332. PMID 14505360. Archived
from the original (PDF) on 2009-03-06.
^ Mattick JS (October 2004). "The hidden genetic program of complex
organisms". Scientific American. 291 (4): 60–7.
doi:10.1038/scientificamerican1004-60. PMID 15487671. [dead
^ a b c Wirta W (2006). Mining the transcriptome – methods and
applications. Stockholm: School of Biotechnology, Royal Institute of
Technology. ISBN 91-7178-436-5. OCLC 185406288.
^ Rossi JJ (July 2004). "
Ribozyme diagnostics comes of age". Chemistry
& Biology. 11 (7): 894–5. doi:10.1016/j.chembiol.2004.07.002.
^ Storz G (May 2002). "An expanding universe of noncoding RNAs".
Science. 296 (5571): 1260–3. Bibcode:2002Sci...296.1260S.
doi:10.1126/science.1072249. PMID 12016301.
^ Fatica A, Bozzoni I (January 2014). "Long non-coding RNAs: new
players in cell differentiation and development". Nature Reviews.
Genetics. 15 (1): 7–21. doi:10.1038/nrg3606.
^ Chen Q, Yan M, Cao Z, Li X, Zhang Y, Shi J, et al. (January 2016).
"Sperm tsRNAs contribute to intergenerational inheritance of an
acquired metabolic disorder". Science. 351 (6271): 397–400.
^ Wei H, Zhou B, Zhang F, Tu Y, Hu Y, Zhang B, Zhai Q (2013).
"Profiling and identification of small rDNA-derived RNAs and their
potential biological functions". PLOS One. 8 (2): e56842.
PMC 3572043 . PMID 23418607.
^ Gueneau de Novoa P, Williams KP (January 2004). "The tm
reductive evolution of tm
RNA in plastids and other endosymbionts".
Nucleic Acids Research. 32 (Database issue): D104–8.
doi:10.1093/nar/gkh102. PMC 308836 . PMID 14681369.
^ Carthew RW, Sontheimer EJ (February 2009). "Origins and Mechanisms
of miRNAs and siRNAs". Cell. 136 (4): 642–55.
doi:10.1016/j.cell.2009.01.035. PMC 2675692 .
^ Liang KH, Yeh CT (May 2013). "A gene expression restriction network
mediated by sense and antisense Alu sequences located on
protein-coding messenger RNAs". BMC Genomics. 14: 325.
doi:10.1186/1471-2164-14-325. PMC 3655826 .
^ Wu L, Belasco JG (January 2008). "Let me count the ways: mechanisms
of gene regulation by miRNAs and siRNAs". Molecular Cell. 29 (1):
1–7. doi:10.1016/j.molcel.2007.12.010. PMID 18206964.
^ Matzke MA, Matzke AJ (May 2004). "Planting the seeds of a new
paradigm". PLoS Biology. 2 (5): E133.
doi:10.1371/journal.pbio.0020133. PMC 406394 .
^ Vazquez F, Vaucheret H, Rajagopalan R, Lepers C, Gasciolli V,
Mallory AC, Hilbert JL, Bartel DP, Crété P (October 2004).
"Endogenous trans-acting siRNAs regulate the accumulation of
Arabidopsis mRNAs". Molecular Cell. 16 (1): 69–79.
doi:10.1016/j.molcel.2004.09.028. PMID 15469823.
^ Watanabe T, Totoki Y, Toyoda A, et al. (May 2008). "Endogenous
siRNAs from naturally formed dsRNAs regulate transcripts in mouse
oocytes". Nature. 453 (7194): 539–43. Bibcode:2008Natur.453..539W.
doi:10.1038/nature06908. PMID 18404146.
^ Sontheimer EJ, Carthew RW (July 2005). "Silence from within:
endogenous siRNAs and miRNAs". Cell. 122 (1): 9–12.
doi:10.1016/j.cell.2005.06.030. PMID 16009127.
^ Doran G (2007). "RNAi – Is one suffix sufficient?". Journal
of RNAi and
Gene Silencing. 3 (1): 217–19. Archived from the
original on 2007-07-16.
^ Pushparaj PN, Aarthi JJ, Kumar SD, Manikandan J (January 2008).
"RNAi and RNAa--the yin and yang of RNAome". Bioinformation. 2 (6):
235–7. doi:10.6026/97320630002235. PMC 2258431 .
^ Horwich MD, Li C, Matranga C, Vagin V, Farley G, Wang P, Zamore PD
(July 2007). "The Drosophila
RNA methyltransferase, DmHen1, modifies
germline piRNAs and single-stranded siRNAs in RISC". Current Biology.
17 (14): 1265–72. doi:10.1016/j.cub.2007.06.030.
^ Girard A, Sachidanandam R, Hannon GJ, Carmell MA (July 2006). "A
germline-specific class of small RNAs binds mammalian Piwi proteins".
Nature. 442 (7099): 199–202. Bibcode:2006Natur.442..199G.
doi:10.1038/nature04917. PMID 16751776.
^ Horvath P, Barrangou R (January 2010). "CRISPR/Cas, the immune
system of bacteria and archaea". Science. 327 (5962): 167–70.
^ Wagner EG, Altuvia S, Romby P (2002). "Antisense RNAs in bacteria
and their genetic elements". Advances in Genetics. Advances in
Genetics. 46: 361–98. doi:10.1016/S0065-2660(02)46013-0.
ISBN 9780120176465. PMID 11931231.
^ Gilbert SF (2003). Developmental Biology (7th ed.). Sinauer.
pp. 101–3. ISBN 0-87893-258-5.
OCLC 154656422. CS1 maint: Uses authors parameter (link)
^ Amaral PP, Mattick JS (August 2008). "Noncoding
RNA in development".
Mammalian Genome. 19 (7–8): 454–92. doi:10.1007/s00335-008-9136-7.
^ Heard E, Mongelard F, Arnaud D, Chureau C, Vourc'h C, Avner P (June
1999). "Human XIST yeast artificial chromosome transgenes show partial
X inactivation center function in mouse embryonic stem cells".
Proceedings of the National Academy of Sciences of the United States
of America. 96 (12): 6841–6. Bibcode:1999PNAS...96.6841H.
doi:10.1073/pnas.96.12.6841. PMC 22003 .
^ Batey RT (June 2006). "Structures of regulatory elements in mRNAs".
Current Opinion in Structural Biology. 16 (3): 299–306.
doi:10.1016/j.sbi.2006.05.001. PMID 16707260.
^ Scotto L, Assoian RK (June 1993). "A GC-rich domain with
bifunctional effects on m
RNA and protein levels: implications for
control of transforming growth factor beta 1 expression". Molecular
and Cellular Biology. 13 (6): 3588–97. PMC 359828 .
^ Steitz TA, Steitz JA (July 1993). "A general two-metal-ion mechanism
for catalytic RNA". Proceedings of the National Academy of Sciences of
the United States of America. 90 (14): 6498–502.
PMC 46959 . PMID 8341661.
^ Xie J, Zhang M, Zhou T, Hua X, Tang L, Wu W (January 2007).
"Sno/scaRNAbase: a curated database for small nucleolar RNAs and cajal
body-specific RNAs". Nucleic Acids Research. 35 (Database issue):
D183–7. doi:10.1093/nar/gkl873. PMC 1669756 .
^ Omer AD, Ziesche S, Decatur WA, Fournier MJ, Dennis PP (May 2003).
"RNA-modifying machines in archaea". Molecular Microbiology. 48 (3):
^ Cavaillé J, Nicoloso M, Bachellerie JP (October 1996). "Targeted
ribose methylation of
RNA in vivo directed by tailored antisense RNA
guides". Nature. 383 (6602): 732–5. Bibcode:1996Natur.383..732C.
doi:10.1038/383732a0. PMID 8878486.
^ Kiss-László Z, Henry Y, Bachellerie JP, Caizergues-Ferrer M, Kiss
T (June 1996). "Site-specific ribose methylation of preribosomal RNA:
a novel function for small nucleolar RNAs". Cell. 85 (7): 1077–88.
doi:10.1016/S0092-8674(00)81308-2. PMID 8674114.
^ Daròs JA, Elena SF, Flores R (June 2006). "Viroids: an Ariadne's
thread into the
RNA labyrinth". EMBO Reports. 7 (6): 593–8.
doi:10.1038/sj.embor.7400706. PMC 1479586 .
^ Kalendar R, Vicient CM, Peleg O, Anamthawat-Jonsson K, Bolshoy A,
Schulman AH (March 2004). "Large retrotransposon derivatives:
abundant, conserved but nonautonomous retroelements of barley and
related genomes". Genetics. 166 (3): 1437–50.
doi:10.1534/genetics.166.3.1437. PMC 1470764 .
^ Podlevsky JD, Bley CJ, Omana RV, Qi X, Chen JJ (January 2008). "The
telomerase database". Nucleic Acids Research. 36 (Database issue):
D339–43. doi:10.1093/nar/gkm700. PMC 2238860 .
^ Blevins T, Rajeswaran R, Shivaprasad PV, Beknazariants D, Si-Ammour
A, Park HS, Vazquez F, Robertson D, Meins F, Hohn T, Pooggin MM
(2006). "Four plant Dicers mediate viral small
RNA biogenesis and DNA
virus induced silencing". Nucleic Acids Research. 34 (21): 6233–46.
doi:10.1093/nar/gkl886. PMC 1669714 . PMID 17090584.
^ Jana S, Chakraborty C, Nandi S, Deb JK (November 2004). "RNA
interference: potential therapeutic targets". Applied Microbiology and
Biotechnology. 65 (6): 649–57. doi:10.1007/s00253-004-1732-1.
^ Schultz U, Kaspers B, Staeheli P (May 2004). "The interferon system
of non-mammalian vertebrates". Developmental and Comparative
Immunology. 28 (5): 499–508. doi:10.1016/j.dci.2003.09.009.
^ Whitehead KA, Dahlman JE, Langer RS, Anderson DG (2011). "Silencing
or stimulation? si
RNA delivery and the immune system". Annual Review
of Chemical and Biomolecular Engineering. 2: 77–96.
^ Hsu, Ming-Ta; Coca-Prados, Miguel (26 July 1979). "Electron
microscopic evidence for the circular form of
RNA in the cytoplasm of
eukaryotic cells". Nature. 280 (5720): 339–340.
doi:10.1038/280339a0. ISSN 1476-4687.
^ Dahm R (February 2005). "
Friedrich Miescher and the discovery of
DNA". Developmental Biology. 278 (2): 274–88.
doi:10.1016/j.ydbio.2004.11.028. PMID 15680349.
^ Caspersson T, Schultz J (1939). "Pentose nucleotides in the
cytoplasm of growing tissues". Nature. 143 (3623): 602–3.
^ Ochoa S (1959). "Enzymatic synthesis of ribonucleic acid" (PDF).
^ Rich A, Davies D (1956). "A New Two-Stranded Helical Structure:
Polyadenylic Acid and Polyuridylic Acid". Journal of the American
Chemical Society. 78 (14): 3548–3549. doi:10.1021/ja01595a086.
^ Holley RW, et al. (March 1965). "Structure of a ribonucleic acid".
Science. 147 (3664): 1462–5. Bibcode:1965Sci...147.1462H.
doi:10.1126/science.147.3664.1462. PMID 14263761.
^ Fiers W, et al. (April 1976). "Complete nucleotide sequence of
bacteriophage MS2 RNA: primary and secondary structure of the
replicase gene". Nature. 260 (5551): 500–7.
^ Napoli C, Lemieux C, Jorgensen R (April 1990). "Introduction of a
Chimeric Chalcone Synthase
Petunia Results in Reversible
Co-Suppression of Homologous Genes in trans". The Plant Cell. 2 (4):
279–289. doi:10.1105/tpc.2.4.279. PMC 159885 .
^ Dafny-Yelin M, Chung SM, Frankman EL, Tzfira T (December 2007).
RNA interference vectors: a modular series for multiple gene
down-regulation in plants". Plant Physiology. 145 (4): 1272–81.
doi:10.1104/pp.107.106062. PMC 2151715 .
^ Ruvkun G (October 2001). "Molecular biology. Glimpses of a tiny RNA
world". Science. 294 (5543): 797–9. doi:10.1126/science.1066315.
^ Fichou Y, Férec C (December 2006). "The potential of
oligonucleotides for therapeutic applications". Trends in
Biotechnology. 24 (12): 563–70. doi:10.1016/j.tibtech.2006.10.003.
^ Siebert S (2006). "Common sequence structure properties and stable
RNA secondary structures" (PDF). Dissertation,
Albert-Ludwigs-Universität, Freiburg im Breisgau. p. 1. Archived
from the original (PDF) on March 9, 2012.
^ Szathmáry E (June 1999). "The origin of the genetic code: amino
acids as cofactors in an
RNA world". Trends in Genetics. 15 (6):
223–9. doi:10.1016/S0168-9525(99)01730-8. PMID 10354582.
^ Marlaire R (3 March 2015). "
NASA Ames Reproduces the Building Blocks
Life in Laboratory". NASA. Retrieved 5 March 2015.
Wikiquote has quotations related to: RNA
Wikimedia Commons has media related to RNA.
RNA World website Link collection (structures, sequences, tools,
Nucleic Acid Database Images of DNA,
RNA and complexes.
Anna Marie Pyle's Seminar:
RNA Structure, Function, and Recognition
the British Isles
the Near East
List of genetics research organizations
DNA → RNA → Protein
Precursor mRNA (pre-mRNA / hnRNA)
Histone acetylation and deacetylation
Transfer RNA (tRNA)
Ribosome-nascent chain complex
Ribosome-nascent chain complex (RNC)
Post-translational modification (functional groups ·
peptides · structural changes)
Gene regulatory network
Types of RNA
Signal recognition particle RNA
Small nuclear RNA
Small nucleolar RNA
Cis-natural antisense transcript
Long noncoding RNA
Small interfering RNA
Small temporal RNA
Short hairpin RNA
Reverse transcribing virus
Types of nucleic acids
precursor, heterogenous nuclear
Small Cajal Body RNAs
Trans-acting small interfering
BNF: cb12175223g (data)
Molecular and cellu