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<p>fRNA</p>
<p>Functional RNA.</p>
<p>A functional<b> RNA</b><span style="color: #000000"> (f<b>RNA</b>) is a non-coding RNA molecule that is not translated into a protein. </span></p>
<p><span style="color: #000000">Less-frequently used synonyms are non-protein-coding RNA (npcRNA), non-messenger RNA (nmRNA), and small non-messenger RNA (snmRNA). The term <b>small RNA</b> (<b>sRNA</b>) is often used for small bacterial ncRNAs. The DNA sequence from which a non-coding RNA is transcribed as the end product is often called an <b>RNA gene</b> or non-coding RNA gene.</span></p>
<p><span style="color: #000000">Non-coding RNA genes include highly abundant and functionally important RNAs such as transfer RNA (tRNA) and ribosomal RNA (rRNA), as well as RNAs such as snoRNAs, microRNAs, siRNAs and piRNAs and the long ncRNAs that include examples such as Xist and HOTAIR (see here for a more complete list of ncRNAs). The number of ncRNAs encoded within the human genome is unknown, however recent transcriptomic and bioinformatic studies suggest the existence of thousands of ncRNAs.<sup id="cite_ref-pmid15790807_0-0" class="reference"><font size="2">[1]</font></sup><sup id="cite_ref-pmid17571346_1-0" class="reference"><font size="2">[2]</font></sup><sup id="cite_ref-pmid17568003_2-0" class="reference"><font size="2">[3]</font></sup><sup><font size="2">, but see</font></sup> <sup id="cite_ref-3" class="reference"><font size="2">[4]</font></sup> Since many of the newly identified ncRNAs have not been validated for their function, it is possible that many are non-functional.<sup id="cite_ref-pmid15851066_4-0" class="reference"><font size="2">[5]</font></sup></span></p>
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<h2><span style="color: #000000"><span id="History_and_discovery" class="mw-headline">History and discovery</span></span></h2>
<div class="rellink boilerplate further"><span style="color: #000000">Further information: History of molecular biology</span></div>
<p><span style="color: #000000">Nucleic acids were first discovered in 1868 by Friedrich Miescher<sup id="cite_ref-5" class="reference"><font size="2">[6]</font></sup> and by 1939 RNA had been implicated in protein synthesis.<sup id="cite_ref-6" class="reference"><font size="2">[7]</font></sup> Two decades later, Francis Crick predicted a functional RNA component which mediated translation; he reasoned that RNA is better suited to base-pair with the mRNA transcript than a pure polypeptide.<sup id="cite_ref-7" class="reference"><font size="2">[8]</font></sup></span></p>
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<div style="width: 252px" class="thumbinner"><span style="color: #000000"><font size="2"><img class="thumbimage" alt="" width="250" height="247" src="http://upload.wikimedia.org/wikipedia/commons/thumb/b/ba/TRNA-Phe_yeast_1ehz.png/250px-TRNA-Phe_yeast_1ehz.png" /></font> </span>
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<span style="color: #000000">The cloverleaf structure of Yeast tRNA<sup><font size="1">Phe</font></sup> (<i>inset</i>) and the 3D structure determined by X-ray analysis.</span></div>
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<p><span style="color: #000000">The first non-coding RNA to be characterised was an alanine tRNA found in baker's yeast, its structure was published in 1965.<sup id="cite_ref-Hol65_8-0" class="reference"><font size="2">[9]</font></sup> To produce a purified alanine tRNA sample, Robert W. Holley <i>et al.</i> used 140kg of commercial baker's yeast to give just 1g of purified tRNA<sup><font size="2">Ala</font></sup> for analysis.<sup id="cite_ref-Nobel68_9-0" class="reference"><font size="2">[10]</font></sup> The 80 nucleotide tRNA was sequenced by first being digested with Pancreatic ribonuclease (producing fragments ending in Cytosine or Uridine) and then with takadiastase ribonuclease Tl (producing fragments which finished with Guanosine). Chromatography and identification of the 5' and 3' ends then helped arrange the fragments to establish the RNA sequence.<sup id="cite_ref-Nobel68_9-1" class="reference"><font size="2">[10]</font></sup> Of the three structures originally proposed for this tRNA,<sup id="cite_ref-Hol65_8-1" class="reference"><font size="2">[9]</font></sup> the 'cloverleaf' structure was independently proposed in several following publications.<sup id="cite_ref-pmid5938777_10-0" class="reference"><font size="2">[11]</font></sup><sup id="cite_ref-11" class="reference"><font size="2">[12]</font></sup><sup id="cite_ref-12" class="reference"><font size="2">[13]</font></sup><sup id="cite_ref-13" class="reference"><font size="2">[14]</font></sup> The cloverleaf secondary structure was finalised following X-ray crystallography anaylsis performed by two independent research groups in 1974.<sup id="cite_ref-14" class="reference"><font size="2">[15]</font></sup><sup id="cite_ref-15" class="reference"><font size="2">[16]</font></sup></span></p>
<p><span style="color: #000000">Ribosomal RNA was next to be discovered, followed by URNA in the early 1980s. Since then, the discovery of new non-coding RNAs has continued with snoRNAs, Xist, CRISPR and many more.<sup id="cite_ref-Edd01_16-0" class="reference"><font size="2">[17]</font></sup> Recent notable additions include riboswitches and miRNA, the discovery of the RNAi mechanism associated with the latter earned Craig C. Mello and Andrew Fire the 2006 Nobel Prize in Physiology or Medicine.<sup id="cite_ref-17" class="reference"><font size="2">[18]</font></sup></span></p>
<h2><span style="color: #000000"><span id="Biological_roles_of_ncRNA" class="mw-headline">Biological roles of ncRNA</span></span></h2>
<p><span style="color: #000000">Noncoding RNAs belong to several groups and are involved in many cellular processes. These range from ncRNAs of central importance that are conserved across all or most cellular life through to more transient ncRNAs specific to one or a few closely related species. The more conserved ncRNAs are thought to be molecular fossils or relics from LUCA and the RNA world.<sup id="cite_ref-pmid9419222_18-0" class="reference"><font size="2">[19]</font></sup><sup id="cite_ref-pmid9419221_19-0" class="reference"><font size="2">[20]</font></sup><sup id="cite_ref-pmid10497339_20-0" class="reference"><font size="2">[21]</font></sup></span></p>
<h3><span style="color: #000000"><span id="ncRNAs_in_translation" class="mw-headline">ncRNAs in translation</span></span></h3>
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<div style="width: 402px" class="thumbinner"><span style="color: #000000"><img class="thumbimage" alt="" width="400" height="203" src="http://upload.wikimedia.org/wikipedia/commons/thumb/e/e6/NcRNAs-central-dogma.svg/400px-NcRNAs-central-dogma.svg.png" /> </span>
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<span style="color: #000000">An illustration of the central dogma of molecular biology annotated with the processes ncRNAs are involved in. RNPs are shown in red, ncRNAs are shown in blue.</span></div>
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<span style="color: #000000">Atomic structure of the 50S Subunit from <i>Haloarcula marismortui</i>. Proteins are shown in blue and the two RNA strands in orange and yellow.<sup id="cite_ref-Ban_21-0" class="reference"><font size="1">[22]</font></sup> The small patch of green in the center of the subunit is the active site.</span></div>
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<p><span style="color: #000000">Many of the conserved, essential and abundant ncRNAs are involved in translation. Ribonucleoprotein (RNP) particles called ribosomes are the 'factories' where translation takes place in the cell. The ribosome consists of more than 60% ribosomal RNA, these are made up of 3 ncRNAs in prokaryotes and 4 ncRNAs in eukaryotes. Ribosomal RNAs catalyse the translation of nucleotide sequences to protein. Another set of ncRNAs, Transfer RNAs, form an 'adaptor molecule' between mRNA and protein. The H/ACA box and C/D box snoRNAs are ncRNAs found in archaea and eukaryotes, RNase MRP is restricted to eukaryotes, both groups of ncRNA are involved in the maturation of rRNA. The snoRNAs guide covalent modifications of rRNA, tRNA and snRNAs, RNase MRP cleaves the internal transcribed spacer 1 between 18S and 5.8S rRNAs. The ubiquitous ncRNA, RNase P, is an evolutionary relative of RNase MRP.<sup id="cite_ref-PMID16540690_22-0" class="reference"><font size="2">[23]</font></sup> RNase P matures tRNA sequences by generating mature 5'-ends of tRNAs through cleaving the 5'-leader elements of precursor-tRNAs. Another ubiquitous RNP called SRP recognizes and transports specific nascent proteins to the endoplasmic reticulum in eukaryotes and the plasma membrane in prokaryotes. In bacteria Transfer-messenger RNA (tmRNA) is an RNP involved in rescuing stalled ribosomes, tagging incomplete polypeptides and promoting the degradation of aberrant mRNA.</span></p>
<h3><span style="color: #000000"><span id="ncRNAs_in_RNA_splicing" class="mw-headline">ncRNAs in RNA splicing</span></span></h3>
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<div style="width: 222px" class="thumbinner"><span style="color: #000000"><img class="thumbimage" alt="" width="220" height="160" src="http://upload.wikimedia.org/wikipedia/commons/thumb/1/18/Yeast_tri-snRNP.jpg/220px-Yeast_tri-snRNP.jpg" /> </span>
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<span style="color: #000000">Electron microscopy images of the yeast spliceosome. Note the bulk of the complex is in fact ncRNA.</span></div>
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<p><span style="color: #000000">In eukaryotes the spliceosome performs the splicing reactions essential for removing intron sequences, this process is required for the formation of mature mRNA. The spliceosome is another RNP often also known as the snRNP or tri-snRNP. There are two different forms of the spliceosome, the major and minor forms. The ncRNA components of the major spliceosome are U1, U2, U4 and U5. The ncRNA components of the minor spliceosome are U11, U12, U5, U4atac and U6atac.</span></p>
<p><span style="color: #000000">Another group of introns can catalyse their own removal from host transcripts, these are called self-splicing RNAs. There are two main groups of self-splicing RNAs, these are the group I catalytic intron and group II catalytic intron. These ncRNAs catalyze their own excision from mRNA, tRNA and rRNA precursors in a wide range of organisms.</span></p>
<p><span style="color: #000000">In mammals it has been found that snoRNAs can also regulate the alternative splicing of mRNA, for example snoRNA HBII-52 regulates the splicing of serotonin receptor 2C.<sup id="cite_ref-Kishore_23-0" class="reference"><font size="2">[24]</font></sup></span></p>
<p><span style="color: #000000">In nematodes the SmY ncRNA appears to be involved in mRNA trans-splicing.</span></p>
<h3><span style="color: #000000"><span id="ncRNAs_in_gene_regulation" class="mw-headline">ncRNAs in gene regulation</span></span></h3>
<p><span style="color: #000000">The expression of many thousands of genes are regulated by ncRNAs. This regulation can occur in trans or in cis.</span></p>
<h4><span style="color: #000000"><span id="trans-acting_ncRNAs" class="mw-headline">trans-acting ncRNAs</span></span></h4>
<p><span style="color: #000000">In higher eukaryotes microRNAs regulate gene expression. A single miRNA can reduce the expression levels of hundreds of genes. The mechanism by which mature miRNA molecules act is through partial complementary to one or more messenger RNA (mRNA) molecules, generally in 3' UTRs. The main function of miRNAs is to down-regulate gene expression.</span></p>
<p><span style="color: #000000">The ncRNA RNase P has also been shown to influence gene expression. In the human nucleus RNase P is required for the normal and efficient transcription of various ncRNAs transcribed by RNA polymerase III. These include tRNA, 5S rRNA, SRP RNA and U6 snRNA genes. RNase P exerts its role in transcription through association with Pol III and chromatin of active tRNA and 5S rRNA genes. <sup id="cite_ref-pmid16778078_24-0" class="reference"><font size="2">[25]</font></sup></span></p>
<p><span style="color: #000000">It has been shown that 7SK RNA, a metazoan ncRNA, acts as a negative regulator of the RNA polymerase II elongation factor P-TEFb, and that this activity is influenced by stress response pathways.</span></p>
<p><span style="color: #000000">The bacterial ncRNA, 6S RNA, specifically associates with RNA polymerase holoenzyme containing the sigma70 specificity factor. This interaction represses expression from a sigma70-dependent promoter during stationary phase.</span></p>
<p><span style="color: #000000">Another bacterial ncRNA, OxyS RNA represses translation by binding to Shine-Dalgarno sequences thereby occluding ribosome binding. OxyS RNA is induced in response to oxidative stress in Escherichia coli.</span></p>
<p><span style="color: #000000">The B2 RNA is a small noncoding RNA polymerase III transcript that represses mRNA transcription in response to heat shock in mouse cells. B2 RNA inhibits transcription by binding to core Pol II. Through this interaction, B2 RNA assembles into preinitiation complexes at the promoter and blocks RNA synthesis. <sup id="cite_ref-pmid15300239_25-0" class="reference"><font size="2">[26]</font></sup></span></p>
<p><span style="color: #000000">A recent study has shown that just the act of transcription of ncRNA sequence can have an influence on gene expression. RNA polymerase II transcription of ncRNAs is required for chromatin remodelling in the Schizosaccharomyces pombe. Chromatin is progressively converted to an open configuration, as several species of ncRNAs are transcribed. <sup id="cite_ref-pmid18820678_26-0" class="reference"><font size="2">[27]</font></sup></span></p>
<h4><span style="color: #000000"><span id="cis-acting_ncRNAs" class="mw-headline">cis-acting ncRNAs</span></span></h4>
<div class="rellink relarticle mainarticle"><span style="color: #000000">Main articles: Five prime untranslated region and Three prime untranslated region</span></div>
<p><span style="color: #000000">A number of ncRNAs are embedded in the 5' UTRs of protein coding genes and influence their expression in various ways. For example, a riboswitch can directly bind a small target molecule, the binding of the target affects the gene's activity.</span></p>
<p><span style="color: #000000">RNA leader sequences are found upstream of the first gene of in amino acid biosynthetic operons. These RNA elements form one of two possible structures in regions encoding very short peptide sequences that are rich in the end product amino acid of the operon. A terminator structure forms when there is an excess of the regulatory amino acid and ribosome movement over the leader transcript is not impeded. When there is a deficiency of the charged tRNA of the regulatory amino acid the ribosome translating the leader peptide stalls and the antiterminator structure forms. This allows RNA polymerase to transcribe the operon. Known RNA leaders are Histidine operon leader, Leucine operon leader, Threonine operon leader and the Tryptophan operon leader.</span></p>
<p><span style="color: #000000">Iron response elements (IRE) are bound by iron response proteins (IRP). The IRE is found in UTRs (Untranslated Regions) of various mRNAs whose products are involved in iron metabolism. When iron concentration is low, IRPs bind the ferritin mRNA IRE leading to translation repression.</span></p>
<p><span style="color: #000000">Internal ribosome entry sites (IRES) are a RNA structure that allow for translation initiation in the middle of a mRNA sequence as part of the process of protein synthesis.</span></p>
<h3><span style="color: #000000"><span id="ncRNAs_and_genome_defense" class="mw-headline">ncRNAs and genome defense</span></span></h3>
<p><span style="color: #000000">Piwi-interacting RNAs (piRNAs) expressed in mammalian testes and somatic cells, they form RNA-protein complexes with Piwi proteins. These piRNA complexes (piRCs) have been linked to transcriptional gene silencing of retrotransposons and other genetic elements in germ line cells, particularly those in spermatogenesis.</span></p>
<p><span style="color: #000000">Clustered Regularly Interspaced Short Palindromic Repeats (CRISPR) are repeats found in the DNA of many bacteria and archaea. The repeats are separated by spacers of similar length. It has been demonstrated that these spacers can be derived from phage and subsequently help protect the cell from infection.</span></p>
<h3><span style="color: #000000"><span id="ncRNAs_and_chromosome_structure" class="mw-headline">ncRNAs and chromosome structure</span></span></h3>
<p><span style="color: #000000">Telomerase is an RNP enzyme that adds specific DNA sequence repeats (&quot;TTAGGG&quot; in vertebrates) to telomeric regions, which are found at the ends of eukaryotic chromosomes. The telomeres contain condensed DNA material, giving stability to the chromosomes. The enzyme is a reverse transcriptase that carries Telomerase RNA, which is used as a template when it elongates telomeres, which are shortened after each replication cycle.</span></p>
<p><span style="color: #000000">Xist (X-inactive-specific transcript) is an long ncRNA gene on the X chromosome of the placental mammals that acts as major effector of the X chromosome inactivation process forming Barr bodies. An antisense RNA, Tsix, is a negative regulator of Xist. X chromosomes lacking Tsix expression (and thus having high levels of Xist transcription) are inactivated more frequently than normal chromosomes. In drosophilids, which also use an XY sex-determination system, the roX (RNA on the X) RNAs are involved in dosage compensation. <sup id="cite_ref-pmid12446910_27-0" class="reference"><font size="2">[28]</font></sup> Both Xist and roX operate by epigenetic regulation of transcription through the recruitment of histone-modifying enzymes.</span></p>
<h3><span style="color: #000000"><span id="Bifunctional_RNA" class="mw-headline">Bifunctional RNA</span></span></h3>
<p><span style="color: #000000"><b>Bifunctional RNAs</b> are RNAs that have two distinct functions, these are also known as dual function RNAs.<sup id="cite_ref-pmid18042713_28-0" class="reference"><font size="2">[29]</font></sup><sup id="cite_ref-pmid19043537_29-0" class="reference"><font size="2">[30]</font></sup> The majority of the known bifunctional RNAs are both mRNAs that encode a protein and ncRNAs. However there are also a growing number of ncRNAs that fall into two different ncRNA categories e.g. H/ACA box snoRNA and miRNA.<sup id="cite_ref-pmid19043559_30-0" class="reference"><font size="2">[31]</font></sup><sup id="cite_ref-pmid19026782_31-0" class="reference"><font size="2">[32]</font></sup></span></p>
<p><span style="color: #000000">Two well known examples of bifunctional RNAs are SgrS RNA and RNAIII. However, a handful of other bifunctional RNAs are known to exist, e.g. SRA (Steroid Receptor Activator) ,<sup id="cite_ref-pmid17710122_32-0" class="reference"><font size="2">[33]</font></sup> VegT RNA ,<sup id="cite_ref-pmid9012531_33-0" class="reference"><font size="2">[34]</font></sup><sup id="cite_ref-pmid16000384_34-0" class="reference"><font size="2">[35]</font></sup> Oskar RNA <sup id="cite_ref-pmid16835436_35-0" class="reference"><font size="2">[36]</font></sup> and ENOD40.<sup id="cite_ref-pmid17452360_36-0" class="reference"><font size="2">[37]</font></sup></span></p>
<h2><span style="color: #000000"><span id="ncRNAs_and_disease" class="mw-headline">ncRNAs and disease</span></span></h2>
<p><span style="color: #000000"><i>See also:</i> Long noncoding RNAs in disease</span></p>
<p><span style="color: #000000">As with proteins, mutations or imbalances in the ncRNA repertoire within the body can cause a variety of diseases.</span></p>
<h3><span style="color: #000000"><span id="Cancer" class="mw-headline">Cancer</span></span></h3>
<p><span style="color: #000000">Many ncRNAs show abnormal expression patterns in cancerous tissues. These include miRNAs,<sup id="cite_ref-pmid15944708_37-0" class="reference"><font size="2">[38]</font></sup> long mRNA-like ncRNAs ,<sup id="cite_ref-pmid11890990_38-0" class="reference"><font size="2">[39]</font></sup><sup id="cite_ref-pmid16569192_39-0" class="reference"><font size="2">[40]</font></sup> GAS5, <sup id="cite_ref-pmid18836484_40-0" class="reference"><font size="2">[41]</font></sup> SNORD50, <sup id="cite_ref-pmid19683667_41-0" class="reference"><font size="2">[42]</font></sup> telomerase RNA and Y RNAs. <sup id="cite_ref-pmid18283318_42-0" class="reference"><font size="2">[43]</font></sup> The miRNAs are involved in the large scale regulation of many protein coding genes,<sup id="cite_ref-pmid16308420_43-0" class="reference"><font size="2">[44]</font></sup><sup id="cite_ref-pmid15685193_44-0" class="reference"><font size="2">[45]</font></sup> the Y RNAs are important for the initiation of DNA replication,<sup id="cite_ref-pmid16943439_45-0" class="reference"><font size="2">[46]</font></sup> telomerase RNA that serves as a primer for telomerase, an RNP that extends telomeric regions at chromosome ends (see telomeres and disease for more information). The direct function of the long mRNA-like ncRNAs is less clear.</span></p>
<p><span style="color: #000000">Germ-line mutations in miR-16-1 and miR-15 primary precursors have been shown to be much more frequent in patients with chronic lymphocytic leukemia compared to control populations.<sup id="cite_ref-pmid16251535_46-0" class="reference"><font size="2">[47]</font></sup><sup id="cite_ref-47" class="reference"><font size="2">[48]</font></sup></span></p>
<p><span style="color: #000000">It has been suggested that a rare SNP (rs11614913) that overlaps hsa-mir-196a2 has been found to be associated with non-small cell lung carcinoma.<sup id="cite_ref-pmid18521189_48-0" class="reference"><font size="2">[49]</font></sup> Likewise, a screen of 17 miRNAs that have been predicted to regulate a number of breast cancer associated genes found variations in the microRNAs miR-17 and miR-30c-1, these patients were noncarriers of BRCA1 or BRCA2 mutations, lending the possibility that familial breast cancer may be caused by variation in these miRNAs.<sup id="cite_ref-pmid19048628_49-0" class="reference"><font size="2">[50]</font></sup></span></p>
<h3><span style="color: #000000"><span id="Prader.E2.80.93Willi_syndrome" class="mw-headline">Prader&ndash;Willi syndrome</span></span></h3>
<p><span style="color: #000000">The deletion of the 48 copies of the C/D box snoRNA SNORD116 has been shown to be the primary cause of Prader&ndash;Willi syndrome.<sup id="cite_ref-pmid18500341_50-0" class="reference"><font size="2">[51]</font></sup><sup id="cite_ref-pmid18320030_51-0" class="reference"><font size="2">[52]</font></sup><sup id="cite_ref-pmid16075369_52-0" class="reference"><font size="2">[53]</font></sup> Prader&ndash;Willi is a developmental disorder associated with over-eating and learning difficulties. SNORD116 has potential target sites within a number of protein-coding genes, and could have a role in regulating alternative splicing.<sup id="cite_ref-pmid18160232_53-0" class="reference"><font size="2">[54]</font></sup></span></p>
<h3><span style="color: #000000"><span id="Autism" class="mw-headline">Autism</span></span></h3>
<p><span style="color: #000000">The chromosomal locus containing the small nucleolar RNA SNORD115 gene cluster has been duplicated in approximately 5% of individuals with autistic traits.<sup id="cite_ref-pmid15318025_54-0" class="reference"><font size="2">[55]</font></sup> <sup id="cite_ref-pmid18923514_55-0" class="reference"><font size="2">[56]</font></sup> A mouse model engineered to have a duplication of the SNORD115 cluster displays autistic-like behaviour. <sup id="cite_ref-pmid19563756_56-0" class="reference"><font size="2">[57]</font></sup></span></p>
<h3><span style="color: #000000"><span id="Cartilage-hair_hypoplasia" class="mw-headline">Cartilage-hair hypoplasia</span></span></h3>
<p><span style="color: #000000">Mutations within RNase MRP have been shown to cause cartilage-hair hypoplasia, a disease associated with an array of symptoms such as short stature, sparse hair, skeletal abnormalities and a suppressed immune system that is frequent among Amish and Finnish.<sup id="cite_ref-pmid11207361_57-0" class="reference"><font size="2">[58]</font></sup><sup id="cite_ref-pmid17189938_58-0" class="reference"><font size="2">[59]</font></sup><sup id="cite_ref-pmid18804272_59-0" class="reference"><font size="2">[60]</font></sup> The best characterised variant is an A-to-G transition at nucleotide 70 that is in a loop region two bases 5' of a conserved pseudoknot. However, many other mutations within RNase MRP also cause CHH.</span></p>
<h3><span style="color: #000000"><span id="Alzheimer.27s_disease" class="mw-headline">Alzheimer's disease</span></span></h3>
<p><span style="color: #000000">The antisense RNA, BACE1-AS is transcribed from the opposite strand to BACE1 and is upregulated in patients with Alzheimer's disease.<sup id="cite_ref-pmid18587408_60-0" class="reference"><font size="2">[61]</font></sup> BACE1-AS regulates the expression of BACE1 by increasing BACE1 mRNA stability and generating additional BACE1 through a post-transcriptional feed-forward mechanism. By the same mechanism it also raises concentrations of beta amyloid, the main constituent of senile plaques. BACE1-AS concentrations are elevated in subjects with Alzheimer's disease and in amyloid precursor protein transgenic mice.</span></p>
<h3><span style="color: #000000"><span id="miR-96_and_hearing_loss" class="mw-headline">miR-96 and hearing loss</span></span></h3>
<p><span style="color: #000000">Variation within the seed region of mature miR-96 has been associated with autosomal dominant, progressive hearing loss in humans and mice. The homozygous mutant mice were profoundly deaf, showing no cochlear responses. Heterozygous mice and humans progressively lose the ability to hear. <sup id="cite_ref-pmid19363479_61-0" class="reference"><font size="2">[62]</font></sup> <sup id="cite_ref-pmid19363478_62-0" class="reference"><font size="2">[63]</font></sup> <sup id="cite_ref-pmid19245798_63-0" class="reference"><font size="2">[64]</font></sup></span></p>
<h2><span style="color: #000000"><span id="Distinction_between_functional_RNA_.28fRNA.29_and_ncRNA" class="mw-headline">Distinction between functional RNA (fRNA) and ncRNA</span></span></h2>
<p><span style="color: #000000">Several publications<sup id="cite_ref-64" class="reference"><font size="2">[65]</font></sup><sup id="cite_ref-65" class="reference"><font size="2">[66]</font></sup><sup id="cite_ref-66" class="reference"><font size="2">[67]</font></sup> have started using the term <b>functional RNA (fRNA)</b>, as opposed to ncRNA, to describe regions functional at the RNA level that may or may not be stand-alone RNA transcripts. Therefore, every ncRNA is a fRNA, but there exist fRNA (such as riboswitches, SECIS elements, and other cis-regulatory regions) that are not ncRNA. Yet the term fRNA could also include mRNA as this is RNA coding for protein and hence is functional. Additionally artificially evolved RNAs also fall under the fRNA umbrella term. Some publications<sup id="cite_ref-Edd01_16-1" class="reference"><font size="2">[17]</font></sup> state that the terms <i>ncRNA</i> and <i>fRNA</i> are nearly synonymous.</span></p>
<h2><span style="color: #000000"><span id="See_also" class="mw-headline">See also</span></span></h2>
<ul>
    <li><span style="color: #000000">List of RNAs</span></li>
    <li><span style="color: #000000">Nucleic acid structure</span></li>
    <li><span style="color: #000000">Rfam</span></li>
    <li><span style="color: #000000">Riboswitch</span></li>
    <li><span style="color: #000000">Ribozyme</span></li>
    <li><span style="color: #000000">RNAs present in environmental samples</span></li>
</ul>
<h2><span style="color: #000000"><span id="References" class="mw-headline">References</span></span></h2>
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</ol>
</div>
<h2><span id="External_links" class="mw-headline">External links</span></h2>
<ul>
    <li><a class="external text" rel="nofollow" href="http://jsm-research.imb.uq.edu.au/rnadb"><font color="#3366bb">Comprehensive database of mammalian ncRNAs</font></a></li>
    <li><a class="external text" rel="nofollow" href="http://rfam.sanger.ac.uk"><font color="#3366bb">The Rfam Database</font></a> &mdash; a curated list of hundreds of families of related ncRNAs.</li>
    <li><a class="external text" rel="nofollow" href="http://www.noncode.org/"><font color="#3366bb">NONCODE.org</font></a> &mdash; a free database of all kinds of noncoding RNAs (except tRNAs and rRNAs).</li>
    <li><a class="external text" rel="nofollow" href="http://www.ncrnadb.trna.ibch.poznan.pl/"><font color="#3366bb">Joint ncRNA Database</font></a> &mdash; over 30,000 individual sequences from 99 species of bacteria, archaea and eukaryota</li>
</ul>
<p>&nbsp;</p>