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data-event-name="pinnable-header.vector-toc.pin">move to sidebar</button> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-unpin-button" data-event-name="pinnable-header.vector-toc.unpin">hide</button> </div> <ul class="vector-toc-contents" id="mw-panel-toc-list"> <li id="toc-mw-content-text" class="vector-toc-list-item vector-toc-level-1"> <a href="#" class="vector-toc-link"> <div class="vector-toc-text">(Top)</div> </a> </li> <li id="toc-1930–1950" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#1930–1950"> <div class="vector-toc-text"> <span class="vector-toc-numb">1</span> <span>1930–1950</span> </div> </a> <button aria-controls="toc-1930–1950-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle 1930–1950 subsection</span> </button> <ul id="toc-1930–1950-sublist" class="vector-toc-list"> <li id="toc-RNA_and_DNA_have_distinct_chemical_properties" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#RNA_and_DNA_have_distinct_chemical_properties"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1</span> <span>RNA and DNA have distinct chemical properties</span> </div> </a> <ul id="toc-RNA_and_DNA_have_distinct_chemical_properties-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Localization_in_cell_and_morphogenetic_role" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Localization_in_cell_and_morphogenetic_role"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>Localization in cell and morphogenetic role</span> </div> </a> <ul id="toc-Localization_in_cell_and_morphogenetic_role-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-1951–1965" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#1951–1965"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>1951–1965</span> </div> </a> <button aria-controls="toc-1951–1965-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle 1951–1965 subsection</span> </button> <ul id="toc-1951–1965-sublist" class="vector-toc-list"> <li id="toc-Messenger_RNA_(mRNA)_carries_genetic_information_that_directs_protein_synthesis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Messenger_RNA_(mRNA)_carries_genetic_information_that_directs_protein_synthesis"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Messenger RNA (mRNA) carries genetic information that directs protein synthesis</span> </div> </a> <ul id="toc-Messenger_RNA_(mRNA)_carries_genetic_information_that_directs_protein_synthesis-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Ribosomes_make_proteins" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Ribosomes_make_proteins"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Ribosomes make proteins</span> </div> </a> <ul id="toc-Ribosomes_make_proteins-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Transfer_RNA_(tRNA)_is_the_physical_link_between_RNA_and_protein" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Transfer_RNA_(tRNA)_is_the_physical_link_between_RNA_and_protein"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Transfer RNA (tRNA) is the physical link between RNA and protein</span> </div> </a> <ul id="toc-Transfer_RNA_(tRNA)_is_the_physical_link_between_RNA_and_protein-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-The_genetic_code_is_solved" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#The_genetic_code_is_solved"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.4</span> <span>The genetic code is solved</span> </div> </a> <ul id="toc-The_genetic_code_is_solved-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-RNA_polymerase_is_purified" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#RNA_polymerase_is_purified"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.5</span> <span>RNA polymerase is purified</span> </div> </a> <ul id="toc-RNA_polymerase_is_purified-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-1966–1975" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#1966–1975"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>1966–1975</span> </div> </a> <button aria-controls="toc-1966–1975-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle 1966–1975 subsection</span> </button> <ul id="toc-1966–1975-sublist" class="vector-toc-list"> <li id="toc-First_complete_nucleotide_sequence_of_a_biological_nucleic_acid_molecule" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#First_complete_nucleotide_sequence_of_a_biological_nucleic_acid_molecule"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>First complete nucleotide sequence of a biological nucleic acid molecule</span> </div> </a> <ul id="toc-First_complete_nucleotide_sequence_of_a_biological_nucleic_acid_molecule-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Evolutionary_variation_of_homologous_RNA_sequences_reveals_folding_patterns" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Evolutionary_variation_of_homologous_RNA_sequences_reveals_folding_patterns"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Evolutionary variation of homologous RNA sequences reveals folding patterns</span> </div> </a> <ul id="toc-Evolutionary_variation_of_homologous_RNA_sequences_reveals_folding_patterns-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-First_complete_genomic_nucleotide_sequence" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#First_complete_genomic_nucleotide_sequence"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.3</span> <span>First complete genomic nucleotide sequence</span> </div> </a> <ul id="toc-First_complete_genomic_nucleotide_sequence-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Reverse_transcriptase_can_copy_RNA_into_DNA" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Reverse_transcriptase_can_copy_RNA_into_DNA"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.4</span> <span>Reverse transcriptase can copy RNA into DNA</span> </div> </a> <ul id="toc-Reverse_transcriptase_can_copy_RNA_into_DNA-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-RNA_replicons_evolve_rapidly" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#RNA_replicons_evolve_rapidly"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.5</span> <span>RNA replicons evolve rapidly</span> </div> </a> <ul id="toc-RNA_replicons_evolve_rapidly-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Ribosomal_RNA_(rRNA)_sequences_provide_a_record_of_the_evolutionary_history_of_all_life_forms" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Ribosomal_RNA_(rRNA)_sequences_provide_a_record_of_the_evolutionary_history_of_all_life_forms"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.6</span> <span>Ribosomal RNA (rRNA) sequences provide a record of the evolutionary history of all life forms</span> </div> </a> <ul id="toc-Ribosomal_RNA_(rRNA)_sequences_provide_a_record_of_the_evolutionary_history_of_all_life_forms-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Non-encoded_nucleotides_are_added_to_the_ends_of_RNA_molecules" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Non-encoded_nucleotides_are_added_to_the_ends_of_RNA_molecules"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.7</span> <span>Non-encoded nucleotides are added to the ends of RNA molecules</span> </div> </a> <ul id="toc-Non-encoded_nucleotides_are_added_to_the_ends_of_RNA_molecules-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-1976–1985" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#1976–1985"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>1976–1985</span> </div> </a> <button aria-controls="toc-1976–1985-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle 1976–1985 subsection</span> </button> <ul id="toc-1976–1985-sublist" class="vector-toc-list"> <li id="toc-Small_RNA_molecules_are_abundant_in_the_eukaryotic_nucleus" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Small_RNA_molecules_are_abundant_in_the_eukaryotic_nucleus"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Small RNA molecules are abundant in the eukaryotic nucleus</span> </div> </a> <ul id="toc-Small_RNA_molecules_are_abundant_in_the_eukaryotic_nucleus-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-RNA_molecules_require_a_specific,_complex_three-dimensional_structure_for_activity" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#RNA_molecules_require_a_specific,_complex_three-dimensional_structure_for_activity"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2</span> <span>RNA molecules require a specific, complex three-dimensional structure for activity</span> </div> </a> <ul id="toc-RNA_molecules_require_a_specific,_complex_three-dimensional_structure_for_activity-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Genes_are_commonly_interrupted_by_introns_that_must_be_removed_by_RNA_splicing" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genes_are_commonly_interrupted_by_introns_that_must_be_removed_by_RNA_splicing"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.3</span> <span>Genes are commonly interrupted by introns that must be removed by RNA splicing</span> </div> </a> <ul id="toc-Genes_are_commonly_interrupted_by_introns_that_must_be_removed_by_RNA_splicing-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Alternative_pre-mRNA_splicing_generates_multiple_proteins_from_a_single_gene" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Alternative_pre-mRNA_splicing_generates_multiple_proteins_from_a_single_gene"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.4</span> <span>Alternative pre-mRNA splicing generates multiple proteins from a single gene</span> </div> </a> <ul id="toc-Alternative_pre-mRNA_splicing_generates_multiple_proteins_from_a_single_gene-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Discovery_of_catalytic_RNA_(ribozymes)" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Discovery_of_catalytic_RNA_(ribozymes)"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.5</span> <span>Discovery of catalytic RNA (ribozymes)</span> </div> </a> <ul id="toc-Discovery_of_catalytic_RNA_(ribozymes)-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-RNA_was_likely_critical_for_prebiotic_evolution" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#RNA_was_likely_critical_for_prebiotic_evolution"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.6</span> <span>RNA was likely critical for prebiotic evolution</span> </div> </a> <ul id="toc-RNA_was_likely_critical_for_prebiotic_evolution-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Introns_can_be_mobile_genetic_elements" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Introns_can_be_mobile_genetic_elements"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.7</span> <span>Introns can be mobile genetic elements</span> </div> </a> <ul id="toc-Introns_can_be_mobile_genetic_elements-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Spliceosomes_mediate_nuclear_pre-mRNA_splicing" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Spliceosomes_mediate_nuclear_pre-mRNA_splicing"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.8</span> <span>Spliceosomes mediate nuclear pre-mRNA splicing</span> </div> </a> <ul id="toc-Spliceosomes_mediate_nuclear_pre-mRNA_splicing-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-1986–2000" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#1986–2000"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>1986–2000</span> </div> </a> <button aria-controls="toc-1986–2000-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle 1986–2000 subsection</span> </button> <ul id="toc-1986–2000-sublist" class="vector-toc-list"> <li id="toc-RNA_sequences_can_be_edited_within_cells" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#RNA_sequences_can_be_edited_within_cells"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1</span> <span>RNA sequences can be edited within cells</span> </div> </a> <ul id="toc-RNA_sequences_can_be_edited_within_cells-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Telomerase_uses_a_built-in_RNA_template_to_maintain_chromosome_ends" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Telomerase_uses_a_built-in_RNA_template_to_maintain_chromosome_ends"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2</span> <span>Telomerase uses a built-in RNA template to maintain chromosome ends</span> </div> </a> <ul id="toc-Telomerase_uses_a_built-in_RNA_template_to_maintain_chromosome_ends-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Ribosomal_RNA_catalyzes_peptide_bond_formation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Ribosomal_RNA_catalyzes_peptide_bond_formation"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.3</span> <span>Ribosomal RNA catalyzes peptide bond formation</span> </div> </a> <ul id="toc-Ribosomal_RNA_catalyzes_peptide_bond_formation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Combinatorial_selection_of_RNA_molecules_enables_in_vitro_evolution" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Combinatorial_selection_of_RNA_molecules_enables_in_vitro_evolution"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.4</span> <span>Combinatorial selection of RNA molecules enables in vitro evolution</span> </div> </a> <ul id="toc-Combinatorial_selection_of_RNA_molecules_enables_in_vitro_evolution-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-2001_–_present" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#2001_–_present"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>2001 – present</span> </div> </a> <button aria-controls="toc-2001_–_present-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle 2001 – present subsection</span> </button> <ul id="toc-2001_–_present-sublist" class="vector-toc-list"> <li id="toc-Many_mobile_DNA_elements_use_an_RNA_intermediate" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Many_mobile_DNA_elements_use_an_RNA_intermediate"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1</span> <span>Many mobile DNA elements use an RNA intermediate</span> </div> </a> <ul id="toc-Many_mobile_DNA_elements_use_an_RNA_intermediate-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Riboswitches_bind_cellular_metabolites_and_control_gene_expression" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Riboswitches_bind_cellular_metabolites_and_control_gene_expression"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.2</span> <span>Riboswitches bind cellular metabolites and control gene expression</span> </div> </a> <ul id="toc-Riboswitches_bind_cellular_metabolites_and_control_gene_expression-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Small_RNA_molecules_regulate_gene_expression_by_post-transcriptional_gene_silencing" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Small_RNA_molecules_regulate_gene_expression_by_post-transcriptional_gene_silencing"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.3</span> <span>Small RNA molecules regulate gene expression by post-transcriptional gene silencing</span> </div> </a> <ul id="toc-Small_RNA_molecules_regulate_gene_expression_by_post-transcriptional_gene_silencing-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Noncoding_RNA_controls_epigenetic_phenomena" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Noncoding_RNA_controls_epigenetic_phenomena"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.4</span> <span>Noncoding RNA controls epigenetic phenomena</span> </div> </a> <ul id="toc-Noncoding_RNA_controls_epigenetic_phenomena-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Nobel_Laureates_in_RNA_biology" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Nobel_Laureates_in_RNA_biology"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Nobel Laureates in RNA biology</span> </div> </a> <ul id="toc-Nobel_Laureates_in_RNA_biology-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> 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data-unpinned-container-id="vector-appearance-unpinned-container" > <div class="vector-pinnable-header-label">Appearance</div> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-pin-button" data-event-name="pinnable-header.vector-appearance.pin">move to sidebar</button> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-unpin-button" data-event-name="pinnable-header.vector-appearance.unpin">hide</button> </div> </div> </div> </nav> </div> </div> <div id="bodyContent" class="vector-body" aria-labelledby="firstHeading" data-mw-ve-target-container> <div class="vector-body-before-content"> <div class="mw-indicators"> </div> <div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><p> Numerous key discoveries in <a href="/wiki/Biology" title="Biology">biology</a> have emerged from studies of <a href="/wiki/RNA" title="RNA">RNA</a> (ribonucleic acid), including seminal work in the fields of <a href="/wiki/Biochemistry" title="Biochemistry">biochemistry</a>, <a href="/wiki/Genetics" title="Genetics">genetics</a>, <a href="/wiki/Microbiology" title="Microbiology">microbiology</a>, <a href="/wiki/Molecular_biology" title="Molecular biology">molecular biology</a>, <a href="/wiki/Molecular_evolution" title="Molecular evolution">molecular evolution</a>, and <a href="/wiki/Structural_biology" title="Structural biology">structural biology</a>. As of 2010, 30 scientists have been awarded <a href="/wiki/Nobel_Prize" title="Nobel Prize">Nobel Prizes</a> for experimental work that includes studies of RNA. Specific discoveries of high biological significance are discussed in this article. </p><p>For related information, see the articles on <a href="/wiki/History_of_molecular_biology" title="History of molecular biology">History of molecular biology</a> and <a href="/wiki/History_of_genetics" title="History of genetics">History of genetics</a>. For background information, see the articles on <a href="/wiki/RNA" title="RNA">RNA</a> and <a href="/wiki/Nucleic_acid" title="Nucleic acid">nucleic acids</a>. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="1930–1950"><span id="1930.E2.80.931950"></span>1930–1950</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=1" title="Edit section: 1930–1950"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="RNA_and_DNA_have_distinct_chemical_properties">RNA and DNA have distinct chemical properties</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=2" title="Edit section: RNA and DNA have distinct chemical properties"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>When first studied in the early 1900s, the chemical and biological differences between RNA and DNA were not apparent, and they were named after the materials from which they were isolated; RNA was initially known as "<a href="/wiki/Yeast" title="Yeast">yeast</a> nucleic acid" and DNA was "<a href="/wiki/Thymus" title="Thymus">thymus</a> nucleic acid".<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Using diagnostic chemical tests, <a href="/wiki/Carbohydrate" title="Carbohydrate">carbohydrate</a> chemists showed that the two nucleic acids contained different <a href="/wiki/Sugars" class="mw-redirect" title="Sugars">sugars</a>, whereupon the common name for RNA became "ribose nucleic acid". Other early biochemical studies showed that RNA was readily broken down at high <a href="/wiki/PH" title="PH">pH</a>, while DNA was stable (although denatured) in <a href="/wiki/Alkali" title="Alkali">alkali</a>. Nucleoside composition analysis showed first that RNA contained similar <a href="/wiki/Nucleobase" class="mw-redirect" title="Nucleobase">nucleobases</a> to DNA, with <a href="/wiki/Uracil" title="Uracil">uracil</a> instead of <a href="/wiki/Thymine" title="Thymine">thymine</a>, and that RNA contained a number of minor nucleobase components, e.g. small amounts of <a href="/wiki/Pseudouridine" title="Pseudouridine">pseudouridine</a> and <a href="/w/index.php?title=Dimethylguanine&action=edit&redlink=1" class="new" title="Dimethylguanine (page does not exist)">dimethylguanine</a>.<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Localization_in_cell_and_morphogenetic_role">Localization in cell and morphogenetic role</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=3" title="Edit section: Localization in cell and morphogenetic role"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 1933, while studying virgin <a href="/wiki/Sea_urchin" title="Sea urchin">sea urchin</a> eggs, <a href="/wiki/Jean_Brachet" title="Jean Brachet">Jean Brachet</a> suggested that <a href="/wiki/DNA" title="DNA">DNA</a> is found in <a href="/wiki/Cell_nucleus" title="Cell nucleus">cell nucleus</a> and that <a href="/wiki/RNA" title="RNA">RNA</a> is present exclusively in the <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a>. At the time, "yeast nucleic acid" (RNA) was thought to occur only in plants, while "thymus nucleic acid" (DNA) only in animals. The latter was thought to be a tetramer, with the function of buffering cellular pH.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> During the 1930s, <a href="/wiki/Joachim_H%C3%A4mmerling" title="Joachim Hämmerling">Joachim Hämmerling</a> conducted experiments with <i><a href="/wiki/Acetabularia" title="Acetabularia">Acetabularia</a></i> in which he began to distinguish the contributions of the nucleus and the cytoplasm substances (later discovered to be DNA and mRNA, respectively) to cell morphogenesis and development.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="1951–1965"><span id="1951.E2.80.931965"></span>1951–1965</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=4" title="Edit section: 1951–1965"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Messenger_RNA_(mRNA)_carries_genetic_information_that_directs_protein_synthesis"><span id="Messenger_RNA_.28mRNA.29_carries_genetic_information_that_directs_protein_synthesis"></span>Messenger RNA (mRNA) carries genetic information that directs protein synthesis</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=5" title="Edit section: Messenger RNA (mRNA) carries genetic information that directs protein synthesis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The concept of messenger RNA emerged during the late 1950s, and is associated with <a href="/wiki/Francis_Crick" title="Francis Crick">Crick</a>'s description of his "Central Dogma of Molecular Biology", which asserted that DNA led to the formation of RNA, which in turn led to the synthesis of <a href="/wiki/Protein" title="Protein">proteins</a>. During the early 1960s, sophisticated genetic analysis of mutations in the <a href="/wiki/Lac_operon" title="Lac operon">lac operon</a> of <a href="/wiki/Escherichia_coli" title="Escherichia coli"><i>E. coli</i></a> and in the rII locus of <a href="/wiki/Enterobacteria_phage_T4" class="mw-redirect" title="Enterobacteria phage T4">bacteriophage T4</a> were instrumental in defining the nature of both <a href="/wiki/Messenger_RNA" title="Messenger RNA">messenger RNA</a> and the <a href="/wiki/Genetic_code" title="Genetic code">genetic code</a>. The short-lived nature of bacterial RNAs, together with the highly complex nature of the cellular mRNA population, made the biochemical isolation of mRNA very challenging. This problem was overcome in the 1960s by the use of <a href="/wiki/Reticulocyte" title="Reticulocyte">reticulocytes</a> in vertebrates,<sup id="cite_ref-pmid16590302_7-0" class="reference"><a href="#cite_note-pmid16590302-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> which produce large quantities of mRNA that are highly enriched in RNA encoding alpha- and beta-globin (the two major protein chains of <a href="/wiki/Hemoglobin" title="Hemoglobin">hemoglobin</a>).<sup id="cite_ref-pmid|4896247_8-0" class="reference"><a href="#cite_note-pmid|4896247-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> The first direct experimental evidence for the existence of mRNA was provided by such a hemoglobin synthesizing system.<sup id="cite_ref-pmid13758530_9-0" class="reference"><a href="#cite_note-pmid13758530-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Ribosomes_make_proteins">Ribosomes make proteins</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=6" title="Edit section: Ribosomes make proteins"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In the 1950s, results of labeling experiments in rat liver showed that radioactive <a href="/wiki/Amino_acid" title="Amino acid">amino acids</a> were found to be associated with "microsomes" (later redefined as <a href="/wiki/Ribosomes" class="mw-redirect" title="Ribosomes">ribosomes</a>) very rapidly after administration, and before they became widely incorporated into cellular proteins. Ribosomes were first visualized using <a href="/wiki/Electron_microscopy" class="mw-redirect" title="Electron microscopy">electron microscopy</a>, and their ribonucleoprotein components were identified by biophysical methods, chiefly sedimentation analysis within <a href="/wiki/Ultracentrifuges" class="mw-redirect" title="Ultracentrifuges">ultracentrifuges</a> capable of generating very high accelerations (equivalent to hundreds of thousands times gravity). <a href="/wiki/Polysomes" class="mw-redirect" title="Polysomes">Polysomes</a> (multiple ribosomes moving along a single mRNA molecule) were identified in the early 1960s, and their study led to an understanding of how ribosomes proceed to read the mRNA in a 5′ to 3′ direction,<sup id="cite_ref-pmid5339691_10-0" class="reference"><a href="#cite_note-pmid5339691-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> processively generating proteins as they do so.<sup id="cite_ref-pmid|5329473_11-0" class="reference"><a href="#cite_note-pmid|5329473-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Transfer_RNA_(tRNA)_is_the_physical_link_between_RNA_and_protein"><span id="Transfer_RNA_.28tRNA.29_is_the_physical_link_between_RNA_and_protein"></span>Transfer RNA (tRNA) is the physical link between RNA and protein</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=7" title="Edit section: Transfer RNA (tRNA) is the physical link between RNA and protein"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Biochemical fractionation experiments showed that radioactive amino acids were rapidly incorporated into small RNA molecules that remained soluble under conditions where larger RNA-containing particles would precipitate. These molecules were termed soluble (sRNA) and were later renamed transfer RNA (<a href="/wiki/TRNA" class="mw-redirect" title="TRNA">tRNA</a>). Subsequent studies showed that (i) every cell has multiple species of tRNA, each of which is associated with a single specific amino acid, (ii) that there are a matching set of <a href="/wiki/Enzyme" title="Enzyme">enzymes</a> responsible for linking tRNAs with the correct amino acids, and (iii) that tRNA <a href="/wiki/Anticodon" class="mw-redirect" title="Anticodon">anticodon</a> sequences form a specific decoding interaction with mRNA <a href="/wiki/Codons" class="mw-redirect" title="Codons">codons</a>.<sup id="cite_ref-pmid|60910_12-0" class="reference"><a href="#cite_note-pmid|60910-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="The_genetic_code_is_solved">The genetic code is solved</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=8" title="Edit section: The genetic code is solved"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <a href="/wiki/Genetic_code" title="Genetic code">genetic code</a> consists of the translation of particular <a href="/wiki/Nucleotide_sequence" class="mw-redirect" title="Nucleotide sequence">nucleotide sequences</a> in mRNA to specific amino acid sequences in <a href="/wiki/Proteins" class="mw-redirect" title="Proteins">proteins</a> (polypeptides). The ability to work out the genetic code emerged from the convergence of three different areas of study: (i) new methods to generate synthetic RNA molecules of defined composition to serve as artificial mRNAs, (ii) development of <i>in vitro</i> translation systems that could be used to translate the synthetic mRNAs into protein, and (iii) experimental and theoretical genetic work which established that the code was written in three letter "words" (<a href="/wiki/Codons" class="mw-redirect" title="Codons">codons</a>). Today, our understanding of the genetic code permits the prediction of the amino sequence of the protein products of the tens of thousands of genes whose sequences are being determined in <a href="/wiki/Genome" title="Genome">genome</a> studies.<sup id="cite_ref-pmid|5322508_13-0" class="reference"><a href="#cite_note-pmid|5322508-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="RNA_polymerase_is_purified">RNA polymerase is purified</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=9" title="Edit section: RNA polymerase is purified"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The biochemical purification and characterization of <a href="/wiki/RNA_polymerase" title="RNA polymerase">RNA polymerase</a> from the bacterium <i><a href="/wiki/Escherichia_coli" title="Escherichia coli">Escherichia coli</a></i> enabled the understanding of the mechanisms through which RNA polymerase initiates and terminates <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcription</a>, and how those processes are regulated to regulate <a href="/wiki/Gene_expression" title="Gene expression">gene expression</a> (i.e. turn genes on and off). Following the isolation of <i>E. coli</i> RNA polymerase, the three RNA polymerases of the eukaryotic nucleus were identified, as well as those associated with viruses and organelles. Studies of transcription also led to the identification of many protein factors that influence transcription, including repressors, activators and enhancers. The availability of purified preparations of RNA polymerase permitted investigators to develop a wide range of novel methods for studying RNA in the test tube, and led directly to many of the subsequent key discoveries in RNA biology.<sup id="cite_ref-pmid|5001045_14-0" class="reference"><a href="#cite_note-pmid|5001045-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="1966–1975"><span id="1966.E2.80.931975"></span>1966–1975</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=10" title="Edit section: 1966–1975"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="First_complete_nucleotide_sequence_of_a_biological_nucleic_acid_molecule">First complete nucleotide sequence of a biological nucleic acid molecule</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=11" title="Edit section: First complete nucleotide sequence of a biological nucleic acid molecule"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Although determining the sequence of proteins was becoming somewhat routine, methods for sequencing of nucleic acids were not available until the mid-1960s. In this seminal work, a specific tRNA was purified in substantial quantities, and then sliced into overlapping fragments using a variety of ribonucleases. Analysis of the detailed nucleotide composition of each fragment provided the information necessary to deduce the sequence of the tRNA. Today, the sequence analysis of much larger nucleic acid molecules is highly automated and enormously faster.<sup id="cite_ref-pmid|4875713_15-0" class="reference"><a href="#cite_note-pmid|4875713-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Evolutionary_variation_of_homologous_RNA_sequences_reveals_folding_patterns">Evolutionary variation of homologous RNA sequences reveals folding patterns</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=12" title="Edit section: Evolutionary variation of homologous RNA sequences reveals folding patterns"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Additional tRNA molecules were purified and sequenced. The first comparative sequence analysis was done and revealed that the sequences varied through evolution in such a way that all of the tRNAs could fold into very similar secondary structures (two-dimensional structures) and had identical sequences at numerous positions (e.g. CCA at the 3′ end). The radial four-arm structure of tRNA molecules is termed the 'cloverleaf structure', and results from the evolution of sequences with common ancestry and common biological function. Since the discovery of the tRNA cloverleaf, comparative analysis of numerous other homologous RNA molecules has led to the identification of common sequences and folding patterns.<sup id="cite_ref-pmid|6163215_16-0" class="reference"><a href="#cite_note-pmid|6163215-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="First_complete_genomic_nucleotide_sequence">First complete genomic nucleotide sequence</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=13" title="Edit section: First complete genomic nucleotide sequence"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The 3569 nucleotide sequence of all of the genes of the RNA <a href="/wiki/Bacteriophage" title="Bacteriophage">bacteriophage</a> <a href="/wiki/MS2_phage" class="mw-redirect" title="MS2 phage">MS2</a> was determined by a large team of researchers over several years, and was reported in a series of scientific papers. These results enabled the analysis of the first complete genome, albeit an extremely tiny one by modern standards. Several surprising features were identified, including genes that partially overlap one another and the first clues that different organisms might have slightly different codon usage patterns.<sup id="cite_ref-pmid|1264203_17-0" class="reference"><a href="#cite_note-pmid|1264203-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Reverse_transcriptase_can_copy_RNA_into_DNA">Reverse transcriptase can copy RNA into DNA</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=14" title="Edit section: Reverse transcriptase can copy RNA into DNA"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Retroviruses were shown to have a single-stranded RNA genome and to replicate via a DNA intermediate, the reverse of the usual DNA-to-RNA transcription pathway. They encode a RNA-dependent DNA polymerase (<a href="/wiki/Reverse_transcriptase" title="Reverse transcriptase">reverse transcriptase</a>) that is essential for this process. Some retroviruses can cause diseases, including several that are associated with cancer, and HIV-1 which causes AIDS. Reverse transcriptase has been widely used as an experimental tool for the analysis of RNA molecules in the laboratory, in particular the conversion of RNA molecules into DNA prior to <a href="/wiki/Molecular_cloning" title="Molecular cloning">molecular cloning</a> and/or <a href="/wiki/Polymerase_chain_reaction" title="Polymerase chain reaction">polymerase chain reaction</a> (PCR).<sup id="cite_ref-pmid|9759480_18-0" class="reference"><a href="#cite_note-pmid|9759480-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="RNA_replicons_evolve_rapidly">RNA replicons evolve rapidly</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=15" title="Edit section: RNA replicons evolve rapidly"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Biochemical and genetic analyses showed that the enzyme systems that replicate viral RNA molecules (reverse transcriptases and RNA replicases) lack molecular proofreading (3′ to 5′ exonuclease) activity, and that RNA sequences do not benefit from extensive repair systems analogous to those that exist for maintaining and repairing DNA sequences. Consequently, RNA genomes appear to be subject to significantly higher mutation rates than DNA genomes. For example, mutations in HIV-1 that lead to the emergence of viral mutants that are insensitive to antiviral drugs are common, and constitute a major clinical challenge.<sup id="cite_ref-pmid|20949639_19-0" class="reference"><a href="#cite_note-pmid|20949639-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Ribosomal_RNA_(rRNA)_sequences_provide_a_record_of_the_evolutionary_history_of_all_life_forms"><span id="Ribosomal_RNA_.28rRNA.29_sequences_provide_a_record_of_the_evolutionary_history_of_all_life_forms"></span>Ribosomal RNA (rRNA) sequences provide a record of the evolutionary history of all life forms</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=16" title="Edit section: Ribosomal RNA (rRNA) sequences provide a record of the evolutionary history of all life forms"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Analysis of <a href="/wiki/Ribosomal_RNA" title="Ribosomal RNA">ribosomal RNA</a> sequences from a large number of organisms demonstrated that all extant forms of life on Earth share common structural and sequence features of the ribosomal RNA, reflecting a <a href="/wiki/Last_common_ancestor" class="mw-redirect" title="Last common ancestor">common ancestry</a>. Mapping the similarities and differences among rRNA molecules from different sources provides clear and quantitative information about the <a href="/wiki/Phylogenetic" class="mw-redirect" title="Phylogenetic">phylogenetic</a> (i.e. evolutionary) relationships among organisms. Analysis of rRNA molecules led to the identification of a third major kingdom of organisms, the <a href="/wiki/Archaea" title="Archaea">archaea</a>, in addition to the <a href="/wiki/Prokaryotes" class="mw-redirect" title="Prokaryotes">prokaryotes</a> and <a href="/wiki/Eukaryotes" class="mw-redirect" title="Eukaryotes">eukaryotes</a>.<sup id="cite_ref-pmid|10900003_20-0" class="reference"><a href="#cite_note-pmid|10900003-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Non-encoded_nucleotides_are_added_to_the_ends_of_RNA_molecules">Non-encoded nucleotides are added to the ends of RNA molecules</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=17" title="Edit section: Non-encoded nucleotides are added to the ends of RNA molecules"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Molecular analysis of mRNA molecules showed that, following transcription, mRNAs have non-DNA-encoded nucleotides added to both their 5′ and 3′ ends (guanosine caps and poly-A, respectively). Enzymes were also identified that add and maintain the universal CCA sequence on the 3′ end of tRNA molecules. These events are among the first discovered examples of <a href="/wiki/RNA_processing" class="mw-redirect" title="RNA processing">RNA processing</a>, a complex series of reactions that are needed to convert RNA primary transcripts into biologically active RNA molecules.<sup id="cite_ref-pmid|1353951_21-0" class="reference"><a href="#cite_note-pmid|1353951-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="1976–1985"><span id="1976.E2.80.931985"></span>1976–1985</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=18" title="Edit section: 1976–1985"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Small_RNA_molecules_are_abundant_in_the_eukaryotic_nucleus">Small RNA molecules are abundant in the eukaryotic nucleus</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=19" title="Edit section: Small RNA molecules are abundant in the eukaryotic nucleus"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Small_nuclear_RNA" title="Small nuclear RNA">Small nuclear RNA</a> molecules (snRNAs) were identified in the eukaryotic <a href="/wiki/Cell_nucleus" title="Cell nucleus">nucleus</a> using immunological studies with autoimmune <a href="/wiki/Antibodies" class="mw-redirect" title="Antibodies">antibodies</a>, which bind to <a href="/wiki/SnRNP" title="SnRNP">small nuclear ribonucleoprotein</a> complexes (snRNPs; complexes of the snRNA and protein). Subsequent biochemical, genetic, and phylogenetic studies established that many of these molecules play key roles in essential <a href="/wiki/RNA_processing" class="mw-redirect" title="RNA processing">RNA processing</a> reactions within the nucleus and <a href="/wiki/Nucleolus" title="Nucleolus">nucleolus</a>, including <a href="/wiki/RNA_splicing" title="RNA splicing">RNA splicing</a>, <a href="/wiki/Polyadenylation" title="Polyadenylation">polyadenylation</a>, and the maturation of <a href="/wiki/Ribosomal_RNA" title="Ribosomal RNA">ribosomal RNAs</a>.<sup id="cite_ref-pmid|6180681_22-0" class="reference"><a href="#cite_note-pmid|6180681-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="RNA_molecules_require_a_specific,_complex_three-dimensional_structure_for_activity"><span id="RNA_molecules_require_a_specific.2C_complex_three-dimensional_structure_for_activity"></span>RNA molecules require a specific, complex three-dimensional structure for activity</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=20" title="Edit section: RNA molecules require a specific, complex three-dimensional structure for activity"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The detailed three-dimensional structure of <a href="/wiki/TRNA" class="mw-redirect" title="TRNA">tRNA</a> molecules was determined using <a href="/wiki/X-ray_crystallography" title="X-ray crystallography">X-ray crystallography</a>, and revealed highly complex, compact three dimensional structures consisting of tertiary interactions laid upon the basic cloverleaf secondary structure. Key features of tRNA tertiary structure include the coaxial stacking of adjacent helices and non-Watson-Crick interactions among nucleotides within the apical loops. Additional crystallographic studies showed that a wide range of RNA molecules (including <a href="/wiki/Ribozymes" class="mw-redirect" title="Ribozymes">ribozymes</a>, <a href="/wiki/Riboswitches" class="mw-redirect" title="Riboswitches">riboswitches</a> and <a href="/wiki/Ribosomal_RNA" title="Ribosomal RNA">ribosomal RNA</a>) also fold into specific structures containing a variety of 3D structural motifs. The ability of RNA molecules to adopt specific tertiary structures is essential for their biological activity, and results from the single-stranded nature of RNA. In many ways, RNA folding is more highly analogous to the folding of proteins rather than to the highly repetitive folded structure of the DNA double helix.<sup id="cite_ref-pmid|60910_12-1" class="reference"><a href="#cite_note-pmid|60910-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Genes_are_commonly_interrupted_by_introns_that_must_be_removed_by_RNA_splicing">Genes are commonly interrupted by introns that must be removed by RNA splicing</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=21" title="Edit section: Genes are commonly interrupted by introns that must be removed by RNA splicing"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Analysis of mature eukaryotic <a href="/wiki/Messenger_RNA" title="Messenger RNA">messenger RNA</a> molecules showed that they are often much smaller than the DNA sequences that encode them. The genes were shown to be discontinuous, composed of sequences that are not present in the final mature RNA (<a href="/wiki/Introns" class="mw-redirect" title="Introns">introns</a>), located between sequences that are retained in the mature RNA (<a href="/wiki/Exons" class="mw-redirect" title="Exons">exons</a>). Introns were shown to be removed after transcription through a process termed <a href="/wiki/RNA_splicing" title="RNA splicing">RNA splicing</a>. Splicing of RNA transcripts requires a highly precise and coordinated sequence of molecular events, consisting of (a) definition of boundaries between exons and introns, (b) RNA strand cleavage at exactly those sites, and (c) covalent linking (ligation) of the RNA exons in the correct order. The discovery of discontinuous genes and RNA splicing was entirely unexpected by the community of RNA biologists, and stands as one of the most shocking findings in molecular biology research.<sup id="cite_ref-pmid|2880558_23-0" class="reference"><a href="#cite_note-pmid|2880558-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Alternative_pre-mRNA_splicing_generates_multiple_proteins_from_a_single_gene">Alternative pre-mRNA splicing generates multiple proteins from a single gene</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=22" title="Edit section: Alternative pre-mRNA splicing generates multiple proteins from a single gene"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The great majority of protein-coding genes encoded within the nucleus of <a href="/wiki/Metazoan" class="mw-redirect" title="Metazoan">metazoan</a> cells contain multiple <a href="/wiki/Introns" class="mw-redirect" title="Introns">introns</a>. In many cases, these introns were shown to be processed in more than one pattern, thus generating a family of related mRNAs that differ, for example, by the inclusion or exclusion of particular exons. The result of <a href="/wiki/Alternative_splicing" title="Alternative splicing">alternative splicing</a> is that a single <a href="/wiki/Gene" title="Gene">gene</a> can encode a number of different protein <a href="/wiki/Isoforms" class="mw-redirect" title="Isoforms">isoforms</a> that can exhibit a variety of (usually related) biological functions. Indeed, most of the proteins encoded by the human genome are generated by alternative splicing.<sup id="cite_ref-pmid|3304142_24-0" class="reference"><a href="#cite_note-pmid|3304142-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Discovery_of_catalytic_RNA_(ribozymes)"><span id="Discovery_of_catalytic_RNA_.28ribozymes.29"></span>Discovery of catalytic RNA (ribozymes)</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=23" title="Edit section: Discovery of catalytic RNA (ribozymes)"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>An experimental system was developed in which an intron-containing rRNA precursor from the nucleus of the ciliated protozoan <i><a href="/wiki/Tetrahymena" title="Tetrahymena">Tetrahymena</a></i> could be spliced <i><a href="/wiki/In_vitro" title="In vitro">in vitro</a></i>. Subsequent biochemical analysis shows that this <a href="/wiki/Group_I_intron" class="mw-redirect" title="Group I intron">group I intron</a> was self-splicing; that is, the precursor RNA is capable of carrying out the complete splicing reaction in the absence of proteins. In separate work, the RNA component of the bacterial enzyme <a href="/wiki/Ribonuclease_P" title="Ribonuclease P">ribonuclease P</a> (a <a href="/wiki/Ribonucleoprotein" class="mw-redirect" title="Ribonucleoprotein">ribonucleoprotein</a> complex) was shown to catalyze its tRNA-processing reaction in the absence of proteins. These experiments represented landmarks in RNA biology, since they revealed that RNA could play an active role in cellular processes, by catalyzing specific biochemical reactions. Before these discoveries, it was believed that biological catalysis was solely the realm of <a href="/wiki/Protein" title="Protein">protein</a> <a href="/wiki/Enzymes" class="mw-redirect" title="Enzymes">enzymes</a>.<sup id="cite_ref-pmid|2197983_25-0" class="reference"><a href="#cite_note-pmid|2197983-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid|9759486_26-0" class="reference"><a href="#cite_note-pmid|9759486-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="RNA_was_likely_critical_for_prebiotic_evolution">RNA was likely critical for prebiotic evolution</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=24" title="Edit section: RNA was likely critical for prebiotic evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The discovery of catalytic RNA (<a href="/wiki/Ribozymes" class="mw-redirect" title="Ribozymes">ribozymes</a>) showed that RNA could both encode genetic information (like DNA) and catalyze specific biochemical reactions (like protein <a href="/wiki/Enzymes" class="mw-redirect" title="Enzymes">enzymes</a>). This realization led to the <a href="/wiki/RNA_World_Hypothesis" class="mw-redirect" title="RNA World Hypothesis">RNA World Hypothesis</a>, a proposal that RNA may have played a critical role in <a href="/wiki/Prebiotic_evolution" class="mw-redirect" title="Prebiotic evolution">prebiotic evolution</a> at a time before the molecules with more specialized functions (DNA and proteins) came to dominate biological information coding and catalysis. Although it is not possible for us to know the course of prebiotic evolution with any certainty, the presence of functional RNA molecules with common ancestry in all modern-day life forms is a strong argument that RNA was widely present at the time of the <a href="/wiki/Last_common_ancestor" class="mw-redirect" title="Last common ancestor">last common ancestor</a>.<sup id="cite_ref-pmid|2466202_27-0" class="reference"><a href="#cite_note-pmid|2466202-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Introns_can_be_mobile_genetic_elements">Introns can be mobile genetic elements</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=25" title="Edit section: Introns can be mobile genetic elements"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Some self-splicing introns can spread through a population of organisms by "homing", inserting copies of themselves into genes at sites that previously lacked an intron. Because they are self-splicing (that is, they remove themselves at the RNA level from genes into which they have inserted), these sequences represent <a href="/wiki/Transposons" class="mw-redirect" title="Transposons">transposons</a> that are genetically silent, i.e. they do not interfere with the expression of the gene into which they become inserted. These introns can be regarded as examples of <a href="/wiki/Selfish_DNA" class="mw-redirect" title="Selfish DNA">selfish DNA</a>. Some mobile introns encode <a href="/wiki/Homing_endonucleases" class="mw-redirect" title="Homing endonucleases">homing endonucleases</a>, enzymes that initiate the homing process by specifically cleaving double-stranded DNA at or near the intron-insertion site of alleles lacking an intron. Mobile introns are frequently members of either the <a href="/wiki/Group_I_intron" class="mw-redirect" title="Group I intron">group I</a> or <a href="/wiki/Group_II_intron" title="Group II intron">group II</a> families of self-splicing introns.<sup id="cite_ref-pmid|8352597_28-0" class="reference"><a href="#cite_note-pmid|8352597-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Spliceosomes_mediate_nuclear_pre-mRNA_splicing">Spliceosomes mediate nuclear pre-mRNA splicing</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=26" title="Edit section: Spliceosomes mediate nuclear pre-mRNA splicing"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Introns are removed from nuclear pre-mRNAs by <a href="/wiki/Spliceosome" title="Spliceosome">spliceosomes</a>, large <a href="/wiki/Ribonucleoprotein" class="mw-redirect" title="Ribonucleoprotein">ribonucleoprotein</a> complexes made up of <a href="/wiki/SnRNA" class="mw-redirect" title="SnRNA">snRNA</a> and protein molecules whose composition and molecular interactions change during the course of the <a href="/wiki/RNA_splicing" title="RNA splicing">RNA splicing</a> reactions. Spliceosomes assemble on and around splice sites (the boundaries between introns and exons in the unspliced pre-mRNA) in mRNA precursors and use RNA-RNA interactions to identify critical nucleotide sequences and, probably, to catalyze the splicing reactions. Nuclear pre-mRNA introns and spliceosome-associated snRNAs show similar structural features to self-splicing group II introns. In addition, the splicing pathway of nuclear pre-mRNA introns and group II introns shares a similar reaction pathway. These similarities have led to the hypothesis that these molecules may share a common ancestor.<sup id="cite_ref-pmid|8811184_29-0" class="reference"><a href="#cite_note-pmid|8811184-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="1986–2000"><span id="1986.E2.80.932000"></span>1986–2000</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=27" title="Edit section: 1986–2000"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="RNA_sequences_can_be_edited_within_cells">RNA sequences can be edited within cells</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=28" title="Edit section: RNA sequences can be edited within cells"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Messenger RNA precursors from a wide range of organisms can be <a href="/wiki/RNA_editing" title="RNA editing">edited</a> before being translated into protein. In this process, non-encoded nucleotides may be inserted into specific sites in the RNA, and encoded nucleotides may be removed or replaced. <a href="/wiki/RNA_editing" title="RNA editing">RNA editing</a> was first discovered within the mitochondria of <a href="/wiki/Kinetoplastida" title="Kinetoplastida">kinetoplastid</a> protozoans, where it has been shown to be extensive.<sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> For example, some protein-coding genes encode fewer than 50% of the nucleotides found within the mature, translated mRNA. Other RNA editing events are found in mammals, plants, bacteria and viruses. These latter editing events involve fewer nucleotide modifications, insertions and deletions than the events within <a href="/wiki/Kinetoplast" title="Kinetoplast">kinetoplast</a> DNA, but still have high biological significance for gene expression and its regulation.<sup id="cite_ref-pmid|11092837_31-0" class="reference"><a href="#cite_note-pmid|11092837-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Telomerase_uses_a_built-in_RNA_template_to_maintain_chromosome_ends">Telomerase uses a built-in RNA template to maintain chromosome ends</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=29" title="Edit section: Telomerase uses a built-in RNA template to maintain chromosome ends"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Telomerase is an enzyme that is present in all eukaryotic nuclei which serves to maintain the ends of the linear DNA in the linear <a href="/wiki/Chromosomes" class="mw-redirect" title="Chromosomes">chromosomes</a> of the eukaryotic nucleus, through the addition of terminal sequences that are lost in each round of DNA replication. Before telomerase was identified, its activity was predicted on the basis of a molecular understanding of DNA replication, which indicated that the DNA polymerases known at that time could not replicate the 3′ end of a linear chromosome, due to the absence of a template strand. Telomerase was shown to be a <a href="/wiki/Ribonucleoprotein" class="mw-redirect" title="Ribonucleoprotein">ribonucleoprotein</a> enzyme that contains an RNA component that serves as a <a href="/wiki/Template_strand" class="mw-redirect" title="Template strand">template strand</a>, and a protein component that has <a href="/wiki/Reverse_transcriptase" title="Reverse transcriptase">reverse transcriptase</a> activity and adds nucleotides to the chromosome ends using the internal RNA template.<sup id="cite_ref-pmid|16756500_32-0" class="reference"><a href="#cite_note-pmid|16756500-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Ribosomal_RNA_catalyzes_peptide_bond_formation">Ribosomal RNA catalyzes peptide bond formation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=30" title="Edit section: Ribosomal RNA catalyzes peptide bond formation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>For years, scientists had worked to identify which protein(s) within the <a href="/wiki/Ribosome" title="Ribosome">ribosome</a> were responsible for <a href="/wiki/Peptidyl_transferase" class="mw-redirect" title="Peptidyl transferase">peptidyl transferase</a> function during <a href="/wiki/Translation_(biology)" title="Translation (biology)">translation</a>, because the covalent linking of amino acids represents one of the most central chemical reactions in all of biology. Careful biochemical studies showed that extensively-deproteinized large ribosomal subunits could still catalyze peptide bond formation, thereby implying that the sought-after activity might lie within ribosomal RNA rather than ribosomal proteins. Structural biologists, using <a href="/wiki/X-ray_crystallography" title="X-ray crystallography">X-ray crystallography</a>, localized the peptidyl transferase center of the ribosome to a highly-<a href="/wiki/Conserved_sequence" title="Conserved sequence">conserved</a> region of the large subunit <a href="/wiki/Ribosomal_RNA" title="Ribosomal RNA">ribosomal RNA</a> (rRNA) that is located at the place within the ribosome where the amino-acid-bearing ends of tRNA bind, and where no proteins are present. These studies led to the conclusion that the <a href="/wiki/Ribosome" title="Ribosome">ribosome</a> is a <a href="/wiki/Ribozyme" title="Ribozyme">ribozyme</a>. The rRNA sequences that make up the ribosomal <a href="/wiki/Active_site" title="Active site">active site</a> represent some of the most highly conserved sequences in the biological world. Together, these observations indicate that peptide bond formation catalyzed by RNA was a feature of the <a href="/wiki/Last_universal_ancestor" class="mw-redirect" title="Last universal ancestor">last common ancestor</a> of all known forms of life.<sup id="cite_ref-pmid|1604315_33-0" class="reference"><a href="#cite_note-pmid|1604315-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Combinatorial_selection_of_RNA_molecules_enables_in_vitro_evolution">Combinatorial selection of RNA molecules enables in vitro evolution</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=31" title="Edit section: Combinatorial selection of RNA molecules enables in vitro evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Experimental methods were invented that allowed investigators to use large, diverse populations of RNA molecules to carry out in vitro molecular experiments that utilized powerful selective replication strategies used by geneticists, and which amount to evolution in the test tube. These experiments have been described using different names, the most common of which are "combinatorial selection", "in vitro selection", and SELEX (for <a href="/wiki/Systematic_Evolution_of_Ligands_by_Exponential_Enrichment" class="mw-redirect" title="Systematic Evolution of Ligands by Exponential Enrichment">Systematic Evolution of Ligands by Exponential Enrichment</a>). These experiments have been used for isolating RNA molecules with a wide range of properties, from binding to particular proteins, to catalyzing particular reactions, to binding low molecular weight organic ligands. They have equal applicability to elucidating interactions and mechanisms that are known properties of naturally occurring RNA molecules to isolating RNA molecules with biochemical properties that are not known in nature. In developing in vitro selection technology for RNA, laboratory systems for synthesizing complex populations of RNA molecules were established, and used in conjunction with the selection of molecules with user-specified biochemical activities, and in vitro schemes for RNA replication. These steps can be viewed as (a) <a href="/wiki/Mutation" title="Mutation">mutation</a>, (b) selection, and (c) <a href="/wiki/Self-replication" title="Self-replication">replication</a>. Together, then, these three processes enable in vitro <a href="/wiki/Molecular_evolution" title="Molecular evolution">molecular evolution</a>.<sup id="cite_ref-pmid|11539574_34-0" class="reference"><a href="#cite_note-pmid|11539574-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="2001_–_present"><span id="2001_.E2.80.93_present"></span>2001 – present</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=32" title="Edit section: 2001 – present"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Many_mobile_DNA_elements_use_an_RNA_intermediate">Many mobile DNA elements use an RNA intermediate</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=33" title="Edit section: Many mobile DNA elements use an RNA intermediate"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Transposon" class="mw-redirect" title="Transposon">Transposable genetic elements</a> (transposons) are found which can replicate via transcription into an <a href="/wiki/Retrotransposon" title="Retrotransposon">RNA intermediate</a> which is subsequently converted to DNA by reverse transcriptase. These sequences, many of which are likely related to retroviruses, constitute much of the DNA of the eukaryotic nucleus, especially so in plants. Genomic sequencing shows that retrotransposons make up 36% of the human genome and over half of the genome of major cereal crops (wheat and maize).<sup id="cite_ref-pmid|18680436_35-0" class="reference"><a href="#cite_note-pmid|18680436-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Riboswitches_bind_cellular_metabolites_and_control_gene_expression">Riboswitches bind cellular metabolites and control gene expression</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=34" title="Edit section: Riboswitches bind cellular metabolites and control gene expression"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Segments of RNA, typically embedded within the 5′-untranslated region of a vast number of bacterial mRNA molecules, have a profound effect on gene expression through a previously-undiscovered mechanism that does not involve the participation of proteins. In many cases, <a href="/wiki/Riboswitch" title="Riboswitch">riboswitches</a> change their folded structure in response to environmental conditions (e.g. ambient temperature or concentrations of specific metabolites), and the structural change controls the translation or stability of the mRNA in which the riboswitch is embedded. In this way, gene expression can be dramatically regulated at the post-transcriptional level.<sup id="cite_ref-pmid|19298181_36-0" class="reference"><a href="#cite_note-pmid|19298181-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Small_RNA_molecules_regulate_gene_expression_by_post-transcriptional_gene_silencing">Small RNA molecules regulate gene expression by post-transcriptional gene silencing</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=35" title="Edit section: Small RNA molecules regulate gene expression by post-transcriptional gene silencing"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Another previously unknown mechanism by which RNA molecules are involved in genetic regulation was discovered in the 1990s. Small RNA molecules termed microRNA (miRNA) and <a href="/wiki/RNA_interference" title="RNA interference">small interfering RNA</a> (siRNA) are abundant in eukaryotic cells and exert post-transcriptional control over mRNA expression. They function by binding to specific sites within the mRNA and inducing cleavage of the mRNA via a specific silencing-associated RNA degradation pathway.<sup id="cite_ref-pmid|19239886_37-0" class="reference"><a href="#cite_note-pmid|19239886-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Noncoding_RNA_controls_epigenetic_phenomena">Noncoding RNA controls epigenetic phenomena</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=36" title="Edit section: Noncoding RNA controls epigenetic phenomena"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In addition to their well-established roles in translation and splicing, members of <a href="/wiki/Noncoding_RNA" class="mw-redirect" title="Noncoding RNA">noncoding RNA</a> (ncRNA) families have recently been found to function in genome defense and chromosome inactivation. For example, <a href="/wiki/Piwi-interacting_RNA" title="Piwi-interacting RNA">piwi-interacting RNAs</a> (piRNAs) prevent genome instability in germ line cells, while <a href="/wiki/Xist" class="mw-redirect" title="Xist">Xist</a> (X-inactive-specific-transcript) is essential for X-chromosome inactivation in mammals.<sup id="cite_ref-pmid|21030644_38-0" class="reference"><a href="#cite_note-pmid|21030644-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Nobel_Laureates_in_RNA_biology">Nobel Laureates in RNA biology</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=37" title="Edit section: Nobel Laureates in RNA biology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/List_of_RNA_biologists" title="List of RNA biologists">List of RNA biologists</a></div> <table class="wikitable sortable" align="center"> <tbody><tr> <th scope="col" style="background:#efefef;">Name </th> <th scope="col" style="background:#efefef;">Dates </th> <th scope="col" style="background:#efefef;">Awards </th></tr> <tr> <td><a href="/wiki/Sidney_Altman" title="Sidney Altman">Altman, Sidney</a> </td> <td>born 1939 </td> <td>1989 Nobel Prize in Chemistry </td></tr> <tr> <td><a href="/wiki/Victor_Ambros" title="Victor Ambros">Ambros, Victor</a> </td> <td>born 1953 </td> <td>2024 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/David_Baltimore" title="David Baltimore">Baltimore, David</a> </td> <td>born 1938 </td> <td>1975 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Fran%C3%A7oise_Barr%C3%A9-Sinoussi" title="Françoise Barré-Sinoussi">Barré-Sinoussi, Françoise</a> </td> <td>born 1947 </td> <td>2008 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Elizabeth_Blackburn" title="Elizabeth Blackburn">Blackburn, Elizabeth</a> </td> <td>born 1948 </td> <td>2009 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Sydney_Brenner" title="Sydney Brenner">Brenner, Sydney</a> </td> <td>born 1927 </td> <td>2002 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Thomas_Cech" title="Thomas Cech">Cech, Thomas</a> </td> <td>born 1947 </td> <td>1989 Nobel Prize in Chemistry </td></tr> <tr> <td><a href="/wiki/Emmanuelle_Charpentier" title="Emmanuelle Charpentier">Charpentier, Emmanuelle</a> </td> <td>born 1968 </td> <td>2020 Nobel Prize in Chemistry </td></tr> <tr> <td><a href="/wiki/Francis_Crick" title="Francis Crick">Crick, Francis</a> </td> <td>1916–2004 </td> <td>1962 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Jennifer_Doudna" title="Jennifer Doudna">Doudna, Jennifer</a> </td> <td>born 1964 </td> <td>2020 Nobel Prize in Chemistry </td></tr> <tr> <td><a href="/wiki/Renato_Dulbecco" title="Renato Dulbecco">Dulbecco, Renato</a> </td> <td>1914–2012 </td> <td>1975 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Andrew_Fire" title="Andrew Fire">Fire, Andrew</a> </td> <td>born 1959 </td> <td>2006 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Walter_Gilbert" title="Walter Gilbert">Gilbert, Walter</a> </td> <td>born 1932 </td> <td>1980 Nobel Prize in Chemistry </td></tr> <tr> <td><a href="/wiki/Carol_W._Greider" title="Carol W. Greider">Greider, Carol</a> </td> <td>born 1961 </td> <td>2009 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Robert_W._Holley" title="Robert W. Holley">Holley, Robert</a> </td> <td>1922–1993 </td> <td>1968 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Fran%C3%A7ois_Jacob" title="François Jacob">Jacob, François</a> </td> <td>1920–2013 </td> <td>1965 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Katalin_Karik%C3%B3" title="Katalin Karikó">Karikó, Katalin</a> </td> <td>born 1955 </td> <td>2023 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Har_Gobind_Khorana" title="Har Gobind Khorana">Khorana, H. Gobind</a> </td> <td>1922–2011 </td> <td>1968 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Aaron_Klug" title="Aaron Klug">Klug, Aaron</a> </td> <td>born 1926 </td> <td>1982 Nobel Prize in Chemistry </td></tr> <tr> <td><a href="/wiki/Roger_D._Kornberg" title="Roger D. Kornberg">Kornberg, Roger</a> </td> <td>born 1947 </td> <td>2006 Nobel Prize in Chemistry </td></tr> <tr> <td><a href="/wiki/Craig_Mello" title="Craig Mello">Mello, Craig</a> </td> <td>born 1960 </td> <td>2006 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Jacques_Monod" title="Jacques Monod">Monod, Jacques</a> </td> <td>1910–1976 </td> <td>1965 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Luc_Montagnier" title="Luc Montagnier">Montagnier, Luc</a> </td> <td>born 1932 </td> <td>2008 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Marshall_Warren_Nirenberg" title="Marshall Warren Nirenberg">Nirenberg, Marshall</a> </td> <td>1927–2010 </td> <td>1968 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Severo_Ochoa" title="Severo Ochoa">Ochoa, Severo</a> </td> <td>1905–1993 </td> <td>1959 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Howard_Temin" class="mw-redirect" title="Howard Temin">Temin, Howard</a> </td> <td>1934–1994 </td> <td>1975 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Venkatraman_Ramakrishnan" class="mw-redirect" title="Venkatraman Ramakrishnan">Ramakrishnan, Venkatraman</a> </td> <td>born 1952 </td> <td>2009 Nobel Prize in Chemistry </td></tr> <tr> <td><a href="/wiki/Richard_J._Roberts" title="Richard J. Roberts">Roberts, Richard</a> </td> <td>born 1943 </td> <td>1993 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Gary_Ruvkun" title="Gary Ruvkun">Ruvkun, Gary</a> </td> <td>born 1952 </td> <td>2024 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Philip_Allen_Sharp" class="mw-redirect" title="Philip Allen Sharp">Sharp, Philip</a> </td> <td>born 1944 </td> <td>1993 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Thomas_A._Steitz" title="Thomas A. Steitz">Steitz, Thomas</a> </td> <td>1940–2018 </td> <td>2009 Nobel Prize in Chemistry </td></tr> <tr> <td><a href="/wiki/Jack_W._Szostak" title="Jack W. Szostak">Szostak, Jack</a> </td> <td>born 1952 </td> <td>2009 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Alexander_R._Todd,_Baron_Todd" class="mw-redirect" title="Alexander R. Todd, Baron Todd">Todd, Alexander</a> </td> <td>1907–1997 </td> <td>1957 Nobel Prize in Chemistry </td></tr> <tr> <td><a href="/wiki/James_D._Watson" class="mw-redirect" title="James D. Watson">Watson, James</a> </td> <td>born 1928 </td> <td>1962 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/w/index.php?title=Drew_Weisman&action=edit&redlink=1" class="new" title="Drew Weisman (page does not exist)">Weissman, Drew</a> </td> <td>born 1959 </td> <td>2023 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Maurice_Wilkins" title="Maurice Wilkins">Wilkins, Maurice</a> </td> <td>1916–2004 </td> <td>1962 Nobel Prize in Physiology or Medicine </td></tr> <tr> <td><a href="/wiki/Ada_Yonath" title="Ada Yonath">Yonath, Ada</a> </td> <td>born 1939 </td> <td>2009 Nobel Prize in Chemistry </td></tr></tbody></table> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=History_of_RNA_biology&action=edit&section=38" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width reflist-columns-2"> <ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></span> <span class="reference-text">Oxford Dictionary of Biochemistry and Molecular Biology. <a rel="nofollow" class="external text" href="http://www.oxfordreference.com/view/10.1093/oi/authority.20110803104532550">"Thymus Nucleic Acid"</a>, <i><a href="/wiki/Oxford_Reference" class="mw-redirect" title="Oxford Reference">Oxford Reference</a></i>. Retrieved on 21 October 2015.</span> </li> <li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFAllen1941" class="citation journal cs1">Allen, F W (June 1941). "The Biochemistry of the Nucleic Acids, Purines, and Pyrimidines". <i><a href="/wiki/Annual_Review_of_Biochemistry" title="Annual Review of Biochemistry">Annual Review of Biochemistry</a></i>. <b>10</b> (1): <span class="nowrap">221–</span>244. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1146%2Fannurev.bi.10.070141.001253">10.1146/annurev.bi.10.070141.001253</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annual+Review+of+Biochemistry&rft.atitle=The+Biochemistry+of+the+Nucleic+Acids%2C+Purines%2C+and+Pyrimidines&rft.volume=10&rft.issue=1&rft.pages=%3Cspan+class%3D%22nowrap%22%3E221-%3C%2Fspan%3E244&rft.date=1941-06&rft_id=info%3Adoi%2F10.1146%2Fannurev.bi.10.070141.001253&rft.aulast=Allen&rft.aufirst=F+W&rfr_id=info%3Asid%2Fen.wikipedia.org%3AHistory+of+RNA+biology" class="Z3988"></span></span> </li> <li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBrachet1933" class="citation journal cs1 cs1-prop-foreign-lang-source">Brachet, J. (1933). "Recherches sur la synthese de l'acide thymonucleique pendant le developpement de l'oeuf d'Oursin" [Research on the synthesis of thymonucleic acid during the development of the sea urchin egg]. <i>Archives de Biologie</i> (in French). <b>44</b>: <span class="nowrap">519–</span>576.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Archives+de+Biologie&rft.atitle=Recherches+sur+la+synthese+de+l%27acide+thymonucleique+pendant+le+developpement+de+l%27oeuf+d%27Oursin&rft.volume=44&rft.pages=%3Cspan+class%3D%22nowrap%22%3E519-%3C%2Fspan%3E576&rft.date=1933&rft.aulast=Brachet&rft.aufirst=J.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AHistory+of+RNA+biology" class="Z3988"></span></span> </li> <li id="cite_note-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-4">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBurian1994" class="citation book cs1">Burian, R. (1994). <a rel="nofollow" class="external text" href="http://www.histcnrs.fr/ColloqDijon/Burian-Brachet.pdf">"Jean Brachet's Cytochemical Embryology: Connections with the Renovation of Biology in France?"</a> <span class="cs1-format">(PDF)</span>. In Debru, C.; Gayon, J.; Picard, J.-F. (eds.). <i>Les sciences biologiques et médicales en France 1920–1950</i>. Cahiers pour I'histoire de la recherche. Vol. 2. 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