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DNA - Wikipedia
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id="toc-Nucleobase_classification" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Nucleobase_classification"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1</span> <span>Nucleobase classification</span> </div> </a> <ul id="toc-Nucleobase_classification-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Non-canonical_bases" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Non-canonical_bases"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>Non-canonical bases</span> </div> </a> <ul id="toc-Non-canonical_bases-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Grooves" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Grooves"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.3</span> <span>Grooves</span> </div> </a> <ul id="toc-Grooves-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Base_pairing" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Base_pairing"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4</span> <span>Base pairing</span> </div> </a> <ul id="toc-Base_pairing-sublist" class="vector-toc-list"> <li id="toc-ssDNA_vs._dsDNA" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#ssDNA_vs._dsDNA"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4.1</span> <span>ssDNA vs. dsDNA</span> </div> </a> <ul id="toc-ssDNA_vs._dsDNA-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Amount" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Amount"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.5</span> <span>Amount</span> </div> </a> <ul id="toc-Amount-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Sense_and_antisense" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Sense_and_antisense"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.6</span> <span>Sense and antisense</span> </div> </a> <ul id="toc-Sense_and_antisense-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Supercoiling" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Supercoiling"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.7</span> <span>Supercoiling</span> </div> </a> <ul id="toc-Supercoiling-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Alternative_DNA_structures" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Alternative_DNA_structures"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.8</span> <span>Alternative DNA structures</span> </div> </a> <ul id="toc-Alternative_DNA_structures-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Alternative_DNA_chemistry" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Alternative_DNA_chemistry"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.9</span> <span>Alternative DNA chemistry</span> </div> </a> <ul id="toc-Alternative_DNA_chemistry-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Quadruplex_structures" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Quadruplex_structures"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.10</span> <span>Quadruplex structures</span> </div> </a> <ul id="toc-Quadruplex_structures-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Branched_DNA" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Branched_DNA"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.11</span> <span>Branched DNA</span> </div> </a> <ul id="toc-Branched_DNA-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Artificial_bases" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Artificial_bases"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.12</span> <span>Artificial bases</span> </div> </a> <ul id="toc-Artificial_bases-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Acidity" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Acidity"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.13</span> <span>Acidity</span> </div> </a> <ul id="toc-Acidity-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Macroscopic_appearance" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Macroscopic_appearance"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.14</span> <span>Macroscopic appearance</span> </div> </a> <ul id="toc-Macroscopic_appearance-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Chemical_modifications_and_altered_DNA_packaging" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Chemical_modifications_and_altered_DNA_packaging"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Chemical modifications and altered DNA packaging</span> </div> </a> <button aria-controls="toc-Chemical_modifications_and_altered_DNA_packaging-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 Chemical modifications and altered DNA packaging subsection</span> </button> <ul id="toc-Chemical_modifications_and_altered_DNA_packaging-sublist" class="vector-toc-list"> <li id="toc-Base_modifications_and_DNA_packaging" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Base_modifications_and_DNA_packaging"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Base modifications and DNA packaging</span> </div> </a> <ul id="toc-Base_modifications_and_DNA_packaging-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Damage" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Damage"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Damage</span> </div> </a> <ul id="toc-Damage-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Biological_functions" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Biological_functions"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Biological functions</span> </div> </a> <button aria-controls="toc-Biological_functions-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 Biological functions subsection</span> </button> <ul id="toc-Biological_functions-sublist" class="vector-toc-list"> <li id="toc-Genes_and_genomes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genes_and_genomes"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Genes and genomes</span> </div> </a> <ul id="toc-Genes_and_genomes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Transcription_and_translation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Transcription_and_translation"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Transcription and translation</span> </div> </a> <ul id="toc-Transcription_and_translation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Replication" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Replication"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.3</span> <span>Replication</span> </div> </a> <ul id="toc-Replication-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Extracellular_nucleic_acids" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Extracellular_nucleic_acids"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.4</span> <span>Extracellular nucleic acids</span> </div> </a> <ul id="toc-Extracellular_nucleic_acids-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Neutrophil_extracellular_traps" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Neutrophil_extracellular_traps"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.5</span> <span>Neutrophil extracellular traps</span> </div> </a> <ul id="toc-Neutrophil_extracellular_traps-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Interactions_with_proteins" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Interactions_with_proteins"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Interactions with proteins</span> </div> </a> <button aria-controls="toc-Interactions_with_proteins-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 Interactions with proteins subsection</span> </button> <ul id="toc-Interactions_with_proteins-sublist" class="vector-toc-list"> <li id="toc-DNA-binding_proteins" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#DNA-binding_proteins"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>DNA-binding proteins</span> </div> </a> <ul id="toc-DNA-binding_proteins-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-DNA-modifying_enzymes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#DNA-modifying_enzymes"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2</span> <span>DNA-modifying enzymes</span> </div> </a> <ul id="toc-DNA-modifying_enzymes-sublist" class="vector-toc-list"> <li id="toc-Nucleases_and_ligases" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Nucleases_and_ligases"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2.1</span> <span>Nucleases and ligases</span> </div> </a> <ul id="toc-Nucleases_and_ligases-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Topoisomerases_and_helicases" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Topoisomerases_and_helicases"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2.2</span> <span>Topoisomerases and helicases</span> </div> </a> <ul id="toc-Topoisomerases_and_helicases-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Polymerases" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Polymerases"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2.3</span> <span>Polymerases</span> </div> </a> <ul id="toc-Polymerases-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-Genetic_recombination" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Genetic_recombination"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Genetic recombination</span> </div> </a> <ul id="toc-Genetic_recombination-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Evolution" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Evolution"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Evolution</span> </div> </a> <ul id="toc-Evolution-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Uses_in_technology" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Uses_in_technology"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Uses in technology</span> </div> </a> <button aria-controls="toc-Uses_in_technology-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 Uses in technology subsection</span> </button> <ul id="toc-Uses_in_technology-sublist" class="vector-toc-list"> <li id="toc-Genetic_engineering" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genetic_engineering"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1</span> <span>Genetic engineering</span> </div> </a> <ul id="toc-Genetic_engineering-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-DNA_profiling" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#DNA_profiling"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.2</span> <span>DNA profiling</span> </div> </a> <ul id="toc-DNA_profiling-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-DNA_enzymes_or_catalytic_DNA" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#DNA_enzymes_or_catalytic_DNA"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.3</span> <span>DNA enzymes or catalytic DNA</span> </div> </a> <ul id="toc-DNA_enzymes_or_catalytic_DNA-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Bioinformatics" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Bioinformatics"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.4</span> <span>Bioinformatics</span> </div> </a> <ul id="toc-Bioinformatics-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-DNA_nanotechnology" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#DNA_nanotechnology"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.5</span> <span>DNA nanotechnology</span> </div> </a> <ul id="toc-DNA_nanotechnology-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-History_and_anthropology" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#History_and_anthropology"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.6</span> <span>History and anthropology</span> </div> </a> <ul id="toc-History_and_anthropology-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Information_storage" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Information_storage"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.7</span> <span>Information storage</span> </div> </a> <ul id="toc-Information_storage-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-History" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#History"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>History</span> </div> </a> <ul id="toc-History-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>See also</span> </div> </a> <ul id="toc-See_also-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">10</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">11</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">12</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " 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Available in 164 languages" > <label id="p-lang-btn-label" for="p-lang-btn-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--action-progressive mw-portlet-lang-heading-164" aria-hidden="true" ><span class="vector-icon mw-ui-icon-language-progressive mw-ui-icon-wikimedia-language-progressive"></span> <span class="vector-dropdown-label-text">164 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-af mw-list-item"><a href="https://af.wikipedia.org/wiki/DNS" title="DNS – Afrikaans" lang="af" hreflang="af" data-title="DNS" data-language-autonym="Afrikaans" data-language-local-name="Afrikaans" class="interlanguage-link-target"><span>Afrikaans</span></a></li><li class="interlanguage-link interwiki-als mw-list-item"><a href="https://als.wikipedia.org/wiki/Desoxyribonukleins%C3%A4ure" title="Desoxyribonukleinsäure – Alemannic" lang="gsw" hreflang="gsw" data-title="Desoxyribonukleinsäure" data-language-autonym="Alemannisch" data-language-local-name="Alemannic" class="interlanguage-link-target"><span>Alemannisch</span></a></li><li class="interlanguage-link interwiki-am mw-list-item"><a href="https://am.wikipedia.org/wiki/%E1%8B%B2_%E1%8A%A4%E1%8A%95_%E1%8A%A4" title="ዲ ኤን ኤ – Amharic" lang="am" hreflang="am" data-title="ዲ ኤን ኤ" data-language-autonym="አማርኛ" data-language-local-name="Amharic" class="interlanguage-link-target"><span>አማርኛ</span></a></li><li class="interlanguage-link interwiki-ar badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://ar.wikipedia.org/wiki/%D8%AD%D9%85%D8%B6_%D9%86%D9%88%D9%88%D9%8A_%D8%B1%D9%8A%D8%A8%D9%88%D8%B2%D9%8A_%D9%85%D9%86%D9%82%D9%88%D8%B5_%D8%A7%D9%84%D8%A3%D9%83%D8%B3%D8%AC%D9%8A%D9%86" title="حمض نووي ريبوزي منقوص الأكسجين – Arabic" lang="ar" hreflang="ar" data-title="حمض نووي ريبوزي منقوص الأكسجين" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-an mw-list-item"><a href="https://an.wikipedia.org/wiki/Acido_desoxirribonucleico" title="Acido desoxirribonucleico – Aragonese" lang="an" hreflang="an" data-title="Acido desoxirribonucleico" data-language-autonym="Aragonés" data-language-local-name="Aragonese" class="interlanguage-link-target"><span>Aragonés</span></a></li><li class="interlanguage-link interwiki-as mw-list-item"><a href="https://as.wikipedia.org/wiki/%E0%A6%A1%E0%A6%BF_%E0%A6%8F%E0%A6%A8_%E0%A6%8F" title="ডি এন এ – Assamese" lang="as" hreflang="as" data-title="ডি এন এ" data-language-autonym="অসমীয়া" data-language-local-name="Assamese" class="interlanguage-link-target"><span>অসমীয়া</span></a></li><li class="interlanguage-link interwiki-ast mw-list-item"><a href="https://ast.wikipedia.org/wiki/%C3%81cidu_desoxirribonucleico" title="Ácidu desoxirribonucleico – Asturian" lang="ast" hreflang="ast" data-title="Ácidu desoxirribonucleico" data-language-autonym="Asturianu" data-language-local-name="Asturian" class="interlanguage-link-target"><span>Asturianu</span></a></li><li class="interlanguage-link interwiki-gn mw-list-item"><a href="https://gn.wikipedia.org/wiki/ADN" title="ADN – Guarani" lang="gn" hreflang="gn" data-title="ADN" data-language-autonym="Avañe'ẽ" data-language-local-name="Guarani" class="interlanguage-link-target"><span>Avañe'ẽ</span></a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/DNT" title="DNT – Azerbaijani" lang="az" hreflang="az" data-title="DNT" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target"><span>Azərbaycanca</span></a></li><li class="interlanguage-link interwiki-azb mw-list-item"><a href="https://azb.wikipedia.org/wiki/%D8%AF%DB%8C%E2%80%8C%D8%A7%D9%86%E2%80%8C%D8%A7%D8%A6%DB%8C" title="دیانائی – South Azerbaijani" lang="azb" hreflang="azb" data-title="دیانائی" data-language-autonym="تۆرکجه" data-language-local-name="South Azerbaijani" class="interlanguage-link-target"><span>تۆرکجه</span></a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%A1%E0%A6%BF%E0%A6%8F%E0%A6%A8%E0%A6%8F" title="ডিএনএ – Bangla" lang="bn" hreflang="bn" data-title="ডিএনএ" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-bjn mw-list-item"><a href="https://bjn.wikipedia.org/wiki/Asam_deoksiribonukleat" title="Asam deoksiribonukleat – Banjar" lang="bjn" hreflang="bjn" data-title="Asam deoksiribonukleat" data-language-autonym="Banjar" data-language-local-name="Banjar" class="interlanguage-link-target"><span>Banjar</span></a></li><li class="interlanguage-link interwiki-zh-min-nan mw-list-item"><a href="https://zh-min-nan.wikipedia.org/wiki/DNA" title="DNA – Minnan" lang="nan" hreflang="nan" data-title="DNA" data-language-autonym="閩南語 / Bân-lâm-gú" data-language-local-name="Minnan" class="interlanguage-link-target"><span>閩南語 / Bân-lâm-gú</span></a></li><li class="interlanguage-link interwiki-ba mw-list-item"><a href="https://ba.wikipedia.org/wiki/%D0%94%D0%9D%D0%9A" title="ДНК – Bashkir" lang="ba" hreflang="ba" data-title="ДНК" data-language-autonym="Башҡортса" data-language-local-name="Bashkir" class="interlanguage-link-target"><span>Башҡортса</span></a></li><li class="interlanguage-link interwiki-be mw-list-item"><a href="https://be.wikipedia.org/wiki/%D0%94%D1%8D%D0%B7%D0%BE%D0%BA%D1%81%D1%96%D1%80%D1%8B%D0%B1%D0%B0%D0%BD%D1%83%D0%BA%D0%BB%D0%B5%D1%96%D0%BD%D0%B0%D0%B2%D0%B0%D1%8F_%D0%BA%D1%96%D1%81%D0%BB%D0%B0%D1%82%D0%B0" title="Дэзоксірыбануклеінавая кіслата – Belarusian" lang="be" hreflang="be" data-title="Дэзоксірыбануклеінавая кіслата" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target"><span>Беларуская</span></a></li><li class="interlanguage-link interwiki-be-x-old mw-list-item"><a href="https://be-tarask.wikipedia.org/wiki/%D0%94%D1%8D%D0%B7%D0%BE%D0%BA%D1%81%D1%8B%D1%80%D1%8B%D0%B1%D0%B0%D0%BD%D1%83%D0%BA%D0%BB%D1%96%D0%B9%D0%BD%D0%B0%D1%8F_%D0%BA%D1%96%D1%81%D1%8C%D0%BB%D1%8F" title="Дэзоксырыбануклійная кісьля – Belarusian (Taraškievica orthography)" lang="be-tarask" hreflang="be-tarask" data-title="Дэзоксырыбануклійная кісьля" data-language-autonym="Беларуская (тарашкевіца)" data-language-local-name="Belarusian (Taraškievica orthography)" class="interlanguage-link-target"><span>Беларуская (тарашкевіца)</span></a></li><li class="interlanguage-link interwiki-bh mw-list-item"><a href="https://bh.wikipedia.org/wiki/%E0%A4%A1%E0%A5%80%E0%A4%8F%E0%A4%A8%E0%A4%8F" title="डीएनए – Bhojpuri" lang="bh" hreflang="bh" data-title="डीएनए" data-language-autonym="भोजपुरी" data-language-local-name="Bhojpuri" class="interlanguage-link-target"><span>भोजपुरी</span></a></li><li class="interlanguage-link interwiki-bcl mw-list-item"><a href="https://bcl.wikipedia.org/wiki/DNA" title="DNA – Central Bikol" lang="bcl" hreflang="bcl" data-title="DNA" data-language-autonym="Bikol Central" data-language-local-name="Central Bikol" class="interlanguage-link-target"><span>Bikol Central</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%94%D0%B5%D0%B7%D0%BE%D0%BA%D1%81%D0%B8%D1%80%D0%B8%D0%B1%D0%BE%D0%BD%D1%83%D0%BA%D0%BB%D0%B5%D0%B8%D0%BD%D0%BE%D0%B2%D0%B0_%D0%BA%D0%B8%D1%81%D0%B5%D0%BB%D0%B8%D0%BD%D0%B0" title="Дезоксирибонуклеинова киселина – Bulgarian" lang="bg" hreflang="bg" data-title="Дезоксирибонуклеинова киселина" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Dezoksiribonukleinska_kiselina" title="Dezoksiribonukleinska kiselina – Bosnian" lang="bs" hreflang="bs" data-title="Dezoksiribonukleinska kiselina" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-br mw-list-item"><a href="https://br.wikipedia.org/wiki/Trenkenn_dezoksiribonukleek" title="Trenkenn dezoksiribonukleek – Breton" lang="br" hreflang="br" data-title="Trenkenn dezoksiribonukleek" data-language-autonym="Brezhoneg" data-language-local-name="Breton" class="interlanguage-link-target"><span>Brezhoneg</span></a></li><li class="interlanguage-link interwiki-bxr mw-list-item"><a href="https://bxr.wikipedia.org/wiki/%D0%94%D0%B5%D0%B7%D0%BE%D0%BA%D1%81%D0%B8%D1%80%D0%B8%D0%B1%D0%BE%D0%BD%D1%83%D0%BA%D0%BB%D0%B5%D0%B8%D0%BD%D0%B0%D0%B9_%D1%85%D2%AF%D1%88%D1%8D%D0%BB" title="Дезоксирибонуклеинай хүшэл – Russia Buriat" lang="bxr" hreflang="bxr" data-title="Дезоксирибонуклеинай хүшэл" data-language-autonym="Буряад" data-language-local-name="Russia Buriat" class="interlanguage-link-target"><span>Буряад</span></a></li><li class="interlanguage-link interwiki-ca badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://ca.wikipedia.org/wiki/%C3%80cid_desoxiribonucleic" title="Àcid desoxiribonucleic – Catalan" lang="ca" hreflang="ca" data-title="Àcid desoxiribonucleic" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cv mw-list-item"><a href="https://cv.wikipedia.org/wiki/%D0%94%D0%B5%D0%B7%D0%BE%D0%BA%D1%81%D0%B8%D1%80%D0%B8%D0%B1%D0%BE%D0%BD%D1%83%D0%BA%D0%BB%D0%B5%D0%B8%D0%BD_%D0%B9%D3%B3%D2%AB%D0%B5%D0%BA%D3%97" title="Дезоксирибонуклеин йӳҫекӗ – Chuvash" lang="cv" hreflang="cv" data-title="Дезоксирибонуклеин йӳҫекӗ" data-language-autonym="Чӑвашла" data-language-local-name="Chuvash" class="interlanguage-link-target"><span>Чӑвашла</span></a></li><li class="interlanguage-link interwiki-ceb mw-list-item"><a href="https://ceb.wikipedia.org/wiki/DNA" title="DNA – Cebuano" lang="ceb" hreflang="ceb" data-title="DNA" data-language-autonym="Cebuano" data-language-local-name="Cebuano" class="interlanguage-link-target"><span>Cebuano</span></a></li><li class="interlanguage-link interwiki-cs badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://cs.wikipedia.org/wiki/DNA" title="DNA – Czech" lang="cs" hreflang="cs" data-title="DNA" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-cy mw-list-item"><a href="https://cy.wikipedia.org/wiki/DNA" title="DNA – Welsh" lang="cy" hreflang="cy" data-title="DNA" data-language-autonym="Cymraeg" data-language-local-name="Welsh" class="interlanguage-link-target"><span>Cymraeg</span></a></li><li class="interlanguage-link interwiki-da badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://da.wikipedia.org/wiki/DNA" title="DNA – Danish" lang="da" hreflang="da" data-title="DNA" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-pdc mw-list-item"><a href="https://pdc.wikipedia.org/wiki/DNA" title="DNA – Pennsylvania German" lang="pdc" hreflang="pdc" data-title="DNA" data-language-autonym="Deitsch" data-language-local-name="Pennsylvania German" class="interlanguage-link-target"><span>Deitsch</span></a></li><li class="interlanguage-link interwiki-de badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://de.wikipedia.org/wiki/Desoxyribonukleins%C3%A4ure" title="Desoxyribonukleinsäure – German" lang="de" hreflang="de" data-title="Desoxyribonukleinsäure" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/DNA" title="DNA – Estonian" lang="et" hreflang="et" data-title="DNA" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/DNA" title="DNA – Greek" lang="el" hreflang="el" data-title="DNA" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-es badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://es.wikipedia.org/wiki/%C3%81cido_desoxirribonucleico" title="Ácido desoxirribonucleico – Spanish" lang="es" hreflang="es" data-title="Ácido desoxirribonucleico" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/DNA" title="DNA – Esperanto" lang="eo" hreflang="eo" data-title="DNA" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-ext mw-list-item"><a href="https://ext.wikipedia.org/wiki/ADN" title="ADN – Extremaduran" lang="ext" hreflang="ext" data-title="ADN" data-language-autonym="Estremeñu" data-language-local-name="Extremaduran" class="interlanguage-link-target"><span>Estremeñu</span></a></li><li class="interlanguage-link interwiki-eu badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://eu.wikipedia.org/wiki/Azido_desoxirribonukleiko" title="Azido desoxirribonukleiko – Basque" lang="eu" hreflang="eu" data-title="Azido desoxirribonukleiko" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%AF%DB%8C%E2%80%8C%D8%A7%D9%86%E2%80%8C%D8%A7%DB%8C" title="دیانای – Persian" lang="fa" hreflang="fa" data-title="دیانای" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-hif mw-list-item"><a href="https://hif.wikipedia.org/wiki/DNA" title="DNA – Fiji Hindi" lang="hif" hreflang="hif" data-title="DNA" data-language-autonym="Fiji Hindi" data-language-local-name="Fiji Hindi" class="interlanguage-link-target"><span>Fiji Hindi</span></a></li><li class="interlanguage-link interwiki-fo mw-list-item"><a href="https://fo.wikipedia.org/wiki/DNA" title="DNA – Faroese" lang="fo" hreflang="fo" data-title="DNA" data-language-autonym="Føroyskt" data-language-local-name="Faroese" class="interlanguage-link-target"><span>Føroyskt</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Acide_d%C3%A9soxyribonucl%C3%A9ique" title="Acide désoxyribonucléique – French" lang="fr" hreflang="fr" data-title="Acide désoxyribonucléique" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-fy mw-list-item"><a href="https://fy.wikipedia.org/wiki/DNA" title="DNA – Western Frisian" lang="fy" hreflang="fy" data-title="DNA" data-language-autonym="Frysk" data-language-local-name="Western Frisian" class="interlanguage-link-target"><span>Frysk</span></a></li><li class="interlanguage-link interwiki-fur mw-list-item"><a href="https://fur.wikipedia.org/wiki/DNA" title="DNA – Friulian" lang="fur" hreflang="fur" data-title="DNA" data-language-autonym="Furlan" data-language-local-name="Friulian" class="interlanguage-link-target"><span>Furlan</span></a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/Aig%C3%A9ad_d%C3%AD-ocsairibean%C3%BAicl%C3%A9asach" title="Aigéad dí-ocsairibeanúicléasach – Irish" lang="ga" hreflang="ga" data-title="Aigéad dí-ocsairibeanúicléasach" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target"><span>Gaeilge</span></a></li><li class="interlanguage-link interwiki-gd mw-list-item"><a href="https://gd.wikipedia.org/wiki/DNA" title="DNA – Scottish Gaelic" lang="gd" hreflang="gd" data-title="DNA" data-language-autonym="Gàidhlig" data-language-local-name="Scottish Gaelic" class="interlanguage-link-target"><span>Gàidhlig</span></a></li><li class="interlanguage-link interwiki-gl badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://gl.wikipedia.org/wiki/%C3%81cido_desoxirribonucleico" title="Ácido desoxirribonucleico – Galician" lang="gl" hreflang="gl" data-title="Ácido desoxirribonucleico" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-inh mw-list-item"><a href="https://inh.wikipedia.org/wiki/%D0%94%D0%B5%D0%B7%D0%BE%D0%BA%D1%81%D0%B8%D1%80%D0%B8%D0%B1%D0%BE%D0%BD%D1%83%D0%BA%D0%BB%D0%B5%D0%B8%D0%BD%D0%B0_%D0%BA%D0%B8%D1%81%D0%BB%D0%BE%D1%82%D0%B0" title="Дезоксирибонуклеина кислота – Ingush" lang="inh" hreflang="inh" data-title="Дезоксирибонуклеина кислота" data-language-autonym="ГӀалгӀай" data-language-local-name="Ingush" class="interlanguage-link-target"><span>ГӀалгӀай</span></a></li><li class="interlanguage-link interwiki-hak mw-list-item"><a href="https://hak.wikipedia.org/wiki/DNA" title="DNA – Hakka Chinese" lang="hak" hreflang="hak" data-title="DNA" data-language-autonym="客家語 / Hak-kâ-ngî" data-language-local-name="Hakka Chinese" class="interlanguage-link-target"><span>客家語 / Hak-kâ-ngî</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/DNA" title="DNA – Korean" lang="ko" hreflang="ko" data-title="DNA" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hy badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://hy.wikipedia.org/wiki/%D4%B4%D5%86%D4%B9" title="ԴՆԹ – Armenian" lang="hy" hreflang="hy" data-title="ԴՆԹ" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%A1%E0%A5%80%E0%A4%91%E0%A4%95%E0%A5%8D%E0%A4%B8%E0%A5%80%E0%A4%B0%E0%A4%BE%E0%A4%87%E0%A4%AC%E0%A5%8B_%E0%A4%A8%E0%A5%8D%E0%A4%AF%E0%A5%82%E0%A4%95%E0%A5%8D%E0%A4%B2%E0%A4%BF%E0%A4%95_%E0%A4%85%E0%A4%AE%E0%A5%8D%E0%A4%B2" title="डीऑक्सीराइबो न्यूक्लिक अम्ल – Hindi" lang="hi" hreflang="hi" data-title="डीऑक्सीराइबो न्यूक्लिक अम्ल" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target"><span>हिन्दी</span></a></li><li class="interlanguage-link interwiki-hr badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://hr.wikipedia.org/wiki/Deoksiribonukleinska_kiselina" title="Deoksiribonukleinska kiselina – Croatian" lang="hr" hreflang="hr" data-title="Deoksiribonukleinska kiselina" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-gor mw-list-item"><a href="https://gor.wikipedia.org/wiki/DNA" title="DNA – Gorontalo" lang="gor" hreflang="gor" data-title="DNA" data-language-autonym="Bahasa Hulontalo" data-language-local-name="Gorontalo" class="interlanguage-link-target"><span>Bahasa Hulontalo</span></a></li><li class="interlanguage-link interwiki-io mw-list-item"><a href="https://io.wikipedia.org/wiki/DNA" title="DNA – Ido" lang="io" hreflang="io" data-title="DNA" data-language-autonym="Ido" data-language-local-name="Ido" class="interlanguage-link-target"><span>Ido</span></a></li><li class="interlanguage-link interwiki-ilo mw-list-item"><a href="https://ilo.wikipedia.org/wiki/Deoksiribonukleiko_nga_asido" title="Deoksiribonukleiko nga asido – Iloko" lang="ilo" hreflang="ilo" data-title="Deoksiribonukleiko nga asido" data-language-autonym="Ilokano" data-language-local-name="Iloko" class="interlanguage-link-target"><span>Ilokano</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Asam_deoksiribonukleat" title="Asam deoksiribonukleat – Indonesian" lang="id" hreflang="id" data-title="Asam deoksiribonukleat" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-ia mw-list-item"><a href="https://ia.wikipedia.org/wiki/Acido_deoxyribonucleic" title="Acido deoxyribonucleic – Interlingua" lang="ia" hreflang="ia" data-title="Acido deoxyribonucleic" data-language-autonym="Interlingua" data-language-local-name="Interlingua" class="interlanguage-link-target"><span>Interlingua</span></a></li><li class="interlanguage-link interwiki-zu mw-list-item"><a href="https://zu.wikipedia.org/wiki/ULibofuzo" title="ULibofuzo – Zulu" lang="zu" hreflang="zu" data-title="ULibofuzo" data-language-autonym="IsiZulu" data-language-local-name="Zulu" class="interlanguage-link-target"><span>IsiZulu</span></a></li><li class="interlanguage-link interwiki-is mw-list-item"><a href="https://is.wikipedia.org/wiki/DNA" title="DNA – Icelandic" lang="is" hreflang="is" data-title="DNA" data-language-autonym="Íslenska" data-language-local-name="Icelandic" class="interlanguage-link-target"><span>Íslenska</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/DNA" title="DNA – Italian" lang="it" hreflang="it" data-title="DNA" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/DNA" title="DNA – Hebrew" lang="he" hreflang="he" data-title="DNA" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-jv mw-list-item"><a href="https://jv.wikipedia.org/wiki/DNA" title="DNA – Javanese" lang="jv" hreflang="jv" data-title="DNA" data-language-autonym="Jawa" data-language-local-name="Javanese" class="interlanguage-link-target"><span>Jawa</span></a></li><li class="interlanguage-link interwiki-kn mw-list-item"><a href="https://kn.wikipedia.org/wiki/%E0%B2%A1%E0%B2%BF%E0%B2%8E%E0%B2%A8%E0%B3%8D%E0%B2%8E_-(DNA)" title="ಡಿಎನ್ಎ -(DNA) – Kannada" lang="kn" hreflang="kn" data-title="ಡಿಎನ್ಎ -(DNA)" data-language-autonym="ಕನ್ನಡ" data-language-local-name="Kannada" class="interlanguage-link-target"><span>ಕನ್ನಡ</span></a></li><li class="interlanguage-link interwiki-pam mw-list-item"><a href="https://pam.wikipedia.org/wiki/DNA" title="DNA – Pampanga" lang="pam" hreflang="pam" data-title="DNA" data-language-autonym="Kapampangan" data-language-local-name="Pampanga" class="interlanguage-link-target"><span>Kapampangan</span></a></li><li class="interlanguage-link interwiki-ka badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://ka.wikipedia.org/wiki/%E1%83%93%E1%83%94%E1%83%96%E1%83%9D%E1%83%A5%E1%83%A1%E1%83%98%E1%83%A0%E1%83%98%E1%83%91%E1%83%9D%E1%83%9C%E1%83%A3%E1%83%99%E1%83%9A%E1%83%94%E1%83%98%E1%83%9C%E1%83%98%E1%83%A1_%E1%83%9B%E1%83%9F%E1%83%90%E1%83%95%E1%83%90" title="დეზოქსირიბონუკლეინის მჟავა – Georgian" lang="ka" hreflang="ka" data-title="დეზოქსირიბონუკლეინის მჟავა" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target"><span>ქართული</span></a></li><li class="interlanguage-link interwiki-ks mw-list-item"><a href="https://ks.wikipedia.org/wiki/%DA%88%DB%8C_%D8%A7%DB%8C%D9%86_%D8%A7%DB%92" title="ڈی این اے – Kashmiri" lang="ks" hreflang="ks" data-title="ڈی این اے" data-language-autonym="कॉशुर / کٲشُر" data-language-local-name="Kashmiri" class="interlanguage-link-target"><span>कॉशुर / کٲشُر</span></a></li><li class="interlanguage-link interwiki-csb mw-list-item"><a href="https://csb.wikipedia.org/wiki/DNA" title="DNA – Kashubian" lang="csb" hreflang="csb" data-title="DNA" data-language-autonym="Kaszëbsczi" data-language-local-name="Kashubian" class="interlanguage-link-target"><span>Kaszëbsczi</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%94%D0%9D%D2%9A" title="ДНҚ – Kazakh" lang="kk" hreflang="kk" data-title="ДНҚ" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-kw mw-list-item"><a href="https://kw.wikipedia.org/wiki/TDN" title="TDN – Cornish" lang="kw" hreflang="kw" data-title="TDN" data-language-autonym="Kernowek" data-language-local-name="Cornish" class="interlanguage-link-target"><span>Kernowek</span></a></li><li class="interlanguage-link interwiki-sw mw-list-item"><a href="https://sw.wikipedia.org/wiki/DNA" title="DNA – Swahili" lang="sw" hreflang="sw" data-title="DNA" data-language-autonym="Kiswahili" data-language-local-name="Swahili" class="interlanguage-link-target"><span>Kiswahili</span></a></li><li class="interlanguage-link interwiki-ht mw-list-item"><a href="https://ht.wikipedia.org/wiki/ADN" title="ADN – Haitian Creole" lang="ht" hreflang="ht" data-title="ADN" data-language-autonym="Kreyòl ayisyen" data-language-local-name="Haitian Creole" class="interlanguage-link-target"><span>Kreyòl ayisyen</span></a></li><li class="interlanguage-link interwiki-gcr mw-list-item"><a href="https://gcr.wikipedia.org/wiki/Asid_d%C3%A9zoksoribonikl%C3%A9yik" title="Asid dézoksoribonikléyik – Guianan Creole" lang="gcr" hreflang="gcr" data-title="Asid dézoksoribonikléyik" data-language-autonym="Kriyòl gwiyannen" data-language-local-name="Guianan Creole" class="interlanguage-link-target"><span>Kriyòl gwiyannen</span></a></li><li class="interlanguage-link interwiki-ku mw-list-item"><a href="https://ku.wikipedia.org/wiki/As%C3%AEda_deoks%C3%AEr%C3%AEbonukley%C3%AE" title="Asîda deoksîrîbonukleyî – Kurdish" lang="ku" hreflang="ku" data-title="Asîda deoksîrîbonukleyî" data-language-autonym="Kurdî" data-language-local-name="Kurdish" class="interlanguage-link-target"><span>Kurdî</span></a></li><li class="interlanguage-link interwiki-ky mw-list-item"><a href="https://ky.wikipedia.org/wiki/%D0%94%D0%B5%D0%B7%D0%BE%D0%BA%D1%81%D0%B8%D1%80%D0%B8%D0%B1%D0%BE%D0%BD%D1%83%D0%BA%D0%BB%D0%B5%D0%B8%D0%BD_%D0%BA%D0%B8%D1%81%D0%BB%D0%BE%D1%82%D0%B0%D1%81%D1%8B" title="Дезоксирибонуклеин кислотасы – Kyrgyz" lang="ky" hreflang="ky" data-title="Дезоксирибонуклеин кислотасы" data-language-autonym="Кыргызча" data-language-local-name="Kyrgyz" class="interlanguage-link-target"><span>Кыргызча</span></a></li><li class="interlanguage-link interwiki-lld mw-list-item"><a href="https://lld.wikipedia.org/wiki/DNA" title="DNA – Ladin" lang="lld" hreflang="lld" data-title="DNA" data-language-autonym="Ladin" data-language-local-name="Ladin" class="interlanguage-link-target"><span>Ladin</span></a></li><li class="interlanguage-link interwiki-lo mw-list-item"><a href="https://lo.wikipedia.org/wiki/%E0%BA%AD%E0%BA%B2%E0%BB%80%E0%BA%94%E0%BB%81%E0%BA%AD%E0%BA%B1%E0%BA%99" title="ອາເດແອັນ – Lao" lang="lo" hreflang="lo" data-title="ອາເດແອັນ" data-language-autonym="ລາວ" data-language-local-name="Lao" class="interlanguage-link-target"><span>ລາວ</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Acidum_desoxyribonucleicum" title="Acidum desoxyribonucleicum – Latin" lang="la" hreflang="la" data-title="Acidum desoxyribonucleicum" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-lv mw-list-item"><a href="https://lv.wikipedia.org/wiki/Dezoksiribonukle%C4%ABnsk%C4%81be" title="Dezoksiribonukleīnskābe – Latvian" lang="lv" hreflang="lv" data-title="Dezoksiribonukleīnskābe" data-language-autonym="Latviešu" data-language-local-name="Latvian" class="interlanguage-link-target"><span>Latviešu</span></a></li><li class="interlanguage-link interwiki-lez mw-list-item"><a href="https://lez.wikipedia.org/wiki/%D0%94%D0%B5%D0%B7%D0%BE%D0%BA%D1%81%D0%B8%D1%80%D0%B8%D0%B1%D0%BE%D0%BD%D1%83%D0%BA%D0%BB%D0%B5%D0%B8%D0%BD_%D1%86%D1%83%D1%80%D1%83%D1%86%D3%80" title="Дезоксирибонуклеин цуруцӀ – Lezghian" lang="lez" hreflang="lez" data-title="Дезоксирибонуклеин цуруцӀ" data-language-autonym="Лезги" data-language-local-name="Lezghian" class="interlanguage-link-target"><span>Лезги</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Deoksiribonukleor%C5%ABg%C5%A1tis" title="Deoksiribonukleorūgštis – Lithuanian" lang="lt" hreflang="lt" data-title="Deoksiribonukleorūgštis" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target"><span>Lietuvių</span></a></li><li class="interlanguage-link interwiki-li mw-list-item"><a href="https://li.wikipedia.org/wiki/DNA" title="DNA – Limburgish" lang="li" hreflang="li" data-title="DNA" data-language-autonym="Limburgs" data-language-local-name="Limburgish" class="interlanguage-link-target"><span>Limburgs</span></a></li><li class="interlanguage-link interwiki-lfn mw-list-item"><a href="https://lfn.wikipedia.org/wiki/Asida_desosiribonucleal" title="Asida desosiribonucleal – Lingua Franca Nova" lang="lfn" hreflang="lfn" data-title="Asida desosiribonucleal" data-language-autonym="Lingua Franca Nova" data-language-local-name="Lingua Franca Nova" class="interlanguage-link-target"><span>Lingua Franca Nova</span></a></li><li class="interlanguage-link interwiki-lmo mw-list-item"><a href="https://lmo.wikipedia.org/wiki/DNA" title="DNA – Lombard" lang="lmo" hreflang="lmo" data-title="DNA" data-language-autonym="Lombard" data-language-local-name="Lombard" class="interlanguage-link-target"><span>Lombard</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Dezoxiribonukleinsav" title="Dezoxiribonukleinsav – Hungarian" lang="hu" hreflang="hu" data-title="Dezoxiribonukleinsav" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%94%D0%9D%D0%9A" title="ДНК – Macedonian" lang="mk" hreflang="mk" data-title="ДНК" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target"><span>Македонски</span></a></li><li class="interlanguage-link interwiki-mg mw-list-item"><a href="https://mg.wikipedia.org/wiki/Asidra_dez%C3%B4ksirib%C3%B4niokleika" title="Asidra dezôksiribôniokleika – Malagasy" lang="mg" hreflang="mg" data-title="Asidra dezôksiribôniokleika" data-language-autonym="Malagasy" data-language-local-name="Malagasy" class="interlanguage-link-target"><span>Malagasy</span></a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%A1%E0%B4%BF.%E0%B4%8E%E0%B5%BB.%E0%B4%8E" title="ഡി.എൻ.എ – Malayalam" lang="ml" hreflang="ml" data-title="ഡി.എൻ.എ" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target"><span>മലയാളം</span></a></li><li class="interlanguage-link interwiki-mr mw-list-item"><a href="https://mr.wikipedia.org/wiki/%E0%A4%A1%E0%A4%BF%E0%A4%91%E0%A4%95%E0%A5%8D%E0%A4%B8%E0%A4%BF%E0%A4%B0%E0%A4%BE%E0%A4%AF%E0%A4%AC%E0%A5%8B_%E0%A4%A8%E0%A5%8D%E0%A4%AF%E0%A5%82%E0%A4%95%E0%A5%8D%E0%A4%B2%E0%A5%87%E0%A4%87%E0%A4%95_%E0%A4%86%E0%A4%AE%E0%A5%8D%E0%A4%B2" title="डिऑक्सिरायबो न्यूक्लेइक आम्ल – Marathi" lang="mr" hreflang="mr" data-title="डिऑक्सिरायबो न्यूक्लेइक आम्ल" data-language-autonym="मराठी" data-language-local-name="Marathi" class="interlanguage-link-target"><span>मराठी</span></a></li><li class="interlanguage-link interwiki-xmf mw-list-item"><a href="https://xmf.wikipedia.org/wiki/%E1%83%93%E1%83%94%E1%83%96%E1%83%9D%E1%83%A5%E1%83%A1%E1%83%98%E1%83%A0%E1%83%98%E1%83%91%E1%83%9D%E1%83%9C%E1%83%A3%E1%83%99%E1%83%9A%E1%83%94%E1%83%98%E1%83%9C%E1%83%98%E1%83%A8_%E1%83%91%E1%83%9F%E1%83%94" title="დეზოქსირიბონუკლეინიშ ბჟე – Mingrelian" lang="xmf" hreflang="xmf" data-title="დეზოქსირიბონუკლეინიშ ბჟე" data-language-autonym="მარგალური" data-language-local-name="Mingrelian" class="interlanguage-link-target"><span>მარგალური</span></a></li><li class="interlanguage-link interwiki-arz mw-list-item"><a href="https://arz.wikipedia.org/wiki/%D8%AD%D9%85%D8%B6_%D9%86%D9%88%D9%88%D9%89" title="حمض نووى – Egyptian Arabic" lang="arz" hreflang="arz" data-title="حمض نووى" data-language-autonym="مصرى" data-language-local-name="Egyptian Arabic" class="interlanguage-link-target"><span>مصرى</span></a></li><li class="interlanguage-link interwiki-mzn mw-list-item"><a href="https://mzn.wikipedia.org/wiki/DNA" title="DNA – Mazanderani" lang="mzn" hreflang="mzn" data-title="DNA" data-language-autonym="مازِرونی" data-language-local-name="Mazanderani" class="interlanguage-link-target"><span>مازِرونی</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Asid_deoksiribonukleik" title="Asid deoksiribonukleik – Malay" lang="ms" hreflang="ms" data-title="Asid deoksiribonukleik" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-cdo mw-list-item"><a href="https://cdo.wikipedia.org/wiki/DNA" title="DNA – Mindong" lang="cdo" hreflang="cdo" data-title="DNA" data-language-autonym="閩東語 / Mìng-dĕ̤ng-ngṳ̄" data-language-local-name="Mindong" class="interlanguage-link-target"><span>閩東語 / Mìng-dĕ̤ng-ngṳ̄</span></a></li><li class="interlanguage-link interwiki-mn badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://mn.wikipedia.org/wiki/%D0%94%D0%B5%D0%B7%D0%BE%D0%BA%D1%81%D0%B8%D1%80%D0%B8%D0%B1%D0%BE%D0%BD%D1%83%D0%BA%D0%BB%D0%B5%D0%B9%D0%BD_%D1%85%D2%AF%D1%87%D0%B8%D0%BB" title="Дезоксирибонуклейн хүчил – Mongolian" lang="mn" hreflang="mn" data-title="Дезоксирибонуклейн хүчил" data-language-autonym="Монгол" data-language-local-name="Mongolian" class="interlanguage-link-target"><span>Монгол</span></a></li><li class="interlanguage-link interwiki-my mw-list-item"><a href="https://my.wikipedia.org/wiki/%E1%80%92%E1%80%AE%E1%80%A1%E1%80%94%E1%80%BA%E1%80%A1%E1%80%B1" title="ဒီအန်အေ – Burmese" lang="my" hreflang="my" data-title="ဒီအန်အေ" data-language-autonym="မြန်မာဘာသာ" data-language-local-name="Burmese" class="interlanguage-link-target"><span>မြန်မာဘာသာ</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/DNA_(biologie)" title="DNA (biologie) – Dutch" lang="nl" hreflang="nl" data-title="DNA (biologie)" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ne mw-list-item"><a href="https://ne.wikipedia.org/wiki/%E0%A4%A1%E0%A5%80%E0%A4%8F%E0%A4%A8%E0%A4%8F" title="डीएनए – Nepali" lang="ne" hreflang="ne" data-title="डीएनए" data-language-autonym="नेपाली" data-language-local-name="Nepali" class="interlanguage-link-target"><span>नेपाली</span></a></li><li class="interlanguage-link interwiki-new mw-list-item"><a href="https://new.wikipedia.org/wiki/%E0%A4%A1%E0%A5%80_%E0%A4%8F%E0%A4%A8_%E0%A4%8F" title="डी एन ए – Newari" lang="new" hreflang="new" data-title="डी एन ए" data-language-autonym="नेपाल भाषा" data-language-local-name="Newari" class="interlanguage-link-target"><span>नेपाल भाषा</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E3%83%87%E3%82%AA%E3%82%AD%E3%82%B7%E3%83%AA%E3%83%9C%E6%A0%B8%E9%85%B8" title="デオキシリボ核酸 – Japanese" lang="ja" hreflang="ja" data-title="デオキシリボ核酸" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-ce mw-list-item"><a href="https://ce.wikipedia.org/wiki/%D0%94%D0%B5%D0%B7%D0%BE%D0%BA%D1%81%D0%B8%D1%80%D0%B8%D0%B1%D0%BE%D0%BD%D1%83%D0%BA%D0%BB%D0%B5%D0%B8%D0%BD%D0%B0%D0%BD_%D0%BC%D1%83%D1%8C%D1%81%D1%82%D0%B0%D0%BB" title="Дезоксирибонуклеинан муьстал – Chechen" lang="ce" hreflang="ce" data-title="Дезоксирибонуклеинан муьстал" data-language-autonym="Нохчийн" data-language-local-name="Chechen" class="interlanguage-link-target"><span>Нохчийн</span></a></li><li class="interlanguage-link interwiki-frr mw-list-item"><a href="https://frr.wikipedia.org/wiki/DNA" title="DNA – Northern Frisian" lang="frr" hreflang="frr" data-title="DNA" data-language-autonym="Nordfriisk" data-language-local-name="Northern Frisian" class="interlanguage-link-target"><span>Nordfriisk</span></a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/DNA" title="DNA – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="DNA" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target"><span>Norsk bokmål</span></a></li><li class="interlanguage-link interwiki-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/DNA" title="DNA – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="DNA" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target"><span>Norsk nynorsk</span></a></li><li class="interlanguage-link interwiki-nov mw-list-item"><a href="https://nov.wikipedia.org/wiki/DNA" title="DNA – Novial" lang="nov" hreflang="nov" data-title="DNA" data-language-autonym="Novial" data-language-local-name="Novial" class="interlanguage-link-target"><span>Novial</span></a></li><li class="interlanguage-link interwiki-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/Acid_desoxiribonucle%C3%AFc" title="Acid desoxiribonucleïc – Occitan" lang="oc" hreflang="oc" data-title="Acid desoxiribonucleïc" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target"><span>Occitan</span></a></li><li class="interlanguage-link interwiki-om mw-list-item"><a href="https://om.wikipedia.org/wiki/DNA" title="DNA – Oromo" lang="om" hreflang="om" data-title="DNA" data-language-autonym="Oromoo" data-language-local-name="Oromo" class="interlanguage-link-target"><span>Oromoo</span></a></li><li class="interlanguage-link interwiki-uz mw-list-item"><a href="https://uz.wikipedia.org/wiki/Dezoksiribonuklein_kislota" title="Dezoksiribonuklein kislota – Uzbek" lang="uz" hreflang="uz" data-title="Dezoksiribonuklein kislota" data-language-autonym="Oʻzbekcha / ўзбекча" data-language-local-name="Uzbek" class="interlanguage-link-target"><span>Oʻzbekcha / ўзбекча</span></a></li><li class="interlanguage-link interwiki-pa mw-list-item"><a href="https://pa.wikipedia.org/wiki/%E0%A8%A1%E0%A9%80.%E0%A8%90%E0%A9%B1%E0%A8%A8.%E0%A8%8F." title="ਡੀ.ਐੱਨ.ਏ. – Punjabi" lang="pa" hreflang="pa" data-title="ਡੀ.ਐੱਨ.ਏ." data-language-autonym="ਪੰਜਾਬੀ" data-language-local-name="Punjabi" class="interlanguage-link-target"><span>ਪੰਜਾਬੀ</span></a></li><li class="interlanguage-link interwiki-pnb mw-list-item"><a href="https://pnb.wikipedia.org/wiki/%DA%88%DB%8C_%D8%A7%DB%8C%D9%86_%D8%A7%DB%92" title="ڈی این اے – Western Punjabi" lang="pnb" hreflang="pnb" data-title="ڈی این اے" data-language-autonym="پنجابی" data-language-local-name="Western Punjabi" class="interlanguage-link-target"><span>پنجابی</span></a></li><li class="interlanguage-link interwiki-blk mw-list-item"><a href="https://blk.wikipedia.org/wiki/%E1%80%92%E1%80%AE%E1%80%A1%E1%80%B2%E1%80%89%E1%80%BA%E1%80%A1%E1%80%B1" title="ဒီအဲဉ်အေ – Pa'O" lang="blk" hreflang="blk" data-title="ဒီအဲဉ်အေ" data-language-autonym="ပအိုဝ်ႏဘာႏသာႏ" data-language-local-name="Pa'O" class="interlanguage-link-target"><span>ပအိုဝ်ႏဘာႏသာႏ</span></a></li><li class="interlanguage-link interwiki-pap mw-list-item"><a href="https://pap.wikipedia.org/wiki/ADN" title="ADN – Papiamento" lang="pap" hreflang="pap" data-title="ADN" data-language-autonym="Papiamentu" data-language-local-name="Papiamento" class="interlanguage-link-target"><span>Papiamentu</span></a></li><li class="interlanguage-link interwiki-ps mw-list-item"><a href="https://ps.wikipedia.org/wiki/%DA%89%D9%8A_%D8%A2%DA%A9%D8%B3%D9%8A_%D8%B1%D8%A7%D9%8A%D8%A8%D9%88%D9%86%D9%8A%D9%88%DA%A9%D9%84%D9%8A%DA%A9_%D8%A7%D8%B3%D9%8A%D8%AF" title="ډي آکسي رايبونيوکليک اسيد – Pashto" lang="ps" hreflang="ps" data-title="ډي آکسي رايبونيوکليک اسيد" data-language-autonym="پښتو" data-language-local-name="Pashto" class="interlanguage-link-target"><span>پښتو</span></a></li><li class="interlanguage-link interwiki-jam mw-list-item"><a href="https://jam.wikipedia.org/wiki/Diaxiraibonyuukliik_asid" title="Diaxiraibonyuukliik asid – Jamaican Creole English" lang="jam" hreflang="jam" data-title="Diaxiraibonyuukliik asid" data-language-autonym="Patois" data-language-local-name="Jamaican Creole English" class="interlanguage-link-target"><span>Patois</span></a></li><li class="interlanguage-link interwiki-km mw-list-item"><a href="https://km.wikipedia.org/wiki/%E1%9E%8C%E1%9E%B8%E1%9E%A2%E1%9E%B7%E1%9E%93%E1%9E%A2%E1%9F%81" title="ឌីអិនអេ – Khmer" lang="km" hreflang="km" data-title="ឌីអិនអេ" data-language-autonym="ភាសាខ្មែរ" data-language-local-name="Khmer" class="interlanguage-link-target"><span>ភាសាខ្មែរ</span></a></li><li class="interlanguage-link interwiki-pms mw-list-item"><a href="https://pms.wikipedia.org/wiki/DNA" title="DNA – Piedmontese" lang="pms" hreflang="pms" data-title="DNA" data-language-autonym="Piemontèis" data-language-local-name="Piedmontese" class="interlanguage-link-target"><span>Piemontèis</span></a></li><li class="interlanguage-link interwiki-tpi mw-list-item"><a href="https://tpi.wikipedia.org/wiki/DNA" title="DNA – Tok Pisin" lang="tpi" hreflang="tpi" data-title="DNA" data-language-autonym="Tok Pisin" data-language-local-name="Tok Pisin" class="interlanguage-link-target"><span>Tok Pisin</span></a></li><li class="interlanguage-link interwiki-nds mw-list-item"><a href="https://nds.wikipedia.org/wiki/Desoxyribonukleins%C3%BCre" title="Desoxyribonukleinsüre – Low German" lang="nds" hreflang="nds" data-title="Desoxyribonukleinsüre" data-language-autonym="Plattdüütsch" data-language-local-name="Low German" class="interlanguage-link-target"><span>Plattdüütsch</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Kwas_deoksyrybonukleinowy" title="Kwas deoksyrybonukleinowy – Polish" lang="pl" hreflang="pl" data-title="Kwas deoksyrybonukleinowy" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-pt badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://pt.wikipedia.org/wiki/%C3%81cido_desoxirribonucleico" title="Ácido desoxirribonucleico – Portuguese" lang="pt" hreflang="pt" data-title="Ácido desoxirribonucleico" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/ADN" title="ADN – Romanian" lang="ro" hreflang="ro" data-title="ADN" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</span></a></li><li class="interlanguage-link interwiki-qu mw-list-item"><a href="https://qu.wikipedia.org/wiki/DNK" title="DNK – Quechua" lang="qu" hreflang="qu" data-title="DNK" data-language-autonym="Runa Simi" data-language-local-name="Quechua" class="interlanguage-link-target"><span>Runa Simi</span></a></li><li class="interlanguage-link interwiki-rue mw-list-item"><a href="https://rue.wikipedia.org/wiki/DNA" title="DNA – Rusyn" lang="rue" hreflang="rue" data-title="DNA" data-language-autonym="Русиньскый" data-language-local-name="Rusyn" class="interlanguage-link-target"><span>Русиньскый</span></a></li><li class="interlanguage-link interwiki-ru badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://ru.wikipedia.org/wiki/%D0%94%D0%B5%D0%B7%D0%BE%D0%BA%D1%81%D0%B8%D1%80%D0%B8%D0%B1%D0%BE%D0%BD%D1%83%D0%BA%D0%BB%D0%B5%D0%B8%D0%BD%D0%BE%D0%B2%D0%B0%D1%8F_%D0%BA%D0%B8%D1%81%D0%BB%D0%BE%D1%82%D0%B0" title="Дезоксирибонуклеиновая кислота – Russian" lang="ru" hreflang="ru" data-title="Дезоксирибонуклеиновая кислота" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sah mw-list-item"><a href="https://sah.wikipedia.org/wiki/%D0%94%D0%B5%D0%B7%D0%BE%D0%BA%D1%81%D0%B8%D1%80%D0%B8%D0%B1%D0%BE%D0%BD%D1%83%D0%BA%D0%BB%D0%B5%D0%B8%D0%BD%D0%BD%D0%B0%D0%B0%D1%85_%D0%B0%D2%BB%D1%8B%D1%8B%D0%B1%D0%B0" title="Дезоксирибонуклеиннаах аһыыба – Yakut" lang="sah" hreflang="sah" data-title="Дезоксирибонуклеиннаах аһыыба" data-language-autonym="Саха тыла" data-language-local-name="Yakut" class="interlanguage-link-target"><span>Саха тыла</span></a></li><li class="interlanguage-link interwiki-sc mw-list-item"><a href="https://sc.wikipedia.org/wiki/ADN" title="ADN – Sardinian" lang="sc" hreflang="sc" data-title="ADN" data-language-autonym="Sardu" data-language-local-name="Sardinian" class="interlanguage-link-target"><span>Sardu</span></a></li><li class="interlanguage-link interwiki-sco mw-list-item"><a href="https://sco.wikipedia.org/wiki/DNA" title="DNA – Scots" lang="sco" hreflang="sco" data-title="DNA" data-language-autonym="Scots" data-language-local-name="Scots" class="interlanguage-link-target"><span>Scots</span></a></li><li class="interlanguage-link interwiki-stq mw-list-item"><a href="https://stq.wikipedia.org/wiki/Desoxyribonukleins%C3%BC%C3%BCre" title="Desoxyribonukleinsüüre – Saterland Frisian" lang="stq" hreflang="stq" data-title="Desoxyribonukleinsüüre" data-language-autonym="Seeltersk" data-language-local-name="Saterland Frisian" class="interlanguage-link-target"><span>Seeltersk</span></a></li><li class="interlanguage-link interwiki-sq mw-list-item"><a href="https://sq.wikipedia.org/wiki/ADN" title="ADN – Albanian" lang="sq" hreflang="sq" data-title="ADN" data-language-autonym="Shqip" data-language-local-name="Albanian" class="interlanguage-link-target"><span>Shqip</span></a></li><li class="interlanguage-link interwiki-scn mw-list-item"><a href="https://scn.wikipedia.org/wiki/DNA" title="DNA – Sicilian" lang="scn" hreflang="scn" data-title="DNA" data-language-autonym="Sicilianu" data-language-local-name="Sicilian" class="interlanguage-link-target"><span>Sicilianu</span></a></li><li class="interlanguage-link interwiki-si mw-list-item"><a href="https://si.wikipedia.org/wiki/%E0%B6%A9%E0%B7%93.%E0%B6%91%E0%B6%B1%E0%B7%8A.%E0%B6%92." title="ඩී.එන්.ඒ. – Sinhala" lang="si" hreflang="si" data-title="ඩී.එන්.ඒ." data-language-autonym="සිංහල" data-language-local-name="Sinhala" class="interlanguage-link-target"><span>සිංහල</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/DNA" title="DNA – Simple English" lang="en-simple" hreflang="en-simple" data-title="DNA" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-sd mw-list-item"><a href="https://sd.wikipedia.org/wiki/%DA%8A%D9%8A_%D8%A7%D9%8A%D9%86_%D8%A7%D9%8A" title="ڊي اين اي – Sindhi" lang="sd" hreflang="sd" data-title="ڊي اين اي" data-language-autonym="سنڌي" data-language-local-name="Sindhi" class="interlanguage-link-target"><span>سنڌي</span></a></li><li class="interlanguage-link interwiki-sk badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://sk.wikipedia.org/wiki/Deoxyribonukleov%C3%A1_kyselina" title="Deoxyribonukleová kyselina – Slovak" lang="sk" hreflang="sk" data-title="Deoxyribonukleová kyselina" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Deoksiribonukleinska_kislina" title="Deoksiribonukleinska kislina – Slovenian" lang="sl" hreflang="sl" data-title="Deoksiribonukleinska kislina" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-szl mw-list-item"><a href="https://szl.wikipedia.org/wiki/DNA" title="DNA – Silesian" lang="szl" hreflang="szl" data-title="DNA" data-language-autonym="Ślůnski" data-language-local-name="Silesian" class="interlanguage-link-target"><span>Ślůnski</span></a></li><li class="interlanguage-link interwiki-so mw-list-item"><a href="https://so.wikipedia.org/wiki/DNA" title="DNA – Somali" lang="so" hreflang="so" data-title="DNA" data-language-autonym="Soomaaliga" data-language-local-name="Somali" class="interlanguage-link-target"><span>Soomaaliga</span></a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D8%AF%DB%8C_%D8%A6%DB%8E%D9%86_%D8%A6%DB%95%DB%8C" title="دی ئێن ئەی – Central Kurdish" lang="ckb" hreflang="ckb" data-title="دی ئێن ئەی" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target"><span>کوردی</span></a></li><li class="interlanguage-link interwiki-sr badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://sr.wikipedia.org/wiki/%D0%94%D0%B5%D0%B7%D0%BE%D0%BA%D1%81%D0%B8%D1%80%D0%B8%D0%B1%D0%BE%D0%BD%D1%83%D0%BA%D0%BB%D0%B5%D0%B8%D0%BD%D1%81%D0%BA%D0%B0_%D0%BA%D0%B8%D1%81%D0%B5%D0%BB%D0%B8%D0%BD%D0%B0" title="Дезоксирибонуклеинска киселина – Serbian" lang="sr" hreflang="sr" data-title="Дезоксирибонуклеинска киселина" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target"><span>Српски / srpski</span></a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/DNK" title="DNK – Serbo-Croatian" lang="sh" hreflang="sh" data-title="DNK" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target"><span>Srpskohrvatski / српскохрватски</span></a></li><li class="interlanguage-link interwiki-su mw-list-item"><a href="https://su.wikipedia.org/wiki/DNA" title="DNA – Sundanese" lang="su" hreflang="su" data-title="DNA" data-language-autonym="Sunda" data-language-local-name="Sundanese" class="interlanguage-link-target"><span>Sunda</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/DNA" title="DNA – Finnish" lang="fi" hreflang="fi" data-title="DNA" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/DNA" title="DNA – Swedish" lang="sv" hreflang="sv" data-title="DNA" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-tl mw-list-item"><a href="https://tl.wikipedia.org/wiki/DNA" title="DNA – Tagalog" lang="tl" hreflang="tl" data-title="DNA" data-language-autonym="Tagalog" data-language-local-name="Tagalog" class="interlanguage-link-target"><span>Tagalog</span></a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%9F%E0%AE%BF._%E0%AE%8E%E0%AE%A9%E0%AF%8D._%E0%AE%8F." title="டி. என். ஏ. – Tamil" lang="ta" hreflang="ta" data-title="டி. என். ஏ." data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target"><span>தமிழ்</span></a></li><li class="interlanguage-link interwiki-tt mw-list-item"><a href="https://tt.wikipedia.org/wiki/%D0%94%D0%B5%D0%B7%D0%BE%D0%BA%D1%81%D0%B8%D1%80%D0%B8%D0%B1%D0%BE%D0%BD%D1%83%D0%BA%D0%BB%D0%B5%D0%B8%D0%BD_%D0%BA%D0%B8%D1%81%D0%BB%D0%BE%D1%82%D0%B0%D1%81%D1%8B" title="Дезоксирибонуклеин кислотасы – Tatar" lang="tt" hreflang="tt" data-title="Дезоксирибонуклеин кислотасы" data-language-autonym="Татарча / tatarça" data-language-local-name="Tatar" class="interlanguage-link-target"><span>Татарча / tatarça</span></a></li><li class="interlanguage-link interwiki-te mw-list-item"><a href="https://te.wikipedia.org/wiki/%E0%B0%A1%E0%B1%80%E0%B0%86%E0%B0%95%E0%B1%8D%E0%B0%B8%E0%B1%80%E0%B0%B0%E0%B1%88%E0%B0%AC%E0%B1%8B_%E0%B0%95%E0%B1%87%E0%B0%82%E0%B0%A6%E0%B1%8D%E0%B0%B0%E0%B0%95_%E0%B0%86%E0%B0%AE%E0%B1%8D%E0%B0%B2%E0%B0%82" title="డీఆక్సీరైబో కేంద్రక ఆమ్లం – Telugu" lang="te" hreflang="te" data-title="డీఆక్సీరైబో కేంద్రక ఆమ్లం" data-language-autonym="తెలుగు" data-language-local-name="Telugu" class="interlanguage-link-target"><span>తెలుగు</span></a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B8%94%E0%B8%B5%E0%B9%80%E0%B8%AD%E0%B9%87%E0%B8%99%E0%B9%80%E0%B8%AD" title="ดีเอ็นเอ – Thai" lang="th" hreflang="th" data-title="ดีเอ็นเอ" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target"><span>ไทย</span></a></li><li class="interlanguage-link interwiki-tg mw-list-item"><a href="https://tg.wikipedia.org/wiki/%D0%9A%D0%B8%D1%81%D0%BB%D0%BE%D1%82%D0%B0%D0%B8_%D0%B4%D0%B5%D0%B7%D0%BE%D0%BA%D1%81%D0%B8%D1%80%D0%B8%D0%B1%D0%BE%D0%BD%D1%83%D0%BA%D0%BB%D0%B5%D0%B0%D1%82" title="Кислотаи дезоксирибонуклеат – Tajik" lang="tg" hreflang="tg" data-title="Кислотаи дезоксирибонуклеат" data-language-autonym="Тоҷикӣ" data-language-local-name="Tajik" class="interlanguage-link-target"><span>Тоҷикӣ</span></a></li><li class="interlanguage-link interwiki-tr badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://tr.wikipedia.org/wiki/DNA" title="DNA – Turkish" lang="tr" hreflang="tr" data-title="DNA" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://uk.wikipedia.org/wiki/%D0%94%D0%B5%D0%B7%D0%BE%D0%BA%D1%81%D0%B8%D1%80%D0%B8%D0%B1%D0%BE%D0%BD%D1%83%D0%BA%D0%BB%D0%B5%D1%97%D0%BD%D0%BE%D0%B2%D0%B0_%D0%BA%D0%B8%D1%81%D0%BB%D0%BE%D1%82%D0%B0" title="Дезоксирибонуклеїнова кислота – Ukrainian" lang="uk" hreflang="uk" data-title="Дезоксирибонуклеїнова кислота" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-ur mw-list-item"><a href="https://ur.wikipedia.org/wiki/%DA%88%DB%8C_%D8%A7%DB%8C%D9%86_%D8%A7%DB%92" title="ڈی این اے – Urdu" lang="ur" hreflang="ur" data-title="ڈی این اے" data-language-autonym="اردو" data-language-local-name="Urdu" class="interlanguage-link-target"><span>اردو</span></a></li><li class="interlanguage-link interwiki-ug mw-list-item"><a href="https://ug.wikipedia.org/wiki/%D8%AF%DB%90%D8%A6%D9%88%D9%83%D8%B3%D9%89%D8%B1%D9%89%D8%A8%D9%88%D9%86%DB%87%D9%83%D9%84%DB%90%D8%A6%D9%89%D9%83_%D9%83%D9%89%D8%B3%D9%84%D8%A7%D8%AA%D8%A7" title="دېئوكسىرىبونۇكلېئىك كىسلاتا – Uyghur" lang="ug" hreflang="ug" data-title="دېئوكسىرىبونۇكلېئىك كىسلاتا" data-language-autonym="ئۇيغۇرچە / Uyghurche" data-language-local-name="Uyghur" class="interlanguage-link-target"><span>ئۇيغۇرچە / Uyghurche</span></a></li><li class="interlanguage-link interwiki-vec mw-list-item"><a href="https://vec.wikipedia.org/wiki/DNA" title="DNA – Venetian" lang="vec" hreflang="vec" data-title="DNA" data-language-autonym="Vèneto" data-language-local-name="Venetian" class="interlanguage-link-target"><span>Vèneto</span></a></li><li class="interlanguage-link interwiki-vi badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://vi.wikipedia.org/wiki/DNA" title="DNA – Vietnamese" lang="vi" hreflang="vi" data-title="DNA" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li><li class="interlanguage-link interwiki-wa mw-list-item"><a href="https://wa.wikipedia.org/wiki/Seur_dezocsiribonucleyike" title="Seur dezocsiribonucleyike – Walloon" lang="wa" hreflang="wa" data-title="Seur dezocsiribonucleyike" data-language-autonym="Walon" data-language-local-name="Walloon" class="interlanguage-link-target"><span>Walon</span></a></li><li class="interlanguage-link interwiki-zh-classical mw-list-item"><a href="https://zh-classical.wikipedia.org/wiki/%E8%84%AB%E6%B0%A7%E6%A0%B8%E7%B3%96%E6%A0%B8%E9%85%B8" title="脫氧核糖核酸 – Literary Chinese" lang="lzh" hreflang="lzh" data-title="脫氧核糖核酸" data-language-autonym="文言" data-language-local-name="Literary Chinese" class="interlanguage-link-target"><span>文言</span></a></li><li class="interlanguage-link interwiki-vls badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://vls.wikipedia.org/wiki/DNA" title="DNA – West Flemish" lang="vls" hreflang="vls" data-title="DNA" data-language-autonym="West-Vlams" data-language-local-name="West Flemish" class="interlanguage-link-target"><span>West-Vlams</span></a></li><li class="interlanguage-link interwiki-war mw-list-item"><a href="https://war.wikipedia.org/wiki/DNA" title="DNA – Waray" lang="war" hreflang="war" data-title="DNA" data-language-autonym="Winaray" data-language-local-name="Waray" class="interlanguage-link-target"><span>Winaray</span></a></li><li class="interlanguage-link interwiki-wuu mw-list-item"><a href="https://wuu.wikipedia.org/wiki/%E8%84%AB%E6%B0%A7%E6%A0%B8%E7%B3%96%E6%A0%B8%E9%85%B8" title="脫氧核糖核酸 – Wu" lang="wuu" hreflang="wuu" data-title="脫氧核糖核酸" data-language-autonym="吴语" data-language-local-name="Wu" class="interlanguage-link-target"><span>吴语</span></a></li><li class="interlanguage-link interwiki-yi mw-list-item"><a href="https://yi.wikipedia.org/wiki/DNA" title="DNA – Yiddish" lang="yi" hreflang="yi" data-title="DNA" data-language-autonym="ייִדיש" data-language-local-name="Yiddish" class="interlanguage-link-target"><span>ייִדיש</span></a></li><li class="interlanguage-link interwiki-yo mw-list-item"><a href="https://yo.wikipedia.org/wiki/DNA" title="DNA – Yoruba" lang="yo" hreflang="yo" data-title="DNA" data-language-autonym="Yorùbá" data-language-local-name="Yoruba" class="interlanguage-link-target"><span>Yorùbá</span></a></li><li class="interlanguage-link interwiki-zh-yue mw-list-item"><a href="https://zh-yue.wikipedia.org/wiki/%E8%84%AB%E6%B0%A7%E6%A0%B8%E7%B3%96%E6%A0%B8%E9%85%B8" title="脫氧核糖核酸 – Cantonese" lang="yue" hreflang="yue" data-title="脫氧核糖核酸" data-language-autonym="粵語" data-language-local-name="Cantonese" class="interlanguage-link-target"><span>粵語</span></a></li><li class="interlanguage-link interwiki-bat-smg mw-list-item"><a href="https://bat-smg.wikipedia.org/wiki/DNR" title="DNR – Samogitian" lang="sgs" hreflang="sgs" 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Click here for more information."><img alt="Featured article" src="//upload.wikimedia.org/wikipedia/en/thumb/e/e7/Cscr-featured.svg/20px-Cscr-featured.svg.png" decoding="async" width="20" height="19" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/e/e7/Cscr-featured.svg/30px-Cscr-featured.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/e/e7/Cscr-featured.svg/40px-Cscr-featured.svg.png 2x" data-file-width="466" data-file-height="443" /></a></span></div></div> <div id="mw-indicator-pp-default" class="mw-indicator"><div class="mw-parser-output"><span typeof="mw:File"><a href="/wiki/Wikipedia:Protection_policy#semi" title="This article is semi-protected due to vandalism"><img alt="Page semi-protected" src="//upload.wikimedia.org/wikipedia/en/thumb/1/1b/Semi-protection-shackle.svg/20px-Semi-protection-shackle.svg.png" decoding="async" width="20" height="20" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/1/1b/Semi-protection-shackle.svg/30px-Semi-protection-shackle.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/1/1b/Semi-protection-shackle.svg/40px-Semi-protection-shackle.svg.png 2x" data-file-width="512" data-file-height="512" /></a></span></div></div> <div id="mw-indicator-spoken-icon" class="mw-indicator"><div class="mw-parser-output"><span typeof="mw:File"><a href="/wiki/File:Dna.ogg" title="Listen to this article"><img alt="Listen to this article" src="//upload.wikimedia.org/wikipedia/commons/thumb/4/47/Sound-icon.svg/20px-Sound-icon.svg.png" decoding="async" width="20" height="15" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/47/Sound-icon.svg/30px-Sound-icon.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/47/Sound-icon.svg/40px-Sound-icon.svg.png 2x" data-file-width="128" data-file-height="96" /></a></span></div></div> </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"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Molecule that carries genetic information</div> <p class="mw-empty-elt"> </p> <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">For a non-technical introduction to the topic, see <a href="/wiki/Introduction_to_genetics" title="Introduction to genetics">Introduction to genetics</a>. For other uses, see <a href="/wiki/DNA_(disambiguation)" class="mw-disambig" title="DNA (disambiguation)">DNA (disambiguation)</a>.</div> <p class="mw-empty-elt"> </p> <table role="presentation" cellpadding="0" style="border-spacing:0; clear:right; float:right;"> <tbody><tr> <td><div class="thumb tright"> <div class="thumbinner" style="width:322px;"> <div style="position:relative; width:320px; height:185px; overflow:hidden; border:solid #ccc 1px; background-color:white;"> <div style="left:0px; top:0px; width:320px; position:absolute"> <span typeof="mw:File"><a href="/wiki/File:Chromosome_DNA_Gene.svg" title="File:Chromosome DNA Gene.svg"><img alt="A chromosome unravelling into a long string of DNA, a section of which is highlighted as the gene" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d0/Chromosome_DNA_Gene_unannotated.svg/320px-Chromosome_DNA_Gene_unannotated.svg.png" decoding="async" width="320" height="175" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d0/Chromosome_DNA_Gene_unannotated.svg/480px-Chromosome_DNA_Gene_unannotated.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d0/Chromosome_DNA_Gene_unannotated.svg/640px-Chromosome_DNA_Gene_unannotated.svg.png 2x" data-file-width="1200" data-file-height="658" /></a></span> </div> <div style="text-align:left; font-size:11px; line-height:110%"> <div style="background-color:transparent; color:black"><div id="annotation_128x52" style="position:absolute; left:128px; top:52px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Chromosome" title="Chromosome">Chromosome </a> </span></div> <div id="annotation_197x52" style="position:absolute; left:197px; top:52px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> (<a href="/wiki/Base_pair#Length_measurements" title="Base pair">10<sup>7</sup> - 10<sup>10</sup> bp</a>)</span></div> <div id="annotation_25x100" style="position:absolute; left:25px; top:100px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a class="mw-selflink selflink">DNA </a> </span></div> <div id="annotation_128x110" style="position:absolute; left:128px; top:110px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Gene" title="Gene">Gene </a> </span></div> <div id="annotation_162x110" style="position:absolute; left:162px; top:110px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> (<a href="/wiki/Base_pair#Length_measurements" title="Base pair">10<sup>3</sup> - 10<sup>6</sup> bp </a>)</span></div> <div id="annotation_154x158" style="position:absolute; left:154px; top:158px; line-height:110%;"><span style="background-color:transparent; color:inherit;"> <a href="/wiki/Phenotype" title="Phenotype">Function </a> </span></div></div> </div> </div> <div class="thumbcaption" style="clear:left"><div style="float:left;margin-right:0.5em"><span typeof="mw:File"><span title="The image above contains clickable links"><img alt="The image above contains clickable links" src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e6/Interactive_icon.svg/18px-Interactive_icon.svg.png" decoding="async" width="18" height="27" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e6/Interactive_icon.svg/27px-Interactive_icon.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e6/Interactive_icon.svg/36px-Interactive_icon.svg.png 2x" data-file-width="133" data-file-height="200" /></span></span></div>A chromosome and its <a href="/wiki/Nuclear_organization" title="Nuclear organization">packaged</a> long strand of DNA unraveled. The DNA's <a href="/wiki/Base_pair" title="Base pair">base pairs</a> encode genes, which provide functions. A human DNA can have up to 500 million base pairs with thousands of genes.</div> </div> </div> </td></tr></tbody></table> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:DNA_Structure%2BKey%2BLabelled.pn_NoBB.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/4c/DNA_Structure%2BKey%2BLabelled.pn_NoBB.png/290px-DNA_Structure%2BKey%2BLabelled.pn_NoBB.png" decoding="async" width="290" height="288" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/4c/DNA_Structure%2BKey%2BLabelled.pn_NoBB.png/435px-DNA_Structure%2BKey%2BLabelled.pn_NoBB.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/4c/DNA_Structure%2BKey%2BLabelled.pn_NoBB.png/580px-DNA_Structure%2BKey%2BLabelled.pn_NoBB.png 2x" data-file-width="3024" data-file-height="3000" /></a><figcaption>The structure of the DNA <a href="/wiki/Double_helix" class="mw-redirect" title="Double helix">double helix</a> (type <a href="/wiki/B-DNA" class="mw-redirect" title="B-DNA">B-DNA</a>). The <a href="/wiki/Atom" title="Atom">atoms</a> in the structure are colour-coded by <a href="/wiki/Chemical_element" title="Chemical element">element</a> and the detailed structures of two <a href="/wiki/Base_pair" title="Base pair">base pairs</a> are shown in the bottom right.</figcaption></figure><figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Phosphate_backbone.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/19/Phosphate_backbone.jpg/220px-Phosphate_backbone.jpg" decoding="async" width="220" height="483" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/19/Phosphate_backbone.jpg/330px-Phosphate_backbone.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/19/Phosphate_backbone.jpg/440px-Phosphate_backbone.jpg 2x" data-file-width="1600" data-file-height="3511" /></a><figcaption>Simplified diagram</figcaption></figure> <style 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.sidebar-list-title,html.skin-theme-clientpref-night .mw-parser-output .sidebar:not(.notheme) .sidebar-title-with-pretitle{background:transparent!important}html.skin-theme-clientpref-night .mw-parser-output .sidebar:not(.notheme) .sidebar-title-with-pretitle a{color:var(--color-progressive)!important}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .sidebar:not(.notheme) .sidebar-list-title,html.skin-theme-clientpref-os .mw-parser-output .sidebar:not(.notheme) .sidebar-title-with-pretitle{background:transparent!important}html.skin-theme-clientpref-os .mw-parser-output .sidebar:not(.notheme) .sidebar-title-with-pretitle a{color:var(--color-progressive)!important}}@media print{body.ns-0 .mw-parser-output .sidebar{display:none!important}}</style><style data-mw-deduplicate="TemplateStyles:r1066933788">.mw-parser-output .excerpt-hat .mw-editsection-like{font-style:normal}</style><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1066933788"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1066933788"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1066933788"><table class="sidebar sidebar-collapse nomobile nowraplinks" style="width:17em;border:2px solid #90db90"><tbody><tr><td class="sidebar-pretitle">Part of a series on</td></tr><tr><th class="sidebar-title-with-pretitle" style="background:#90db90;font-size:175%;font-weight:bold"><a href="/wiki/Genetics" title="Genetics">Genetics</a></th></tr><tr><td class="sidebar-image"><span typeof="mw:File"><a href="/wiki/File:Hybridogenesis_in_water_frogs_gametes.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f0/Hybridogenesis_in_water_frogs_gametes.svg/150px-Hybridogenesis_in_water_frogs_gametes.svg.png" decoding="async" width="150" height="74" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f0/Hybridogenesis_in_water_frogs_gametes.svg/225px-Hybridogenesis_in_water_frogs_gametes.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f0/Hybridogenesis_in_water_frogs_gametes.svg/300px-Hybridogenesis_in_water_frogs_gametes.svg.png 2x" data-file-width="225" data-file-height="111" /></a></span></td></tr><tr><td class="sidebar-content" style="background:#e7f4e7;padding:0.2em 0.4em 0.4em;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90db90;font-size:100%;font-weight:bold;;color: var(--color-base)">Key components</div><div class="sidebar-list-content mw-collapsible-content hlist"><div class="excerpt-block"><div class="excerpt"> <ul><li><a href="/wiki/Chromosome" title="Chromosome">Chromosome</a></li> <li>DNA</li> <li><a href="/wiki/RNA" title="RNA">RNA</a></li> <li><a href="/wiki/Genome" title="Genome">Genome</a></li> <li><a href="/wiki/Heredity" title="Heredity">Heredity</a></li> <li><a href="/wiki/Nucleotide" title="Nucleotide">Nucleotide</a></li> <li><a href="/wiki/Mutation" title="Mutation">Mutation</a></li> <li><a href="/wiki/Genetic_variation" title="Genetic variation">Genetic variation</a></li> <li><a href="/wiki/Allele" title="Allele">Allele</a></li> <li><a href="/wiki/Amino_acid" title="Amino acid">Amino acid</a></li></ul></div></div> <hr /> <ul><li><a href="/wiki/Outline_of_genetics" title="Outline of genetics">Outline</a></li> <li><a href="/wiki/Index_of_genetics_articles" title="Index of genetics articles">Index</a></li></ul></div></div></td> </tr><tr><td class="sidebar-content" style="background:#e7f4e7;padding:0.2em 0.4em 0.4em;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90db90;font-size:100%;font-weight:bold;;color: var(--color-base)">History and topics</div><div class="sidebar-list-content mw-collapsible-content hlist"> <ul><li><a href="/wiki/Introduction_to_genetics" title="Introduction to genetics">Introduction</a></li> <li><a href="/wiki/History_of_genetics" title="History of genetics">History</a></li> <li><a href="/wiki/Evolution" title="Evolution">Evolution</a> (<a href="/wiki/Molecular_evolution" title="Molecular evolution">molecular</a>)</li> <li><a href="/wiki/Population_genetics" title="Population genetics">Population genetics</a></li> <li><a href="/wiki/Mendelian_inheritance" title="Mendelian inheritance">Mendelian inheritance</a></li> <li><a href="/wiki/Quantitative_genetics" title="Quantitative genetics">Quantitative genetics</a></li> <li><a href="/wiki/Molecular_genetics" title="Molecular genetics">Molecular genetics</a></li></ul></div></div></td> </tr><tr><td class="sidebar-content" style="background:#e7f4e7;padding:0.2em 0.4em 0.4em;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90db90;font-size:100%;font-weight:bold;;color: var(--color-base)">Research</div><div class="sidebar-list-content mw-collapsible-content hlist"> <ul><li><a href="/wiki/Geneticist" title="Geneticist">Geneticist</a></li> <li><a href="/wiki/DNA_sequencing" title="DNA sequencing">DNA sequencing</a></li> <li><a href="/wiki/Genetic_engineering" title="Genetic engineering">Genetic engineering</a></li> <li><span class="nowrap"><a href="/wiki/Genomics" title="Genomics">Genomics</a> (<span class="noviewer" typeof="mw:File"><span title="Template"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/83/Symbol_template_class_pink.svg/12px-Symbol_template_class_pink.svg.png" decoding="async" width="12" height="12" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/83/Symbol_template_class_pink.svg/18px-Symbol_template_class_pink.svg.png 1.5x, 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title="Classical genetics">Classical</a></li> <li><a href="/wiki/Conservation_genetics" title="Conservation genetics">Conservation</a></li> <li><a href="/wiki/Cytogenetics" title="Cytogenetics">Cytogenetics</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological</a></li> <li><a href="/wiki/Immunogenetics" title="Immunogenetics">Immunogenetics</a></li> <li><a href="/wiki/Microbial_genetics" title="Microbial genetics">Microbial</a></li> <li><a href="/wiki/Molecular_genetics" title="Molecular genetics">Molecular</a></li> <li><a href="/wiki/Population_genetics" title="Population genetics">Population</a></li> <li><a href="/wiki/Quantitative_genetics" title="Quantitative genetics">Quantitative</a></li></ul></div></div></div></div></td> </tr><tr><td class="sidebar-content" style="background:#e7f4e7;padding:0.2em 0.4em 0.4em;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90db90;font-size:100%;font-weight:bold;;color: var(--color-base)">Personalized medicine</div><div class="sidebar-list-content mw-collapsible-content hlist"> <ul><li><a href="/wiki/Personalized_medicine" title="Personalized medicine">Personalized medicine</a></li></ul></div></div></td> </tr><tr><td class="sidebar-below hlist"> <ul><li><span class="nowrap"><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Genetics" title="Category:Genetics">Category</a></span></li></ul></td></tr><tr><td class="sidebar-navbar"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Genetics_sidebar" title="Template:Genetics sidebar"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Genetics_sidebar" title="Template talk:Genetics sidebar"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Genetics_sidebar" title="Special:EditPage/Template:Genetics sidebar"><abbr title="Edit this template">e</abbr></a></li></ul></div></td></tr></tbody></table> <p><b>Deoxyribonucleic acid</b> (<span class="rt-commentedText nowrap"><span class="IPA nopopups noexcerpt" lang="en-fonipa"><a href="/wiki/Help:IPA/English" title="Help:IPA/English">/<span style="border-bottom:1px dotted"><span title="'d' in 'dye'">d</span><span title="/iː/: 'ee' in 'fleece'">iː</span><span title="/ˈ/: primary stress follows">ˈ</span><span title="/ɒ/: 'o' in 'body'">ɒ</span><span title="'k' in 'kind'">k</span><span title="'s' in 'sigh'">s</span><span title="/ɪ/: 'i' in 'kit'">ɪ</span><span title="/ˌ/: secondary stress follows">ˌ</span><span title="'r' in 'rye'">r</span><span title="/aɪ/: 'i' in 'tide'">aɪ</span><span title="'b' in 'buy'">b</span><span title="/oʊ/: 'o' in 'code'">oʊ</span><span title="/nj/: 'n' in 'new'">nj</span><span title="/uː/: 'oo' in 'goose'">uː</span><span title="/ˌ/: secondary stress follows">ˌ</span><span title="'k' in 'kind'">k</span><span title="'l' in 'lie'">l</span><span title="/iː/: 'ee' in 'fleece'">iː</span><span title="/ɪ/: 'i' in 'kit'">ɪ</span><span title="'k' in 'kind'">k</span></span>,<span class="wrap"> </span>-<span style="border-bottom:1px dotted"><span title="/ˌ/: secondary stress follows">ˌ</span><span title="'k' in 'kind'">k</span><span title="'l' in 'lie'">l</span><span title="/eɪ/: 'a' in 'face'">eɪ</span></span>-/</a></span> <span class="noprint"><span class="ext-phonos"><span data-nosnippet="" id="ooui-php-1" class="ext-phonos-PhonosButton noexcerpt ext-phonos-PhonosButton-emptylabel oo-ui-widget oo-ui-widget-enabled oo-ui-buttonElement oo-ui-buttonElement-frameless oo-ui-iconElement oo-ui-buttonWidget" data-ooui="{"_":"mw.Phonos.PhonosButton","href":"\/\/upload.wikimedia.org\/wikipedia\/commons\/transcoded\/f\/f2\/En-us-Deoxyribonucleic_acid.ogg\/En-us-Deoxyribonucleic_acid.ogg.mp3","rel":["nofollow"],"framed":false,"icon":"volumeUp","data":{"ipa":"","text":"","lang":"en","wikibase":"","file":"En-us-Deoxyribonucleic acid.ogg"},"classes":["ext-phonos-PhonosButton","noexcerpt","ext-phonos-PhonosButton-emptylabel"]}"><a role="button" tabindex="0" href="//upload.wikimedia.org/wikipedia/commons/transcoded/f/f2/En-us-Deoxyribonucleic_acid.ogg/En-us-Deoxyribonucleic_acid.ogg.mp3" rel="nofollow" aria-label="Play audio" title="Play audio" class="oo-ui-buttonElement-button"><span class="oo-ui-iconElement-icon oo-ui-icon-volumeUp"></span><span class="oo-ui-labelElement-label"></span><span class="oo-ui-indicatorElement-indicator oo-ui-indicatorElement-noIndicator"></span></a></span><sup class="ext-phonos-attribution noexcerpt navigation-not-searchable"><a href="/wiki/File:En-us-Deoxyribonucleic_acid.ogg" title="File:En-us-Deoxyribonucleic acid.ogg">ⓘ</a></sup></span></span></span>;<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> <b>DNA</b>) is a <a href="/wiki/Polymer" title="Polymer">polymer</a> composed of two <a href="/wiki/Polynucleotide" title="Polynucleotide">polynucleotide</a> chains that coil around each other to form a <a href="/wiki/Nucleic_acid_double_helix" title="Nucleic acid double helix">double helix</a>. The polymer carries <a href="/wiki/Genetics" title="Genetics">genetic</a> instructions for the development, functioning, growth and <a href="/wiki/Reproduction" title="Reproduction">reproduction</a> of all known <a href="/wiki/Organism" title="Organism">organisms</a> and many <a href="/wiki/Virus" title="Virus">viruses</a>. DNA and <a href="/wiki/Ribonucleic_acid" class="mw-redirect" title="Ribonucleic acid">ribonucleic acid</a> (RNA) are <a href="/wiki/Nucleic_acid" title="Nucleic acid">nucleic acids</a>. Alongside <a href="/wiki/Protein" title="Protein">proteins</a>, <a href="/wiki/Lipids" class="mw-redirect" title="Lipids">lipids</a> and complex carbohydrates (<a href="/wiki/Polysaccharide" title="Polysaccharide">polysaccharides</a>), nucleic acids are one of the four major types of <a href="/wiki/Macromolecule" title="Macromolecule">macromolecules</a> that are essential for all known forms of <a href="/wiki/Life" title="Life">life</a>. </p><p>The two DNA strands are known as polynucleotides as they are composed of simpler <a href="/wiki/Monomer" title="Monomer">monomeric</a> units called <a href="/wiki/Nucleotide" title="Nucleotide">nucleotides</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><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> Each nucleotide is composed of one of four <a href="/wiki/Nitrogenous_base" class="mw-redirect" title="Nitrogenous base">nitrogen-containing</a> <a href="/wiki/Nucleobase" class="mw-redirect" title="Nucleobase">nucleobases</a> (<a href="/wiki/Cytosine" title="Cytosine">cytosine</a> [C], <a href="/wiki/Guanine" title="Guanine">guanine</a> [G], <a href="/wiki/Adenine" title="Adenine">adenine</a> [A] or <a href="/wiki/Thymine" title="Thymine">thymine</a> [T]), a <a href="/wiki/Monosaccharide" title="Monosaccharide">sugar</a> called <a href="/wiki/Deoxyribose" title="Deoxyribose">deoxyribose</a>, and a <a href="/wiki/Organophosphate" title="Organophosphate">phosphate group</a>. The nucleotides are joined to one another in a chain by <a href="/wiki/Covalent_bond" title="Covalent bond">covalent bonds</a> (known as the <a href="/wiki/Phosphodiester_bond" title="Phosphodiester bond">phosphodiester linkage</a>) between the sugar of one nucleotide and the phosphate of the next, resulting in an alternating <a href="/wiki/Backbone_chain" class="mw-redirect" title="Backbone chain">sugar-phosphate backbone</a>. The nitrogenous bases of the two separate polynucleotide strands are bound together, according to <a href="/wiki/Base_pair" title="Base pair">base pairing</a> rules (A with T and C with G), with <a href="/wiki/Hydrogen_bond" title="Hydrogen bond">hydrogen bonds</a> to make double-stranded DNA. The complementary nitrogenous bases are divided into two groups, the single-ringed <a href="/wiki/Pyrimidine" title="Pyrimidine">pyrimidines</a> and the double-ringed <a href="/wiki/Purine" title="Purine">purines</a>. In DNA, the pyrimidines are thymine and cytosine; the purines are adenine and guanine. </p><p>Both strands of double-stranded DNA store the same <a href="/wiki/Central_dogma_of_molecular_biology#Biological_sequence_information" title="Central dogma of molecular biology">biological information</a>. This information is <a href="/wiki/DNA_replication" title="DNA replication">replicated</a> when the two strands separate. A large part of DNA (more than 98% for humans) is <a href="/wiki/Non-coding_DNA" title="Non-coding DNA">non-coding</a>, meaning that these sections do not serve as patterns for <a href="/wiki/Primary_protein_structure" class="mw-redirect" title="Primary protein structure">protein sequences</a>. The two strands of DNA run in opposite directions to each other and are thus <a href="/wiki/Antiparallel_(biochemistry)" title="Antiparallel (biochemistry)">antiparallel</a>. Attached to each sugar is one of four types of nucleobases (or <i>bases</i>). It is the <a href="/wiki/Nucleic_acid_sequence" title="Nucleic acid sequence">sequence</a> of these four nucleobases along the backbone that encodes genetic information. <a href="/wiki/RNA" title="RNA">RNA</a> strands are created using DNA strands as a template in a process called <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcription</a>, where DNA bases are exchanged for their corresponding bases except in the case of thymine (T), for which RNA substitutes <a href="/wiki/Uracil" title="Uracil">uracil</a> (U).<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> Under the <a href="/wiki/Genetic_code" title="Genetic code">genetic code</a>, these RNA strands specify the sequence of <a href="/wiki/Amino_acid" title="Amino acid">amino acids</a> within proteins in a process called <a href="/wiki/Translation_(genetics)" class="mw-redirect" title="Translation (genetics)">translation</a>. </p><p>Within eukaryotic cells, DNA is organized into long structures called <a href="/wiki/Chromosome" title="Chromosome">chromosomes</a>. Before typical <a href="/wiki/Cell_division" title="Cell division">cell division</a>, these chromosomes are duplicated in the process of DNA replication, providing a complete set of chromosomes for each daughter cell. <a href="/wiki/Eukaryote" title="Eukaryote">Eukaryotic organisms</a> (<a href="/wiki/Animal" title="Animal">animals</a>, <a href="/wiki/Plant" title="Plant">plants</a>, <a href="/wiki/Fungus" title="Fungus">fungi</a> and <a href="/wiki/Protist" title="Protist">protists</a>) store most of their DNA inside the <a href="/wiki/Cell_nucleus" title="Cell nucleus">cell nucleus</a> as <a href="/wiki/Nuclear_DNA" title="Nuclear DNA">nuclear DNA</a>, and some in the <a href="/wiki/Mitochondrion" title="Mitochondrion">mitochondria</a> as <a href="/wiki/Mitochondrial_DNA" title="Mitochondrial DNA">mitochondrial DNA</a> or in <a href="/wiki/Chloroplast" title="Chloroplast">chloroplasts</a> as <a href="/wiki/Chloroplast_DNA" title="Chloroplast DNA">chloroplast DNA</a>.<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> In contrast, <a href="/wiki/Prokaryote" title="Prokaryote">prokaryotes</a> (<a href="/wiki/Bacteria" title="Bacteria">bacteria</a> and <a href="/wiki/Archaea" title="Archaea">archaea</a>) store their DNA only in the <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a>, in <a href="/wiki/Circular_chromosome" title="Circular chromosome">circular chromosomes</a>. Within eukaryotic chromosomes, <a href="/wiki/Chromatin" title="Chromatin">chromatin</a> proteins, such as <a href="/wiki/Histone" title="Histone">histones</a>, compact and organize DNA. These compacting structures guide the interactions between DNA and other proteins, helping control which parts of the DNA are transcribed. </p> <style data-mw-deduplicate="TemplateStyles:r886046785">.mw-parser-output .toclimit-2 .toclevel-1 ul,.mw-parser-output .toclimit-3 .toclevel-2 ul,.mw-parser-output .toclimit-4 .toclevel-3 ul,.mw-parser-output .toclimit-5 .toclevel-4 ul,.mw-parser-output .toclimit-6 .toclevel-5 ul,.mw-parser-output .toclimit-7 .toclevel-6 ul{display:none}</style><div class="toclimit-3"><meta property="mw:PageProp/toc" /></div> <div class="mw-heading mw-heading2"><h2 id="Properties">Properties</h2></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:DNA_chemical_structure.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e4/DNA_chemical_structure.svg/300px-DNA_chemical_structure.svg.png" decoding="async" width="300" height="350" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e4/DNA_chemical_structure.svg/450px-DNA_chemical_structure.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e4/DNA_chemical_structure.svg/600px-DNA_chemical_structure.svg.png 2x" data-file-width="1500" data-file-height="1750" /></a><figcaption>Chemical structure of DNA; <a href="/wiki/Hydrogen_bond" title="Hydrogen bond">hydrogen bonds</a> shown as dotted lines. Each end of the double helix has an exposed <a href="/wiki/Directionality_(molecular_biology)#5′-end" title="Directionality (molecular biology)">5'</a> phosphate on one strand and an exposed <a href="/wiki/Directionality_(molecular_biology)#3′-end" title="Directionality (molecular biology)">3′</a> hydroxyl group (—OH) on the other.</figcaption></figure> <p>DNA is a long <a href="/wiki/Polymer" title="Polymer">polymer</a> made from repeating units called <a href="/wiki/Nucleotide" title="Nucleotide">nucleotides</a>.<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><sup id="cite_ref-Alberts_7-0" class="reference"><a href="#cite_note-Alberts-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> The structure of DNA is dynamic along its length, being capable of coiling into tight loops and other shapes.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> In all species it is composed of two helical chains, bound to each other by <a href="/wiki/Hydrogen_bonds" class="mw-redirect" title="Hydrogen bonds">hydrogen bonds</a>. Both chains are coiled around the same axis, and have the same <a href="/wiki/Pitch_(screw)" class="mw-redirect" title="Pitch (screw)">pitch</a> of 34 <a href="/wiki/%C3%85ngstr%C3%B6m" class="mw-redirect" title="Ångström">ångströms</a> (3.4 <a href="/wiki/Nanometre" title="Nanometre">nm</a>). The pair of chains have a radius of 10 Å (1.0 nm).<sup id="cite_ref-Watson-1953_9-0" class="reference"><a href="#cite_note-Watson-1953-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> According to another study, when measured in a different solution, the DNA chain measured 22–26 Å (2.2–2.6 nm) wide, and one nucleotide unit measured 3.3 Å (0.33 nm) long.<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> The buoyant density of most DNA is 1.7g/cm<sup>3</sup>.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p><p>DNA does not usually exist as a single strand, but instead as a pair of strands that are held tightly together.<sup id="cite_ref-Watson-1953_9-1" class="reference"><a href="#cite_note-Watson-1953-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-berg_12-0" class="reference"><a href="#cite_note-berg-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> These two long strands coil around each other, in the shape of a <a href="/wiki/Double_helix" class="mw-redirect" title="Double helix">double helix</a>. The nucleotide contains both a segment of the <a href="/wiki/Backbone_chain" class="mw-redirect" title="Backbone chain">backbone</a> of the molecule (which holds the chain together) and a <a href="/wiki/Nucleobase" class="mw-redirect" title="Nucleobase">nucleobase</a> (which interacts with the other DNA strand in the helix). A nucleobase linked to a sugar is called a <a href="/wiki/Nucleoside" title="Nucleoside">nucleoside</a>, and a base linked to a sugar and to one or more phosphate groups is called a <a href="/wiki/Nucleotide" title="Nucleotide">nucleotide</a>. A <a href="/wiki/Biopolymer" title="Biopolymer">biopolymer</a> comprising multiple linked nucleotides (as in DNA) is called a <a href="/wiki/Polynucleotide" title="Polynucleotide">polynucleotide</a>.<sup id="cite_ref-IUPAC_13-0" class="reference"><a href="#cite_note-IUPAC-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p>The backbone of the DNA strand is made from alternating <a href="/wiki/Phosphate" title="Phosphate">phosphate</a> and <a href="/wiki/Carbohydrate" title="Carbohydrate">sugar</a> groups.<sup id="cite_ref-Ghosh_14-0" class="reference"><a href="#cite_note-Ghosh-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> The sugar in DNA is <a href="/wiki/Deoxyribose" title="Deoxyribose">2-deoxyribose</a>, which is a <a href="/wiki/Pentose" title="Pentose">pentose</a> (five-<a href="/wiki/Carbon" title="Carbon">carbon</a>) sugar. The sugars are joined by phosphate groups that form <a href="/wiki/Phosphodiester_bond" title="Phosphodiester bond">phosphodiester bonds</a> between the third and fifth carbon <a href="/wiki/Atom" title="Atom">atoms</a> of adjacent sugar rings. These are known as the <a href="/wiki/Directionality_(molecular_biology)#3′-end" title="Directionality (molecular biology)">3′-end</a> (three prime end), and <a href="/wiki/Directionality_(molecular_biology)#5′-end" title="Directionality (molecular biology)">5′-end</a> (five prime end) carbons, the prime symbol being used to distinguish these carbon atoms from those of the base to which the deoxyribose forms a <a href="/wiki/Glycosidic_bond" title="Glycosidic bond">glycosidic bond</a>.<sup id="cite_ref-berg_12-1" class="reference"><a href="#cite_note-berg-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p><p>Therefore, any DNA strand normally has one end at which there is a phosphate group attached to the 5′ carbon of a ribose (the 5′ phosphoryl) and another end at which there is a free hydroxyl group attached to the 3′ carbon of a ribose (the 3′ hydroxyl). The orientation of the 3′ and 5′ carbons along the sugar-phosphate backbone confers <a href="/wiki/Directionality_(molecular_biology)" title="Directionality (molecular biology)">directionality</a> (sometimes called polarity) to each DNA strand. In a <a href="/wiki/Nucleic_acid_double_helix" title="Nucleic acid double helix">nucleic acid double helix</a>, the direction of the nucleotides in one strand is opposite to their direction in the other strand: the strands are <a href="/wiki/Antiparallel_(biochemistry)" title="Antiparallel (biochemistry)">antiparallel</a>. The asymmetric ends of DNA strands are said to have a directionality of five prime end (5′ ), and three prime end (3′), with the 5′ end having a terminal phosphate group and the 3′ end a terminal hydroxyl group. One major difference between DNA and <a href="/wiki/RNA" title="RNA">RNA</a> is the sugar, with the 2-deoxyribose in DNA being replaced by the related pentose sugar <a href="/wiki/Ribose" title="Ribose">ribose</a> in RNA.<sup id="cite_ref-berg_12-2" class="reference"><a href="#cite_note-berg-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:DNA_animation.gif" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0c/DNA_animation.gif/170px-DNA_animation.gif" decoding="async" width="170" height="294" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/0/0c/DNA_animation.gif 1.5x" data-file-width="181" data-file-height="313" /></a><figcaption>A section of DNA. The bases lie horizontally between the two spiraling strands<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> (<a href="/wiki/File:DNA_orbit_animated.gif" title="File:DNA orbit animated.gif">animated version</a>).</figcaption></figure> <p>The DNA double helix is stabilized primarily by two forces: <a href="/wiki/Hydrogen_bond" title="Hydrogen bond">hydrogen bonds</a> between nucleotides and <a href="/wiki/Stacking_(chemistry)" title="Stacking (chemistry)">base-stacking</a> interactions among <a href="/wiki/Aromatic" class="mw-redirect" title="Aromatic">aromatic</a> nucleobases.<sup id="cite_ref-Yakovchuk2006_16-0" class="reference"><a href="#cite_note-Yakovchuk2006-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> The four bases found in DNA are <a href="/wiki/Adenine" title="Adenine">adenine</a> (<style data-mw-deduplicate="TemplateStyles:r886049734">.mw-parser-output .monospaced{font-family:monospace,monospace}</style><span class="monospaced">A</span>), <a href="/wiki/Cytosine" title="Cytosine">cytosine</a> (<link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">C</span>), <a href="/wiki/Guanine" title="Guanine">guanine</a> (<link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">G</span>) and <a href="/wiki/Thymine" title="Thymine">thymine</a> (<link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">T</span>). These four bases are attached to the sugar-phosphate to form the complete nucleotide, as shown for <a href="/wiki/Adenosine_monophosphate" title="Adenosine monophosphate">adenosine monophosphate</a>. Adenine pairs with thymine and guanine pairs with cytosine, forming <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">A-T</span> and <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">G-C</span> <a href="/wiki/Base_pair" title="Base pair">base pairs</a>.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Nucleobase_classification">Nucleobase classification</h3></div> <p>The nucleobases are classified into two types: the <a href="/wiki/Purine" title="Purine">purines</a>, <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">A</span> and <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">G</span>, which are fused five- and six-membered <a href="/wiki/Heterocyclic_compound" title="Heterocyclic compound">heterocyclic compounds</a>, and the <a href="/wiki/Pyrimidine" title="Pyrimidine">pyrimidines</a>, the six-membered rings <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">C</span> and <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">T</span>.<sup id="cite_ref-berg_12-3" class="reference"><a href="#cite_note-berg-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> A fifth pyrimidine nucleobase, <a href="/wiki/Uracil" title="Uracil">uracil</a> (<link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">U</span>), usually takes the place of thymine in RNA and differs from thymine by lacking a <a href="/wiki/Methyl_group" title="Methyl group">methyl group</a> on its ring. In addition to RNA and DNA, many artificial <a href="/wiki/Nucleic_acid_analogue" title="Nucleic acid analogue">nucleic acid analogues</a> have been created to study the properties of nucleic acids, or for use in biotechnology.<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Non-canonical_bases">Non-canonical bases</h3></div> <p>Modified bases occur in DNA. The first of these recognized was <a href="/wiki/5-methylcytosine" class="mw-redirect" title="5-methylcytosine">5-methylcytosine</a>, which was found in the <a href="/wiki/Genome" title="Genome">genome</a> of <i><a href="/wiki/Mycobacterium_tuberculosis" title="Mycobacterium tuberculosis">Mycobacterium tuberculosis</a></i> in 1925.<sup id="cite_ref-Johnson1925_20-0" class="reference"><a href="#cite_note-Johnson1925-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> The reason for the presence of these noncanonical bases in bacterial viruses (<a href="/wiki/Bacteriophage" title="Bacteriophage">bacteriophages</a>) is to avoid the <a href="/wiki/Restriction_enzyme" title="Restriction enzyme">restriction enzymes</a> present in bacteria. This enzyme system acts at least in part as a molecular immune system protecting bacteria from infection by viruses.<sup id="cite_ref-pmid27319741_21-0" class="reference"><a href="#cite_note-pmid27319741-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> Modifications of the bases cytosine and adenine, the more common and modified DNA bases, play vital roles in the <a href="/wiki/Epigenetics" title="Epigenetics">epigenetic</a> control of gene expression in plants and animals.<sup id="cite_ref-pmid30619465_22-0" class="reference"><a href="#cite_note-pmid30619465-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p><p>A number of noncanonical bases are known to occur in DNA.<sup id="cite_ref-pmid28941008_23-0" class="reference"><a href="#cite_note-pmid28941008-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> Most of these are modifications of the canonical bases plus uracil. </p> <ul><li>Modified <b>Adenine</b> <ul><li>N6-carbamoyl-methyladenine</li> <li>N6-methyadenine</li></ul></li> <li>Modified <b>Guanine</b> <ul><li>7-Deazaguanine</li> <li>7-Methylguanine</li></ul></li> <li>Modified <b>Cytosine</b> <ul><li>N4-Methylcytosine</li> <li>5-Carboxylcytosine</li> <li>5-Formylcytosine</li> <li>5-Glycosylhydroxymethylcytosine</li> <li>5-Hydroxycytosine</li> <li>5-Methylcytosine</li></ul></li> <li>Modified <b>Thymidine</b> <ul><li>α-Glutamythymidine</li> <li>α-Putrescinylthymine</li></ul></li> <li><b>Uracil</b> and modifications <ul><li>Base J</li> <li>Uracil</li> <li>5-Dihydroxypentauracil</li> <li>5-Hydroxymethyldeoxyuracil</li></ul></li> <li>Others <ul><li>Deoxyarchaeosine</li> <li>2,6-Diaminopurine (2-Aminoadenine)</li></ul></li></ul> <div class="mw-heading mw-heading3"><h3 id="Grooves">Grooves</h3></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:DNA-ligand-by-Abalone.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8b/DNA-ligand-by-Abalone.png/220px-DNA-ligand-by-Abalone.png" decoding="async" width="220" height="148" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8b/DNA-ligand-by-Abalone.png/330px-DNA-ligand-by-Abalone.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8b/DNA-ligand-by-Abalone.png/440px-DNA-ligand-by-Abalone.png 2x" data-file-width="1057" data-file-height="713" /></a><figcaption>DNA major and minor grooves. The latter is a binding site for the <a href="/wiki/Hoechst_stain" title="Hoechst stain">Hoechst stain</a> dye 33258.</figcaption></figure> <p>Twin helical strands form the DNA backbone. Another double helix may be found tracing the spaces, or grooves, between the strands. These voids are adjacent to the base pairs and may provide a <a href="/wiki/Binding_site" title="Binding site">binding site</a>. As the strands are not symmetrically located with respect to each other, the grooves are unequally sized. The major groove is 22 ångströms (2.2 nm) wide, while the minor groove is 12 Å (1.2 nm) in width.<sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> Due to the larger width of the major groove, the edges of the bases are more accessible in the major groove than in the minor groove. As a result, proteins such as <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factors</a> that can bind to specific sequences in double-stranded DNA usually make contact with the sides of the bases exposed in the major groove.<sup id="cite_ref-Pabo1984_25-0" class="reference"><a href="#cite_note-Pabo1984-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> This situation varies in unusual conformations of DNA within the cell <i>(see below)</i>, but the major and minor grooves are always named to reflect the differences in width that would be seen if the DNA was twisted back into the ordinary <a href="/wiki/B-DNA" class="mw-redirect" title="B-DNA">B form</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Base_pairing">Base pairing</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Base_pair" title="Base pair">Base pair</a></div> <div class="thumb tright" style="background:#f9f9f9; border:1px solid #ccc; margin:0.5em;"> <table border="0" cellpadding="2" cellspacing="0" style="width:230px; font-size:85%; border:1px solid #ccc; margin:0.3em;"> <tbody><tr> <td><span typeof="mw:File"><a href="/wiki/File:Base_pair_GC.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/64/Base_pair_GC.svg/282px-Base_pair_GC.svg.png" decoding="async" width="282" height="168" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/64/Base_pair_GC.svg/423px-Base_pair_GC.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/64/Base_pair_GC.svg/564px-Base_pair_GC.svg.png 2x" data-file-width="620" data-file-height="369" /></a></span> </td></tr></tbody></table> <table border="0" cellpadding="2" cellspacing="0" style="width:230px; font-size:85%; border:1px solid #ccc; margin:0.3em;"> <tbody><tr> <td><span typeof="mw:File"><a href="/wiki/File:Base_pair_AT.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/db/Base_pair_AT.svg/282px-Base_pair_AT.svg.png" decoding="async" width="282" height="160" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/db/Base_pair_AT.svg/423px-Base_pair_AT.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/db/Base_pair_AT.svg/564px-Base_pair_AT.svg.png 2x" data-file-width="620" data-file-height="352" /></a></span> </td></tr></tbody></table> <div style="border: none; width:282px;"><div class="thumbcaption">Top, a <b><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">GC</span></b> base pair with three <a href="/wiki/Hydrogen_bond" title="Hydrogen bond">hydrogen bonds</a>. Bottom, an <b><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">AT</span></b> base pair with two hydrogen bonds. Non-covalent hydrogen bonds between the pairs are shown as dashed lines.</div></div></div> <p>In a DNA double helix, each type of nucleobase on one strand bonds with just one type of nucleobase on the other strand. This is called <a href="/wiki/Complementarity_(molecular_biology)" title="Complementarity (molecular biology)">complementary</a> <a href="/wiki/Base_pair" title="Base pair">base pairing</a>. Purines form <a href="/wiki/Hydrogen_bond" title="Hydrogen bond">hydrogen bonds</a> to pyrimidines, with adenine bonding only to thymine in two hydrogen bonds, and cytosine bonding only to guanine in three hydrogen bonds. This arrangement of two nucleotides binding together across the double helix (from six-carbon ring to six-carbon ring) is called a Watson-Crick base pair. DNA with high <a href="/wiki/GC-content" title="GC-content">GC-content</a> is more stable than DNA with low <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">GC</span>-content. A <a href="/wiki/Hoogsteen_base_pair" title="Hoogsteen base pair">Hoogsteen base pair</a> (hydrogen bonding the 6-carbon ring to the 5-carbon ring) is a rare variation of base-pairing.<sup id="cite_ref-pmid23818176_26-0" class="reference"><a href="#cite_note-pmid23818176-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> As hydrogen bonds are not <a href="/wiki/Covalent_bond" title="Covalent bond">covalent</a>, they can be broken and rejoined relatively easily. The two strands of DNA in a double helix can thus be pulled apart like a zipper, either by a mechanical force or high <a href="/wiki/Temperature" title="Temperature">temperature</a>.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> As a result of this base pair complementarity, all the information in the double-stranded sequence of a DNA helix is duplicated on each strand, which is vital in DNA replication. This reversible and specific interaction between complementary base pairs is critical for all the functions of DNA in organisms.<sup id="cite_ref-Alberts_7-1" class="reference"><a href="#cite_note-Alberts-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p><p><span class="anchor" id="ssDNA"></span> </p> <div class="mw-heading mw-heading4"><h4 id="ssDNA_vs._dsDNA">ssDNA vs. dsDNA</h4></div> <p>Most DNA molecules are actually two polymer strands, bound together in a helical fashion by noncovalent bonds; this double-stranded (dsDNA) structure is maintained largely by the intrastrand base stacking interactions, which are strongest for <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">G,C</span> stacks. The two strands can come apart—a process known as melting—to form two single-stranded DNA (ssDNA) molecules. Melting occurs at high temperatures, low salt and high <a href="/wiki/PH" title="PH">pH</a> (low pH also melts DNA, but since DNA is unstable due to acid depurination, low pH is rarely used). </p><p>The stability of the dsDNA form depends not only on the <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">GC</span>-content (% <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">G,C</span> basepairs) but also on sequence (since stacking is sequence specific) and also length (longer molecules are more stable). The stability can be measured in various ways; a common way is the <a href="/wiki/DNA_melting" class="mw-redirect" title="DNA melting">melting temperature</a> (also called <i>T<sub>m</sub></i> value), which is the temperature at which 50% of the double-strand molecules are converted to single-strand molecules; melting temperature is dependent on ionic strength and the concentration of DNA. As a result, it is both the percentage of <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">GC</span> base pairs and the overall length of a DNA double helix that determines the strength of the association between the two strands of DNA. Long DNA helices with a high <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">GC</span>-content have more strongly interacting strands, while short helices with high <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">AT</span> content have more weakly interacting strands.<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> In biology, parts of the DNA double helix that need to separate easily, such as the <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">TATAAT</span> <a href="/wiki/Pribnow_box" title="Pribnow box">Pribnow box</a> in some <a href="/wiki/Promoter_(biology)" class="mw-redirect" title="Promoter (biology)">promoters</a>, tend to have a high <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r886049734"><span class="monospaced">AT</span> content, making the strands easier to pull apart.<sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> </p><p>In the laboratory, the strength of this interaction can be measured by finding the melting temperature <i>T<sub>m</sub></i> necessary to break half of the hydrogen bonds. When all the base pairs in a DNA double helix melt, the strands separate and exist in solution as two entirely independent molecules. These single-stranded DNA molecules have no single common shape, but some conformations are more stable than others.<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> </p> <div class="mw-heading mw-heading3"><h3 id="Amount">Amount</h3></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Human_karyotype_with_bands_and_sub-bands.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b1/Human_karyotype_with_bands_and_sub-bands.png/220px-Human_karyotype_with_bands_and_sub-bands.png" decoding="async" width="220" height="364" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b1/Human_karyotype_with_bands_and_sub-bands.png/330px-Human_karyotype_with_bands_and_sub-bands.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b1/Human_karyotype_with_bands_and_sub-bands.png/440px-Human_karyotype_with_bands_and_sub-bands.png 2x" data-file-width="9684" data-file-height="16008" /></a><figcaption>Schematic <a href="/wiki/Karyotype" title="Karyotype">karyogram</a> of a human. It shows 22 <a href="/wiki/Homologous_chromosome" title="Homologous chromosome">homologous chromosomes</a>, both the female (XX) and male (XY) versions of the <a href="/wiki/Sex_chromosome" title="Sex chromosome">sex chromosome</a> (bottom right), as well as the <a href="/wiki/Human_mitochondrial_genetics" title="Human mitochondrial genetics">mitochondrial genome</a> (to scale at bottom left). The blue scale to the left of each chromosome pair (and the mitochondrial genome) shows its length in terms of millions of DNA <a href="/wiki/Base_pair" title="Base pair">base pairs</a>.<link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Karyotype" title="Karyotype">Karyotype</a></div></figcaption></figure> <p>In humans, the total female <a href="/wiki/Diploid" class="mw-redirect" title="Diploid">diploid</a> <a href="/wiki/Nuclear_genome" class="mw-redirect" title="Nuclear genome">nuclear genome</a> per cell extends for 6.37 Gigabase pairs (Gbp), is 208.23 cm long and weighs 6.51 picograms (pg).<sup id="cite_ref-pmid30813969_31-0" class="reference"><a href="#cite_note-pmid30813969-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> Male values are 6.27 Gbp, 205.00 cm, 6.41 pg.<sup id="cite_ref-pmid30813969_31-1" class="reference"><a href="#cite_note-pmid30813969-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> Each DNA polymer can contain hundreds of millions of nucleotides, such as in <a href="/wiki/Chromosome_1" title="Chromosome 1">chromosome 1</a>. Chromosome 1 is the largest human <a href="/wiki/Chromosome" title="Chromosome">chromosome</a> with approximately 220 million <a href="/wiki/Base_pair" title="Base pair">base pairs</a>, and would be <span class="nowrap"><span data-sort-value="6998850000000000000♠"></span>85 mm</span> long if straightened.<sup id="cite_ref-Gregory_2006_32-0" class="reference"><a href="#cite_note-Gregory_2006-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p><p>In <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotes</a>, in addition to <a href="/wiki/Nuclear_DNA" title="Nuclear DNA">nuclear DNA</a>, there is also <a href="/wiki/Mitochondrial_DNA" title="Mitochondrial DNA">mitochondrial DNA</a> (mtDNA) which encodes certain proteins used by the mitochondria. The mtDNA is usually relatively small in comparison to the nuclear DNA. For example, the <a href="/wiki/Human_mitochondrial_genetics" title="Human mitochondrial genetics">human mitochondrial DNA</a> forms closed circular molecules, each of which contains 16,569<sup id="cite_ref-Anderson_1981_33-0" class="reference"><a href="#cite_note-Anderson_1981-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> DNA base pairs,<sup id="cite_ref-Satoh1991_35-0" class="reference"><a href="#cite_note-Satoh1991-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> with each such molecule normally containing a full set of the mitochondrial genes. Each human mitochondrion contains, on average, approximately 5 such mtDNA molecules.<sup id="cite_ref-Satoh1991_35-1" class="reference"><a href="#cite_note-Satoh1991-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> Each human <a href="/wiki/Cell_(biology)" title="Cell (biology)">cell</a> contains approximately 100 mitochondria, giving a total number of mtDNA molecules per human cell of approximately 500.<sup id="cite_ref-Satoh1991_35-2" class="reference"><a href="#cite_note-Satoh1991-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> However, the amount of mitochondria per cell also varies by cell type, and an <a href="/wiki/Egg_cell" title="Egg cell">egg cell</a> can contain 100,000 mitochondria, corresponding to up to 1,500,000 copies of the mitochondrial genome (constituting up to 90% of the DNA of the cell).<sup id="cite_ref-pmid28721182_36-0" class="reference"><a href="#cite_note-pmid28721182-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Sense_and_antisense">Sense and antisense</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Sense_(molecular_biology)" title="Sense (molecular biology)">Sense (molecular biology)</a></div> <p>A <a href="/wiki/DNA_sequencing" title="DNA sequencing">DNA sequence</a> is called a "sense" sequence if it is the same as that of a <a href="/wiki/Messenger_RNA" title="Messenger RNA">messenger RNA</a> copy that is translated into protein.<sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> The sequence on the opposite strand is called the "antisense" sequence. Both sense and antisense sequences can exist on different parts of the same strand of DNA (i.e. both strands can contain both sense and antisense sequences). In both prokaryotes and eukaryotes, antisense RNA sequences are produced, but the functions of these RNAs are not entirely clear.<sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> One proposal is that antisense RNAs are involved in regulating <a href="/wiki/Gene_expression" title="Gene expression">gene expression</a> through RNA-RNA base pairing.<sup id="cite_ref-39" class="reference"><a href="#cite_note-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> </p><p>A few DNA sequences in prokaryotes and eukaryotes, and more in <a href="/wiki/Plasmid" title="Plasmid">plasmids</a> and <a href="/wiki/Virus" title="Virus">viruses</a>, blur the distinction between sense and antisense strands by having <a href="/wiki/Overlapping_gene" title="Overlapping gene">overlapping genes</a>.<sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> In these cases, some DNA sequences do double duty, encoding one protein when read along one strand, and a second protein when read in the opposite direction along the other strand. In <a href="/wiki/Bacteria" title="Bacteria">bacteria</a>, this overlap may be involved in the regulation of gene transcription,<sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> while in viruses, overlapping genes increase the amount of information that can be encoded within the small viral genome.<sup id="cite_ref-42" class="reference"><a href="#cite_note-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Supercoiling">Supercoiling</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/DNA_supercoil" title="DNA supercoil">DNA supercoil</a></div> <p>DNA can be twisted like a rope in a process called <a href="/wiki/DNA_supercoil" title="DNA supercoil">DNA supercoiling</a>. With DNA in its "relaxed" state, a strand usually circles the axis of the double helix once every 10.4 base pairs, but if the DNA is twisted the strands become more tightly or more loosely wound.<sup id="cite_ref-43" class="reference"><a href="#cite_note-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> If the DNA is twisted in the direction of the helix, this is positive supercoiling, and the bases are held more tightly together. If they are twisted in the opposite direction, this is negative supercoiling, and the bases come apart more easily. In nature, most DNA has slight negative supercoiling that is introduced by <a href="/wiki/Enzyme" title="Enzyme">enzymes</a> called <a href="/wiki/Topoisomerase" title="Topoisomerase">topoisomerases</a>.<sup id="cite_ref-Champoux_44-0" class="reference"><a href="#cite_note-Champoux-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> These enzymes are also needed to relieve the twisting stresses introduced into DNA strands during processes such as <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcription</a> and <a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a>.<sup id="cite_ref-Wang_45-0" class="reference"><a href="#cite_note-Wang-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Alternative_DNA_structures">Alternative DNA structures</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Molecular_Structure_of_Nucleic_Acids:_A_Structure_for_Deoxyribose_Nucleic_Acid" title="Molecular Structure of Nucleic Acids: A Structure for Deoxyribose Nucleic Acid">Molecular Structure of Nucleic Acids: A Structure for Deoxyribose Nucleic Acid</a>, <a href="/wiki/Molecular_models_of_DNA" title="Molecular models of DNA">Molecular models of DNA</a>, and <a href="/wiki/DNA_structure" class="mw-redirect" title="DNA structure">DNA structure</a></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Dnaconformations.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/00/Dnaconformations.png/220px-Dnaconformations.png" decoding="async" width="220" height="255" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/00/Dnaconformations.png/330px-Dnaconformations.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/00/Dnaconformations.png/440px-Dnaconformations.png 2x" data-file-width="1131" data-file-height="1310" /></a><figcaption>From left to right, the structures of <a href="/wiki/A-DNA" title="A-DNA">A</a>, <a href="/wiki/B-DNA" class="mw-redirect" title="B-DNA">B</a> and <a href="/wiki/Z-DNA" title="Z-DNA">Z-DNA</a></figcaption></figure> <p>DNA exists in many possible <a href="/wiki/Conformational_isomerism" title="Conformational isomerism">conformations</a> that include <a href="/wiki/A-DNA" title="A-DNA">A-DNA</a>, <a href="/wiki/B-DNA" class="mw-redirect" title="B-DNA">B-DNA</a>, and <a href="/wiki/Z-DNA" title="Z-DNA">Z-DNA</a> forms, although only B-DNA and Z-DNA have been directly observed in functional organisms.<sup id="cite_ref-Ghosh_14-1" class="reference"><a href="#cite_note-Ghosh-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> The conformation that DNA adopts depends on the hydration level, DNA sequence, the amount and direction of supercoiling, chemical modifications of the bases, the type and concentration of metal <a href="/wiki/Ion" title="Ion">ions</a>, and the presence of <a href="/wiki/Polyamine" title="Polyamine">polyamines</a> in solution.<sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> </p><p>The first published reports of A-DNA <a href="/w/index.php?title=X-ray_diffraction_patterns&action=edit&redlink=1" class="new" title="X-ray diffraction patterns (page does not exist)">X-ray diffraction patterns</a>—and also B-DNA—used analyses based on <a href="/wiki/Patterson_function" title="Patterson function">Patterson functions</a> that provided only a limited amount of structural information for oriented fibers of DNA.<sup id="cite_ref-47" class="reference"><a href="#cite_note-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-NatFranGos_48-0" class="reference"><a href="#cite_note-NatFranGos-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> An alternative analysis was proposed by Wilkins <i>et al.</i> in 1953 for the <i><a href="/wiki/In_vivo" title="In vivo">in vivo</a></i> B-DNA X-ray diffraction-scattering patterns of highly hydrated DNA fibers in terms of squares of <a href="/wiki/Bessel_function" title="Bessel function">Bessel functions</a>.<sup id="cite_ref-NatWilk_49-0" class="reference"><a href="#cite_note-NatWilk-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> In the same journal, <a href="/wiki/James_Watson" title="James Watson">James Watson</a> and <a href="/wiki/Francis_Crick" title="Francis Crick">Francis Crick</a> presented their <a href="/wiki/Molecular_models_of_DNA" title="Molecular models of DNA">molecular modeling</a> analysis of the DNA X-ray diffraction patterns to suggest that the structure was a double helix.<sup id="cite_ref-Watson-1953_9-2" class="reference"><a href="#cite_note-Watson-1953-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p><p>Although the <i>B-DNA form</i> is most common under the conditions found in cells,<sup id="cite_ref-50" class="reference"><a href="#cite_note-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> it is not a well-defined conformation but a family of related DNA conformations<sup id="cite_ref-51" class="reference"><a href="#cite_note-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> that occur at the high hydration levels present in cells. Their corresponding X-ray diffraction and scattering patterns are characteristic of molecular <a href="/wiki/Paracrystalline" class="mw-redirect" title="Paracrystalline">paracrystals</a> with a significant degree of disorder.<sup id="cite_ref-52" class="reference"><a href="#cite_note-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-53" class="reference"><a href="#cite_note-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> </p><p>Compared to B-DNA, the A-DNA form is a wider <a href="/wiki/Helix#Properties_and_types" title="Helix">right-handed</a> spiral, with a shallow, wide minor groove and a narrower, deeper major groove. The A form occurs under non-physiological conditions in partly dehydrated samples of DNA, while in the cell it may be produced in hybrid pairings of DNA and RNA strands, and in enzyme-DNA complexes.<sup id="cite_ref-54" class="reference"><a href="#cite_note-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-55" class="reference"><a href="#cite_note-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> Segments of DNA where the bases have been chemically modified by <a href="/wiki/Methylation" title="Methylation">methylation</a> may undergo a larger change in conformation and adopt the <a href="/wiki/Z-DNA" title="Z-DNA">Z form</a>. Here, the strands turn about the helical axis in a left-handed spiral, the opposite of the more common B form.<sup id="cite_ref-56" class="reference"><a href="#cite_note-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> These unusual structures can be recognized by specific Z-DNA binding proteins and may be involved in the regulation of transcription.<sup id="cite_ref-57" class="reference"><a href="#cite_note-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Alternative_DNA_chemistry">Alternative DNA chemistry</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Hypothetical_types_of_biochemistry" title="Hypothetical types of biochemistry">hypothetical types of biochemistry</a></div> <p>For many years, <a href="/wiki/Astrobiology" title="Astrobiology">exobiologists</a> have proposed the existence of a <a href="/wiki/Shadow_biosphere" title="Shadow biosphere">shadow biosphere</a>, a postulated microbial <a href="/wiki/Biosphere" title="Biosphere">biosphere</a> of Earth that uses radically different biochemical and molecular processes than currently known life. One of the proposals was the existence of lifeforms that use <a href="/wiki/Arsenic_DNA" class="mw-redirect" title="Arsenic DNA">arsenic instead of phosphorus in DNA</a>. A report in 2010 of the possibility in the <a href="/wiki/Bacterium" class="mw-redirect" title="Bacterium">bacterium</a> <a href="/wiki/GFAJ-1" title="GFAJ-1">GFAJ-1</a> was announced,<sup id="cite_ref-arsenic_extremophile_58-0" class="reference"><a href="#cite_note-arsenic_extremophile-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Space_59-0" class="reference"><a href="#cite_note-Space-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> though the research was disputed,<sup id="cite_ref-Space_59-1" class="reference"><a href="#cite_note-Space-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-60" class="reference"><a href="#cite_note-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> and evidence suggests the bacterium actively prevents the incorporation of arsenic into the DNA backbone and other biomolecules.<sup id="cite_ref-Nature_61-0" class="reference"><a href="#cite_note-Nature-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Quadruplex_structures">Quadruplex structures</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/G-quadruplex" title="G-quadruplex">G-quadruplex</a></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Parallel_telomere_quadruple.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/9e/Parallel_telomere_quadruple.png/220px-Parallel_telomere_quadruple.png" decoding="async" width="220" height="201" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/9e/Parallel_telomere_quadruple.png/330px-Parallel_telomere_quadruple.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/9e/Parallel_telomere_quadruple.png/440px-Parallel_telomere_quadruple.png 2x" data-file-width="1316" data-file-height="1200" /></a><figcaption>DNA quadruplex formed by <a href="/wiki/Telomere" title="Telomere">telomere</a> repeats. The looped conformation of the DNA backbone is very different from the typical DNA helix. The green spheres in the center represent potassium ions.<sup id="cite_ref-62" class="reference"><a href="#cite_note-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>At the ends of the linear chromosomes are specialized regions of DNA called <a href="/wiki/Telomere" title="Telomere">telomeres</a>. The main function of these regions is to allow the cell to replicate chromosome ends using the enzyme <a href="/wiki/Telomerase" title="Telomerase">telomerase</a>, as the enzymes that normally replicate DNA cannot copy the extreme 3′ ends of chromosomes.<sup id="cite_ref-Greider_63-0" class="reference"><a href="#cite_note-Greider-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup> These specialized chromosome caps also help protect the DNA ends, and stop the <a href="/wiki/DNA_repair" title="DNA repair">DNA repair</a> systems in the cell from treating them as damage to be corrected.<sup id="cite_ref-Nugent_64-0" class="reference"><a href="#cite_note-Nugent-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> In <a href="/wiki/List_of_distinct_cell_types_in_the_adult_human_body" class="mw-redirect" title="List of distinct cell types in the adult human body">human cells</a>, telomeres are usually lengths of single-stranded DNA containing several thousand repeats of a simple TTAGGG sequence.<sup id="cite_ref-65" class="reference"><a href="#cite_note-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> </p><p>These guanine-rich sequences may stabilize chromosome ends by forming structures of stacked sets of four-base units, rather than the usual base pairs found in other DNA molecules. Here, four guanine bases, known as a <a href="/wiki/Guanine_tetrad" title="Guanine tetrad">guanine tetrad</a>, form a flat plate. These flat four-base units then stack on top of each other to form a stable <a href="/wiki/G-quadruplex" title="G-quadruplex">G-quadruplex</a> structure.<sup id="cite_ref-Burge_66-0" class="reference"><a href="#cite_note-Burge-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> These structures are stabilized by hydrogen bonding between the edges of the bases and <a href="/wiki/Chelation" title="Chelation">chelation</a> of a metal ion in the centre of each four-base unit.<sup id="cite_ref-67" class="reference"><a href="#cite_note-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> Other structures can also be formed, with the central set of four bases coming from either a single strand folded around the bases, or several different parallel strands, each contributing one base to the central structure. </p><p>In addition to these stacked structures, telomeres also form large loop structures called telomere loops, or T-loops. Here, the single-stranded DNA curls around in a long circle stabilized by telomere-binding proteins.<sup id="cite_ref-68" class="reference"><a href="#cite_note-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup> At the very end of the T-loop, the single-stranded telomere DNA is held onto a region of double-stranded DNA by the telomere strand disrupting the double-helical DNA and base pairing to one of the two strands. This <a href="/wiki/Triple-stranded_DNA" title="Triple-stranded DNA">triple-stranded</a> structure is called a displacement loop or <a href="/wiki/D-loop" title="D-loop">D-loop</a>.<sup id="cite_ref-Burge_66-1" class="reference"><a href="#cite_note-Burge-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Branched_DNA">Branched DNA</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Branched_DNA" class="mw-redirect" title="Branched DNA">Branched DNA</a> and <a href="/wiki/DNA_nanotechnology" title="DNA nanotechnology">DNA nanotechnology</a></div> <div class="thumb tright" style="background:#f9f9f9; border:1px solid #ccc; margin:0.5em;"> <table border="0" cellpadding="2" cellspacing="0" style="width:200px; font-size:85%; border:1px solid #ccc; margin:0.3em;"> <tbody><tr> <td><span typeof="mw:File"><a href="/wiki/File:Branch-dna-single.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/54/Branch-dna-single.svg/95px-Branch-dna-single.svg.png" decoding="async" width="95" height="71" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/54/Branch-dna-single.svg/143px-Branch-dna-single.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/54/Branch-dna-single.svg/190px-Branch-dna-single.svg.png 2x" data-file-width="200" data-file-height="150" /></a></span> </td> <td><span typeof="mw:File"><a href="/wiki/File:Branch-DNA-multiple.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/08/Branch-DNA-multiple.svg/95px-Branch-DNA-multiple.svg.png" decoding="async" width="95" height="71" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/08/Branch-DNA-multiple.svg/143px-Branch-DNA-multiple.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/08/Branch-DNA-multiple.svg/190px-Branch-DNA-multiple.svg.png 2x" data-file-width="200" data-file-height="150" /></a></span> </td></tr> <tr> <td align="center">Single branch </td> <td align="center">Multiple branches </td></tr></tbody></table> <div style="border: none; width:200px;font-size: 90%;"><div class="thumbcaption"><a href="/wiki/Branched_DNA" class="mw-redirect" title="Branched DNA">Branched DNA</a> can form networks containing multiple branches.</div></div></div> <p>In DNA, <a href="/wiki/DNA_end#Frayed_ends" class="mw-redirect" title="DNA end">fraying</a> occurs when non-complementary regions exist at the end of an otherwise complementary double-strand of DNA. However, branched DNA can occur if a third strand of DNA is introduced and contains adjoining regions able to hybridize with the frayed regions of the pre-existing double-strand. Although the simplest example of branched DNA involves only three strands of DNA, complexes involving additional strands and multiple branches are also possible.<sup id="cite_ref-69" class="reference"><a href="#cite_note-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup> Branched DNA can be used in <a href="/wiki/Nanotechnology" title="Nanotechnology">nanotechnology</a> to construct geometric shapes, see the section on <a href="#Uses_in_technology">uses in technology</a> below. </p> <div class="mw-heading mw-heading3"><h3 id="Artificial_bases">Artificial bases</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Nucleic_acid_analogue" title="Nucleic acid analogue">Nucleic acid analogue</a></div> <p>Several artificial nucleobases have been synthesized, and successfully incorporated in the eight-base DNA analogue named <a href="/wiki/Hachimoji_DNA" title="Hachimoji DNA">Hachimoji DNA</a>. Dubbed S, B, P, and Z, these artificial bases are capable of bonding with each other in a predictable way (S–B and P–Z), maintain the double helix structure of DNA, and be transcribed to RNA. Their existence could be seen as an indication that there is nothing special about the four natural nucleobases that evolved on Earth.<sup id="cite_ref-70" class="reference"><a href="#cite_note-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-71" class="reference"><a href="#cite_note-71"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup> On the other hand, DNA is tightly related to <a href="/wiki/RNA" title="RNA">RNA</a> which does not only act as a transcript of DNA but also performs as molecular machines many tasks in cells. For this purpose it has to fold into a structure. It has been shown that to allow to create all possible structures at least four bases are required for the corresponding <a href="/wiki/RNA" title="RNA">RNA</a>,<sup id="cite_ref-72" class="reference"><a href="#cite_note-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> while a higher number is also possible but this would be against the natural <a href="/wiki/Principle_of_least_effort" title="Principle of least effort">principle of least effort</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Acidity">Acidity</h3></div> <p>The phosphate groups of DNA give it similar <a href="/wiki/Acid" title="Acid">acidic</a> properties to <a href="/wiki/Phosphoric_acid" title="Phosphoric acid">phosphoric acid</a> and it can be considered as a <a href="/wiki/Acid_strength" title="Acid strength">strong acid</a>. It will be fully ionized at a normal cellular pH, releasing <a href="/wiki/Proton" title="Proton">protons</a> which leave behind negative charges on the phosphate groups. These negative charges protect DNA from breakdown by <a href="/wiki/Hydrolysis" title="Hydrolysis">hydrolysis</a> by repelling <a href="/wiki/Nucleophile" title="Nucleophile">nucleophiles</a> which could hydrolyze it.<sup id="cite_ref-Reusch_73-0" class="reference"><a href="#cite_note-Reusch-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Macroscopic_appearance">Macroscopic appearance</h3></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Estrazione_DNA_(cropped).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a4/Estrazione_DNA_%28cropped%29.jpg/220px-Estrazione_DNA_%28cropped%29.jpg" decoding="async" width="220" height="136" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a4/Estrazione_DNA_%28cropped%29.jpg/330px-Estrazione_DNA_%28cropped%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a4/Estrazione_DNA_%28cropped%29.jpg/440px-Estrazione_DNA_%28cropped%29.jpg 2x" data-file-width="1049" data-file-height="648" /></a><figcaption>Impure DNA extracted from an orange</figcaption></figure> <p>Pure DNA extracted from cells forms white, stringy clumps.<sup id="cite_ref-74" class="reference"><a href="#cite_note-74"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Chemical_modifications_and_altered_DNA_packaging">Chemical modifications and altered DNA packaging</h2></div> <div class="mw-heading mw-heading3"><h3 id="Base_modifications_and_DNA_packaging">Base modifications and DNA packaging</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/DNA_methylation" title="DNA methylation">DNA methylation</a> and <a href="/wiki/Chromatin_remodeling" title="Chromatin remodeling">Chromatin remodeling</a></div> <div class="thumb tright" style="background:#f9f9f9; border:1px solid #ccc; margin:0.5em;"> <table border="0" cellpadding="2" cellspacing="0" style="width:300px; font-size:85%; border:1px solid #ccc; margin:0.3em;"> <tbody><tr> <td><span typeof="mw:File"><a href="/wiki/File:Cytosin.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/dd/Cytosin.svg/75px-Cytosin.svg.png" decoding="async" width="75" height="102" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/dd/Cytosin.svg/113px-Cytosin.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/dd/Cytosin.svg/150px-Cytosin.svg.png 2x" data-file-width="112" data-file-height="153" /></a></span> </td> <td><span typeof="mw:File"><a href="/wiki/File:5-Methylcytosine.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3a/5-Methylcytosine.svg/95px-5-Methylcytosine.svg.png" decoding="async" width="95" height="83" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/3a/5-Methylcytosine.svg/143px-5-Methylcytosine.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/3a/5-Methylcytosine.svg/190px-5-Methylcytosine.svg.png 2x" data-file-width="620" data-file-height="543" /></a></span> </td> <td><span typeof="mw:File"><a href="/wiki/File:Thymin.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/15/Thymin.svg/97px-Thymin.svg.png" decoding="async" width="97" height="83" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/15/Thymin.svg/146px-Thymin.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/15/Thymin.svg/194px-Thymin.svg.png 2x" data-file-width="180" data-file-height="154" /></a></span> </td></tr> <tr> <td align="center"><a href="/wiki/Cytosine" title="Cytosine">cytosine</a> </td> <td align="center"><a href="/wiki/5-Methylcytosine" title="5-Methylcytosine">5-methylcytosine</a> </td> <td align="center"><a href="/wiki/Thymine" title="Thymine">thymine</a> </td></tr></tbody></table> <div style="border: none; width:300px;font-size: 90%;"><div class="thumbcaption">Structure of cytosine with and without the 5-methyl group. <a href="/wiki/Deamination" title="Deamination">Deamination</a> converts 5-methylcytosine into thymine.</div></div></div> <p>The expression of genes is influenced by how the DNA is packaged in chromosomes, in a structure called <a href="/wiki/Chromatin" title="Chromatin">chromatin</a>. Base modifications can be involved in packaging, with regions that have low or no gene expression usually containing high levels of <a href="/wiki/Methylation" title="Methylation">methylation</a> of <a href="/wiki/Cytosine" title="Cytosine">cytosine</a> bases. DNA packaging and its influence on gene expression can also occur by covalent modifications of the <a href="/wiki/Histone" title="Histone">histone</a> protein core around which DNA is wrapped in the chromatin structure or else by remodeling carried out by chromatin remodeling complexes (see <a href="/wiki/Chromatin_remodeling" title="Chromatin remodeling">Chromatin remodeling</a>). There is, further, <a href="/wiki/Crosstalk_(biology)" title="Crosstalk (biology)">crosstalk</a> between DNA methylation and histone modification, so they can coordinately affect chromatin and gene expression.<sup id="cite_ref-75" class="reference"><a href="#cite_note-75"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup> </p><p>For one example, cytosine methylation produces <a href="/wiki/5-Methylcytosine" title="5-Methylcytosine">5-methylcytosine</a>, which is important for <a href="/wiki/X-inactivation" title="X-inactivation">X-inactivation</a> of chromosomes.<sup id="cite_ref-76" class="reference"><a href="#cite_note-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup> The average level of methylation varies between organisms—the worm <i><a href="/wiki/Caenorhabditis_elegans" title="Caenorhabditis elegans">Caenorhabditis elegans</a></i> lacks cytosine methylation, while <a href="/wiki/Vertebrate" title="Vertebrate">vertebrates</a> have higher levels, with up to 1% of their DNA containing 5-methylcytosine.<sup id="cite_ref-77" class="reference"><a href="#cite_note-77"><span class="cite-bracket">[</span>77<span class="cite-bracket">]</span></a></sup> Despite the importance of 5-methylcytosine, it can <a href="/wiki/Deamination" title="Deamination">deaminate</a> to leave a thymine base, so methylated cytosines are particularly prone to <a href="/wiki/Mutation" title="Mutation">mutations</a>.<sup id="cite_ref-78" class="reference"><a href="#cite_note-78"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup> Other base modifications include adenine methylation in bacteria, the presence of <a href="/wiki/5-hydroxymethylcytosine" class="mw-redirect" title="5-hydroxymethylcytosine">5-hydroxymethylcytosine</a> in the <a href="/wiki/Brain" title="Brain">brain</a>,<sup id="cite_ref-79" class="reference"><a href="#cite_note-79"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> and the <a href="/wiki/Glycosylation" title="Glycosylation">glycosylation</a> of uracil to produce the "J-base" in <a href="/wiki/Kinetoplastida" title="Kinetoplastida">kinetoplastids</a>.<sup id="cite_ref-80" class="reference"><a href="#cite_note-80"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-81" class="reference"><a href="#cite_note-81"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Damage">Damage</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/DNA_damage_(naturally_occurring)" title="DNA damage (naturally occurring)">DNA damage (naturally occurring)</a>, <a href="/wiki/Mutation" title="Mutation">Mutation</a>, and <a href="/wiki/DNA_damage_theory_of_aging" title="DNA damage theory of aging">DNA damage theory of aging</a></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Benzopyrene_DNA_adduct_1JDG.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d8/Benzopyrene_DNA_adduct_1JDG.png/220px-Benzopyrene_DNA_adduct_1JDG.png" decoding="async" width="220" height="305" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d8/Benzopyrene_DNA_adduct_1JDG.png/330px-Benzopyrene_DNA_adduct_1JDG.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d8/Benzopyrene_DNA_adduct_1JDG.png/440px-Benzopyrene_DNA_adduct_1JDG.png 2x" data-file-width="1131" data-file-height="1566" /></a><figcaption>A <a href="/wiki/Covalent" class="mw-redirect" title="Covalent">covalent</a> <a href="/wiki/Adduct" title="Adduct">adduct</a> between a <a href="/wiki/Cytochrome_P450,_family_1,_member_A1" class="mw-redirect" title="Cytochrome P450, family 1, member A1">metabolically activated</a> form of <a href="/wiki/Benzo(a)pyrene" title="Benzo(a)pyrene">benzo[<i>a</i>]pyrene</a>, the major <a href="/wiki/Mutagen" title="Mutagen">mutagen</a> in <a href="/wiki/Tobacco_smoking" title="Tobacco smoking">tobacco smoke</a>, and DNA<sup id="cite_ref-82" class="reference"><a href="#cite_note-82"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>DNA can be damaged by many sorts of <a href="/wiki/Mutagen" title="Mutagen">mutagens</a>, which change the <a href="/wiki/DNA_sequencing" title="DNA sequencing">DNA sequence</a>. Mutagens include <a href="/wiki/Oxidizing_agent" title="Oxidizing agent">oxidizing agents</a>, <a href="/wiki/Alkylation" title="Alkylation">alkylating agents</a> and also high-energy <a href="/wiki/Electromagnetic_radiation" title="Electromagnetic radiation">electromagnetic radiation</a> such as <a href="/wiki/Ultraviolet" title="Ultraviolet">ultraviolet</a> light and <a href="/wiki/X-ray" title="X-ray">X-rays</a>. The type of DNA damage produced depends on the type of mutagen. For example, UV light can damage DNA by producing <a href="/wiki/Thymine_dimer" class="mw-redirect" title="Thymine dimer">thymine dimers</a>, which are cross-links between pyrimidine bases.<sup id="cite_ref-83" class="reference"><a href="#cite_note-83"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> On the other hand, oxidants such as <a href="/wiki/Radical_(chemistry)" title="Radical (chemistry)">free radicals</a> or <a href="/wiki/Hydrogen_peroxide" title="Hydrogen peroxide">hydrogen peroxide</a> produce multiple forms of damage, including base modifications, particularly of guanosine, and double-strand breaks.<sup id="cite_ref-84" class="reference"><a href="#cite_note-84"><span class="cite-bracket">[</span>84<span class="cite-bracket">]</span></a></sup> A typical human cell contains about 150,000 bases that have suffered oxidative damage.<sup id="cite_ref-85" class="reference"><a href="#cite_note-85"><span class="cite-bracket">[</span>85<span class="cite-bracket">]</span></a></sup> Of these oxidative lesions, the most dangerous are double-strand breaks, as these are difficult to repair and can produce <a href="/wiki/Point_mutation" title="Point mutation">point mutations</a>, <a href="/wiki/Genetic_insertion" class="mw-redirect" title="Genetic insertion">insertions</a>, <a href="/wiki/Deletion_(genetics)" title="Deletion (genetics)">deletions</a> from the DNA sequence, and <a href="/wiki/Chromosomal_translocation" title="Chromosomal translocation">chromosomal translocations</a>.<sup id="cite_ref-86" class="reference"><a href="#cite_note-86"><span class="cite-bracket">[</span>86<span class="cite-bracket">]</span></a></sup> These mutations can cause <a href="/wiki/Cancer" title="Cancer">cancer</a>. Because of inherent limits in the DNA repair mechanisms, if humans lived long enough, they would all eventually develop cancer.<sup id="cite_ref-Weinberg_87-0" class="reference"><a href="#cite_note-Weinberg-87"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-88" class="reference"><a href="#cite_note-88"><span class="cite-bracket">[</span>88<span class="cite-bracket">]</span></a></sup> DNA damages that are <a href="/wiki/DNA_damage_(naturally_occurring)" title="DNA damage (naturally occurring)">naturally occurring</a>, due to normal cellular processes that produce reactive oxygen species, the hydrolytic activities of cellular water, etc., also occur frequently. Although most of these damages are repaired, in any cell some DNA damage may remain despite the action of repair processes. These remaining DNA damages accumulate with age in mammalian postmitotic tissues. This accumulation appears to be an important underlying cause of aging.<sup id="cite_ref-89" class="reference"><a href="#cite_note-89"><span class="cite-bracket">[</span>89<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-90" class="reference"><a href="#cite_note-90"><span class="cite-bracket">[</span>90<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-91" class="reference"><a href="#cite_note-91"><span class="cite-bracket">[</span>91<span class="cite-bracket">]</span></a></sup> </p><p>Many mutagens fit into the space between two adjacent base pairs, this is called <i><a href="/wiki/Intercalation_(biochemistry)" title="Intercalation (biochemistry)">intercalation</a></i>. Most intercalators are <a href="/wiki/Aromaticity" title="Aromaticity">aromatic</a> and planar molecules; examples include <a href="/wiki/Ethidium_bromide" title="Ethidium bromide">ethidium bromide</a>, <a href="/wiki/Acridine" title="Acridine">acridines</a>, <a href="/wiki/Daunorubicin" title="Daunorubicin">daunomycin</a>, and <a href="/wiki/Doxorubicin" title="Doxorubicin">doxorubicin</a>. For an intercalator to fit between base pairs, the bases must separate, distorting the DNA strands by unwinding of the double helix. This inhibits both transcription and DNA replication, causing toxicity and mutations.<sup id="cite_ref-92" class="reference"><a href="#cite_note-92"><span class="cite-bracket">[</span>92<span class="cite-bracket">]</span></a></sup> As a result, DNA intercalators may be <a href="/wiki/Carcinogen" title="Carcinogen">carcinogens</a>, and in the case of thalidomide, a <a href="/wiki/Teratogen" class="mw-redirect" title="Teratogen">teratogen</a>.<sup id="cite_ref-93" class="reference"><a href="#cite_note-93"><span class="cite-bracket">[</span>93<span class="cite-bracket">]</span></a></sup> Others such as <a href="/wiki/Benzo(a)pyrene" title="Benzo(a)pyrene">benzo[<i>a</i>]pyrene diol epoxide</a> and <a href="/wiki/Aflatoxin" title="Aflatoxin">aflatoxin</a> form DNA adducts that induce errors in replication.<sup id="cite_ref-94" class="reference"><a href="#cite_note-94"><span class="cite-bracket">[</span>94<span class="cite-bracket">]</span></a></sup> Nevertheless, due to their ability to inhibit DNA transcription and replication, other similar toxins are also used in <a href="/wiki/Chemotherapy" title="Chemotherapy">chemotherapy</a> to inhibit rapidly growing <a href="/wiki/Cancer" title="Cancer">cancer</a> cells.<sup id="cite_ref-95" class="reference"><a href="#cite_note-95"><span class="cite-bracket">[</span>95<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Biological_functions">Biological functions</h2></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Eukaryote_DNA-en.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e2/Eukaryote_DNA-en.svg/320px-Eukaryote_DNA-en.svg.png" decoding="async" width="320" height="198" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e2/Eukaryote_DNA-en.svg/480px-Eukaryote_DNA-en.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e2/Eukaryote_DNA-en.svg/640px-Eukaryote_DNA-en.svg.png 2x" data-file-width="1189" data-file-height="734" /></a><figcaption>Location of eukaryote <a href="/wiki/Nuclear_DNA" title="Nuclear DNA">nuclear DNA</a> within the chromosomes</figcaption></figure> <p>DNA usually occurs as linear <a href="/wiki/Chromosome" title="Chromosome">chromosomes</a> in <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotes</a>, and <a href="/wiki/Circular_prokaryote_chromosome" class="mw-redirect" title="Circular prokaryote chromosome">circular chromosomes</a> in <a href="/wiki/Prokaryote" title="Prokaryote">prokaryotes</a>. The set of chromosomes in a cell makes up its <a href="/wiki/Genome" title="Genome">genome</a>; the <a href="/wiki/Human_genome" title="Human genome">human genome</a> has approximately 3 billion base pairs of DNA arranged into 46 chromosomes.<sup id="cite_ref-Venter_2001_96-0" class="reference"><a href="#cite_note-Venter_2001-96"><span class="cite-bracket">[</span>96<span class="cite-bracket">]</span></a></sup> The information carried by DNA is held in the <a href="/wiki/DNA_sequence" class="mw-redirect" title="DNA sequence">sequence</a> of pieces of DNA called <a href="/wiki/Gene" title="Gene">genes</a>. <a href="/wiki/Transmission_(genetics)" class="mw-redirect" title="Transmission (genetics)">Transmission</a> of genetic information in genes is achieved via complementary base pairing. For example, in transcription, when a cell uses the information in a gene, the DNA sequence is copied into a complementary RNA sequence through the attraction between the DNA and the correct RNA nucleotides. Usually, this RNA copy is then used to make a matching <a href="/wiki/Peptide_sequence" class="mw-redirect" title="Peptide sequence">protein sequence</a> in a process called <a href="/wiki/Translation_(biology)" title="Translation (biology)">translation</a>, which depends on the same interaction between RNA nucleotides. In an alternative fashion, a cell may copy its genetic information in a process called <a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a>. The details of these functions are covered in other articles; here the focus is on the interactions between DNA and other molecules that mediate the function of the genome. </p> <div class="mw-heading mw-heading3"><h3 id="Genes_and_genomes">Genes and genomes</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Cell_nucleus" title="Cell nucleus">Cell nucleus</a>, <a href="/wiki/Chromatin" title="Chromatin">Chromatin</a>, <a href="/wiki/Chromosome" title="Chromosome">Chromosome</a>, <a href="/wiki/Gene" title="Gene">Gene</a>, and <a href="/wiki/Noncoding_DNA" class="mw-redirect" title="Noncoding DNA">Noncoding DNA</a></div> <p>Genomic DNA is tightly and orderly packed in the process called <a href="/wiki/DNA_condensation" title="DNA condensation">DNA condensation</a>, to fit the small available volumes of the cell. In eukaryotes, DNA is located in the <a href="/wiki/Cell_nucleus" title="Cell nucleus">cell nucleus</a>, with small amounts in <a href="/wiki/Mitochondrion" title="Mitochondrion">mitochondria</a> and <a href="/wiki/Chloroplast" title="Chloroplast">chloroplasts</a>. In prokaryotes, the DNA is held within an irregularly shaped body in the cytoplasm called the <a href="/wiki/Nucleoid" title="Nucleoid">nucleoid</a>.<sup id="cite_ref-97" class="reference"><a href="#cite_note-97"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup> The genetic information in a genome is held within genes, and the complete set of this information in an organism is called its <a href="/wiki/Genotype" title="Genotype">genotype</a>. A gene is a unit of <a href="/wiki/Heredity" title="Heredity">heredity</a> and is a region of DNA that influences a particular characteristic in an organism. Genes contain an <a href="/wiki/Open_reading_frame" title="Open reading frame">open reading frame</a> that can be transcribed, and <a href="/wiki/Regulatory_sequence" title="Regulatory sequence">regulatory sequences</a> such as <a href="/wiki/Promoter_(biology)" class="mw-redirect" title="Promoter (biology)">promoters</a> and <a href="/wiki/Enhancer_(genetics)" title="Enhancer (genetics)">enhancers</a>, which control transcription of the open reading frame. </p><p>In many <a href="/wiki/Species" title="Species">species</a>, only a small fraction of the total sequence of the <a href="/wiki/Genome" title="Genome">genome</a> encodes protein. For example, only about 1.5% of the human genome consists of protein-coding <a href="/wiki/Exon" title="Exon">exons</a>, with over 50% of human DNA consisting of non-coding <a href="/wiki/Repeated_sequence_(DNA)" title="Repeated sequence (DNA)">repetitive sequences</a>.<sup id="cite_ref-98" class="reference"><a href="#cite_note-98"><span class="cite-bracket">[</span>98<span class="cite-bracket">]</span></a></sup> The reasons for the presence of so much <a href="/wiki/Noncoding_DNA" class="mw-redirect" title="Noncoding DNA">noncoding DNA</a> in eukaryotic genomes and the extraordinary differences in <a href="/wiki/Genome_size" title="Genome size">genome size</a>, or <i><a href="/wiki/C-value" title="C-value">C-value</a></i>, among species, represent a long-standing puzzle known as the "<a href="/wiki/C-value_enigma" class="mw-redirect" title="C-value enigma">C-value enigma</a>".<sup id="cite_ref-99" class="reference"><a href="#cite_note-99"><span class="cite-bracket">[</span>99<span class="cite-bracket">]</span></a></sup> However, some DNA sequences that do not code protein may still encode functional <a href="/wiki/Non-coding_RNA" title="Non-coding RNA">non-coding RNA</a> molecules, which are involved in the <a href="/wiki/Regulation_of_gene_expression" title="Regulation of gene expression">regulation of gene expression</a>.<sup id="cite_ref-Birney_2007_100-0" class="reference"><a href="#cite_note-Birney_2007-100"><span class="cite-bracket">[</span>100<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:T7_RNA_polymerase.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e3/T7_RNA_polymerase.jpg/220px-T7_RNA_polymerase.jpg" decoding="async" width="220" height="170" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e3/T7_RNA_polymerase.jpg/330px-T7_RNA_polymerase.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e3/T7_RNA_polymerase.jpg/440px-T7_RNA_polymerase.jpg 2x" data-file-width="1061" data-file-height="819" /></a><figcaption><a href="/wiki/T7_RNA_polymerase" title="T7 RNA polymerase">T7 RNA polymerase</a> (blue) producing an <a href="/wiki/Messenger_RNA" title="Messenger RNA">mRNA</a> (green) from a DNA template (orange)<sup id="cite_ref-101" class="reference"><a href="#cite_note-101"><span class="cite-bracket">[</span>101<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>Some noncoding DNA sequences play structural roles in chromosomes. <a href="/wiki/Telomere" title="Telomere">Telomeres</a> and <a href="/wiki/Centromere" title="Centromere">centromeres</a> typically contain few genes but are important for the function and stability of chromosomes.<sup id="cite_ref-Nugent_64-1" class="reference"><a href="#cite_note-Nugent-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-102" class="reference"><a href="#cite_note-102"><span class="cite-bracket">[</span>102<span class="cite-bracket">]</span></a></sup> An abundant form of noncoding DNA in humans are <a href="/wiki/Pseudogene" title="Pseudogene">pseudogenes</a>, which are copies of genes that have been disabled by mutation.<sup id="cite_ref-103" class="reference"><a href="#cite_note-103"><span class="cite-bracket">[</span>103<span class="cite-bracket">]</span></a></sup> These sequences are usually just molecular <a href="/wiki/Fossil" title="Fossil">fossils</a>, although they can occasionally serve as raw <a href="/wiki/Genome" title="Genome">genetic material</a> for the creation of new genes through the process of <a href="/wiki/Gene_duplication" title="Gene duplication">gene duplication</a> and <a href="/wiki/Divergent_evolution" title="Divergent evolution">divergence</a>.<sup id="cite_ref-104" class="reference"><a href="#cite_note-104"><span class="cite-bracket">[</span>104<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Transcription_and_translation">Transcription and translation</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Genetic_code" title="Genetic code">Genetic code</a>, <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">Transcription (genetics)</a>, and <a href="/wiki/Protein_biosynthesis" title="Protein biosynthesis">Protein biosynthesis</a></div> <p>A gene is a sequence of DNA that contains genetic information and can influence the <a href="/wiki/Phenotype" title="Phenotype">phenotype</a> of an organism. Within a gene, the sequence of bases along a DNA strand defines a <a href="/wiki/Messenger_RNA" title="Messenger RNA">messenger RNA</a> sequence, which then defines one or more protein sequences. The relationship between the nucleotide sequences of genes and the <a href="/wiki/Amino_acid" title="Amino acid">amino-acid</a> sequences of proteins is determined by the rules of <a href="/wiki/Translation_(biology)" title="Translation (biology)">translation</a>, known collectively as the <a href="/wiki/Genetic_code" title="Genetic code">genetic code</a>. The genetic code consists of three-letter 'words' called <i>codons</i> formed from a sequence of three nucleotides (e.g. ACT, CAG, TTT). </p><p>In transcription, the codons of a gene are copied into messenger RNA by <a href="/wiki/RNA_polymerase" title="RNA polymerase">RNA polymerase</a>. This RNA copy is then decoded by a <a href="/wiki/Ribosome" title="Ribosome">ribosome</a> that reads the RNA sequence by base-pairing the messenger RNA to <a href="/wiki/Transfer_RNA" title="Transfer RNA">transfer RNA</a>, which carries amino acids. Since there are 4 bases in 3-letter combinations, there are 64 possible codons (4<sup>3</sup> combinations). These encode the twenty <a href="/wiki/List_of_standard_amino_acids" class="mw-redirect" title="List of standard amino acids">standard amino acids</a>, giving most amino acids more than one possible codon. There are also three 'stop' or 'nonsense' codons signifying the end of the coding region; these are the TAG, TAA, and TGA codons, (UAG, UAA, and UGA on the mRNA). </p> <div class="mw-heading mw-heading3"><h3 id="Replication">Replication</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:DNA_replication_en.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8f/DNA_replication_en.svg/390px-DNA_replication_en.svg.png" decoding="async" width="390" height="190" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8f/DNA_replication_en.svg/585px-DNA_replication_en.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8f/DNA_replication_en.svg/780px-DNA_replication_en.svg.png 2x" data-file-width="691" data-file-height="336" /></a><figcaption>DNA replication: The double helix is unwound by a <a href="/wiki/Helicase" title="Helicase">helicase</a> and <a href="/wiki/Topoisomerase" title="Topoisomerase">topo­iso­merase</a>. Next, one <a href="/wiki/DNA_polymerase" title="DNA polymerase">DNA polymerase</a> produces the <a href="/wiki/Replication_fork" class="mw-redirect" title="Replication fork">leading strand</a> copy. Another DNA polymerase binds to the <a href="/wiki/Replication_fork" class="mw-redirect" title="Replication fork">lagging strand</a>. This enzyme makes discontinuous segments (called <a href="/wiki/Okazaki_fragment" class="mw-redirect" title="Okazaki fragment">Okazaki fragments</a>) before <a href="/wiki/DNA_ligase" title="DNA ligase">DNA ligase</a> joins them together.</figcaption></figure> <p><a href="/wiki/Cell_division" title="Cell division">Cell division</a> is essential for an organism to grow, but, when a cell divides, it must replicate the DNA in its genome so that the two daughter cells have the same genetic information as their parent. The double-stranded structure of DNA provides a simple mechanism for <a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a>. Here, the two strands are separated and then each strand's <a href="/wiki/Complementary_DNA" title="Complementary DNA">complementary DNA</a> sequence is recreated by an <a href="/wiki/Enzyme" title="Enzyme">enzyme</a> called <a href="/wiki/DNA_polymerase" title="DNA polymerase">DNA polymerase</a>. This enzyme makes the complementary strand by finding the correct base through complementary base pairing and bonding it onto the original strand. As DNA polymerases can only extend a DNA strand in a 5′ to 3′ direction, different mechanisms are used to copy the antiparallel strands of the double helix.<sup id="cite_ref-105" class="reference"><a href="#cite_note-105"><span class="cite-bracket">[</span>105<span class="cite-bracket">]</span></a></sup> In this way, the base on the old strand dictates which base appears on the new strand, and the cell ends up with a perfect copy of its DNA. </p> <div class="mw-heading mw-heading3"><h3 id="Extracellular_nucleic_acids">Extracellular nucleic acids</h3></div> <p>Naked extracellular DNA (eDNA), most of it released by cell death, is nearly ubiquitous in the environment. Its concentration in soil may be as high as 2 μg/L, and its concentration in natural aquatic environments may be as high at 88 μg/L.<sup id="cite_ref-Tani_2010_106-0" class="reference"><a href="#cite_note-Tani_2010-106"><span class="cite-bracket">[</span>106<span class="cite-bracket">]</span></a></sup> Various possible functions have been proposed for eDNA: it may be involved in <a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">horizontal gene transfer</a>;<sup id="cite_ref-Vlassov_2007_107-0" class="reference"><a href="#cite_note-Vlassov_2007-107"><span class="cite-bracket">[</span>107<span class="cite-bracket">]</span></a></sup> it may provide nutrients;<sup id="cite_ref-pmid11591672_108-0" class="reference"><a href="#cite_note-pmid11591672-108"><span class="cite-bracket">[</span>108<span class="cite-bracket">]</span></a></sup> and it may act as a buffer to recruit or titrate ions or antibiotics.<sup id="cite_ref-Mulcahy_2008_109-0" class="reference"><a href="#cite_note-Mulcahy_2008-109"><span class="cite-bracket">[</span>109<span class="cite-bracket">]</span></a></sup> Extracellular DNA acts as a functional extracellular matrix component in the <a href="/wiki/Biofilm" title="Biofilm">biofilms</a> of several bacterial species. It may act as a recognition factor to regulate the attachment and dispersal of specific cell types in the biofilm;<sup id="cite_ref-Berne_2010_110-0" class="reference"><a href="#cite_note-Berne_2010-110"><span class="cite-bracket">[</span>110<span class="cite-bracket">]</span></a></sup> it may contribute to biofilm formation;<sup id="cite_ref-Whitchurch_2002_111-0" class="reference"><a href="#cite_note-Whitchurch_2002-111"><span class="cite-bracket">[</span>111<span class="cite-bracket">]</span></a></sup> and it may contribute to the biofilm's physical strength and resistance to biological stress.<sup id="cite_ref-Hu_2012_112-0" class="reference"><a href="#cite_note-Hu_2012-112"><span class="cite-bracket">[</span>112<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Cell-free_fetal_DNA" title="Cell-free fetal DNA">Cell-free fetal DNA</a> is found in the blood of the mother, and can be sequenced to determine a great deal of information about the developing fetus.<sup id="cite_ref-Hui_2013_113-0" class="reference"><a href="#cite_note-Hui_2013-113"><span class="cite-bracket">[</span>113<span class="cite-bracket">]</span></a></sup> </p><p>Under the name of <a href="/wiki/Environmental_DNA" title="Environmental DNA">environmental DNA</a> eDNA has seen increased use in the natural sciences as a survey tool for <a href="/wiki/Ecology" title="Ecology">ecology</a>, monitoring the movements and presence of species in water, air, or on land, and assessing an area's biodiversity.<sup id="cite_ref-114" class="reference"><a href="#cite_note-114"><span class="cite-bracket">[</span>114<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-115" class="reference"><a href="#cite_note-115"><span class="cite-bracket">[</span>115<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Neutrophil_extracellular_traps">Neutrophil extracellular traps</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Neutrophil_extracellular_traps" title="Neutrophil extracellular traps">Neutrophil extracellular traps</a></div> <p>Neutrophil extracellular traps (NETs) are networks of extracellular fibers, primarily composed of DNA, which allow <a href="/wiki/Neutrophils" class="mw-redirect" title="Neutrophils">neutrophils</a>, a type of white blood cell, to kill extracellular pathogens while minimizing damage to the host cells. </p> <div class="mw-heading mw-heading2"><h2 id="Interactions_with_proteins">Interactions with proteins</h2></div> <p>All the functions of DNA depend on interactions with proteins. These <a href="/wiki/Protein_interactions" class="mw-redirect" title="Protein interactions">protein interactions</a> can be non-specific, or the protein can bind specifically to a single DNA sequence. Enzymes can also bind to DNA and of these, the polymerases that copy the DNA base sequence in transcription and DNA replication are particularly important. </p> <div class="mw-heading mw-heading3"><h3 id="DNA-binding_proteins">DNA-binding proteins</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/DNA-binding_protein" title="DNA-binding protein">DNA-binding protein</a></div> <figure class="mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Nucleosome1.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/be/Nucleosome1.png/260px-Nucleosome1.png" decoding="async" width="260" height="200" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/be/Nucleosome1.png/390px-Nucleosome1.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/be/Nucleosome1.png/520px-Nucleosome1.png 2x" data-file-width="1984" data-file-height="1525" /></a><figcaption>Interaction of DNA (in orange) with <a href="/wiki/Histone" title="Histone">histones</a> (in blue). These proteins' basic amino acids bind to the acidic phosphate groups on DNA.</figcaption></figure> <p>Structural proteins that bind DNA are well-understood examples of non-specific DNA-protein interactions. Within chromosomes, DNA is held in complexes with structural proteins. These proteins organize the DNA into a compact structure called <a href="/wiki/Chromatin" title="Chromatin">chromatin</a>. In eukaryotes, this structure involves DNA binding to a complex of small basic proteins called <a href="/wiki/Histone" title="Histone">histones</a>, while in prokaryotes multiple types of proteins are involved.<sup id="cite_ref-116" class="reference"><a href="#cite_note-116"><span class="cite-bracket">[</span>116<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-117" class="reference"><a href="#cite_note-117"><span class="cite-bracket">[</span>117<span class="cite-bracket">]</span></a></sup> The histones form a disk-shaped complex called a <a href="/wiki/Nucleosome" title="Nucleosome">nucleosome</a>, which contains two complete turns of double-stranded DNA wrapped around its surface. These non-specific interactions are formed through basic residues in the histones, making <a href="/wiki/Ionic_bond" class="mw-redirect" title="Ionic bond">ionic bonds</a> to the acidic sugar-phosphate backbone of the DNA, and are thus largely independent of the base sequence.<sup id="cite_ref-118" class="reference"><a href="#cite_note-118"><span class="cite-bracket">[</span>118<span class="cite-bracket">]</span></a></sup> Chemical modifications of these basic amino acid residues include <a href="/wiki/Methylation" title="Methylation">methylation</a>, <a href="/wiki/Phosphorylation" title="Phosphorylation">phosphorylation</a>, and <a href="/wiki/Acetylation" title="Acetylation">acetylation</a>.<sup id="cite_ref-119" class="reference"><a href="#cite_note-119"><span class="cite-bracket">[</span>119<span class="cite-bracket">]</span></a></sup> These chemical changes alter the strength of the interaction between the DNA and the histones, making the DNA more or less accessible to <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factors</a> and changing the rate of transcription.<sup id="cite_ref-120" class="reference"><a href="#cite_note-120"><span class="cite-bracket">[</span>120<span class="cite-bracket">]</span></a></sup> Other non-specific DNA-binding proteins in chromatin include the high-mobility group proteins, which bind to bent or distorted DNA.<sup id="cite_ref-121" class="reference"><a href="#cite_note-121"><span class="cite-bracket">[</span>121<span class="cite-bracket">]</span></a></sup> These proteins are important in bending arrays of nucleosomes and arranging them into the larger structures that make up chromosomes.<sup id="cite_ref-122" class="reference"><a href="#cite_note-122"><span class="cite-bracket">[</span>122<span class="cite-bracket">]</span></a></sup> </p><p>A distinct group of DNA-binding proteins is the DNA-binding proteins that specifically bind single-stranded DNA. In humans, replication <a href="/wiki/Protein_A" title="Protein A">protein A</a> is the best-understood member of this family and is used in processes where the double helix is separated, including DNA replication, recombination, and DNA repair.<sup id="cite_ref-123" class="reference"><a href="#cite_note-123"><span class="cite-bracket">[</span>123<span class="cite-bracket">]</span></a></sup> These binding proteins seem to stabilize single-stranded DNA and protect it from forming <a href="/wiki/Stem-loop" title="Stem-loop">stem-loops</a> or being degraded by <a href="/wiki/Nuclease" title="Nuclease">nucleases</a>. </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Lambda_repressor_1LMB.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8f/Lambda_repressor_1LMB.png/240px-Lambda_repressor_1LMB.png" decoding="async" width="240" height="353" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8f/Lambda_repressor_1LMB.png/360px-Lambda_repressor_1LMB.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8f/Lambda_repressor_1LMB.png/480px-Lambda_repressor_1LMB.png 2x" data-file-width="1068" data-file-height="1569" /></a><figcaption>The lambda repressor <a href="/wiki/Helix-turn-helix" title="Helix-turn-helix">helix-turn-helix</a> transcription factor bound to its DNA target<sup id="cite_ref-124" class="reference"><a href="#cite_note-124"><span class="cite-bracket">[</span>124<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>In contrast, other proteins have evolved to bind to particular DNA sequences. The most intensively studied of these are the various <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factors</a>, which are proteins that regulate transcription. Each transcription factor binds to one particular set of DNA sequences and activates or inhibits the transcription of genes that have these sequences close to their promoters. The transcription factors do this in two ways. Firstly, they can bind the RNA polymerase responsible for transcription, either directly or through other mediator proteins; this locates the polymerase at the promoter and allows it to begin transcription.<sup id="cite_ref-125" class="reference"><a href="#cite_note-125"><span class="cite-bracket">[</span>125<span class="cite-bracket">]</span></a></sup> Alternatively, transcription factors can bind <a href="/wiki/Enzyme" title="Enzyme">enzymes</a> that modify the histones at the promoter. This changes the accessibility of the DNA template to the polymerase.<sup id="cite_ref-126" class="reference"><a href="#cite_note-126"><span class="cite-bracket">[</span>126<span class="cite-bracket">]</span></a></sup> </p><p>As these DNA targets can occur throughout an organism's genome, changes in the activity of one type of transcription factor can affect thousands of genes.<sup id="cite_ref-127" class="reference"><a href="#cite_note-127"><span class="cite-bracket">[</span>127<span class="cite-bracket">]</span></a></sup> Consequently, these proteins are often the targets of the <a href="/wiki/Signal_transduction" title="Signal transduction">signal transduction</a> processes that control responses to environmental changes or <a href="/wiki/Cellular_differentiation" title="Cellular differentiation">cellular differentiation</a> and development. The specificity of these transcription factors' interactions with DNA come from the proteins making multiple contacts to the edges of the DNA bases, allowing them to "read" the DNA sequence. Most of these base-interactions are made in the major groove, where the bases are most accessible.<sup id="cite_ref-Pabo1984_25-1" class="reference"><a href="#cite_note-Pabo1984-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="DNA-modifying_enzymes">DNA-modifying enzymes</h3></div> <div class="mw-heading mw-heading4"><h4 id="Nucleases_and_ligases">Nucleases and ligases</h4></div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:EcoRV_1RVA.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/dc/EcoRV_1RVA.png/240px-EcoRV_1RVA.png" decoding="async" width="240" height="168" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/dc/EcoRV_1RVA.png/360px-EcoRV_1RVA.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/dc/EcoRV_1RVA.png/480px-EcoRV_1RVA.png 2x" data-file-width="1452" data-file-height="1017" /></a><figcaption>The <a href="/wiki/Restriction_enzyme" title="Restriction enzyme">restriction enzyme</a> <a href="/wiki/EcoRV" title="EcoRV">EcoRV</a> (green) in a complex with its substrate DNA<sup id="cite_ref-128" class="reference"><a href="#cite_note-128"><span class="cite-bracket">[</span>128<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p><a href="/wiki/Nuclease" title="Nuclease">Nucleases</a> are <a href="/wiki/Enzyme" title="Enzyme">enzymes</a> that cut DNA strands by catalyzing the <a href="/wiki/Hydrolysis" title="Hydrolysis">hydrolysis</a> of the <a href="/wiki/Phosphodiester_bond" title="Phosphodiester bond">phosphodiester bonds</a>. Nucleases that hydrolyse nucleotides from the ends of DNA strands are called <a href="/wiki/Exonuclease" title="Exonuclease">exonucleases</a>, while <a href="/wiki/Endonuclease" title="Endonuclease">endonucleases</a> cut within strands. The most frequently used nucleases in <a href="/wiki/Molecular_biology" title="Molecular biology">molecular biology</a> are the <a href="/wiki/Restriction_enzyme" title="Restriction enzyme">restriction endonucleases</a>, which cut DNA at specific sequences. For instance, the EcoRV enzyme shown to the left recognizes the 6-base sequence 5′-GATATC-3′ and makes a cut at the horizontal line. In nature, these enzymes protect <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> against <a href="/wiki/Bacteriophage" title="Bacteriophage">phage</a> infection by digesting the phage DNA when it enters the bacterial cell, acting as part of the <a href="/wiki/Restriction_modification_system" title="Restriction modification system">restriction modification system</a>.<sup id="cite_ref-129" class="reference"><a href="#cite_note-129"><span class="cite-bracket">[</span>129<span class="cite-bracket">]</span></a></sup> In technology, these sequence-specific nucleases are used in <a href="/wiki/Molecular_cloning" title="Molecular cloning">molecular cloning</a> and <a href="/wiki/Genetic_fingerprinting" class="mw-redirect" title="Genetic fingerprinting">DNA fingerprinting</a>. </p><p>Enzymes called <a href="/wiki/DNA_ligase" title="DNA ligase">DNA ligases</a> can rejoin cut or broken DNA strands.<sup id="cite_ref-Doherty_130-0" class="reference"><a href="#cite_note-Doherty-130"><span class="cite-bracket">[</span>130<span class="cite-bracket">]</span></a></sup> Ligases are particularly important in <a href="/wiki/Replication_fork" class="mw-redirect" title="Replication fork">lagging strand</a> DNA replication, as they join the short segments of DNA produced at the <a href="/wiki/Replication_fork" class="mw-redirect" title="Replication fork">replication fork</a> into a complete copy of the DNA template. They are also used in <a href="/wiki/DNA_repair" title="DNA repair">DNA repair</a> and <a href="/wiki/Genetic_recombination" title="Genetic recombination">genetic recombination</a>.<sup id="cite_ref-Doherty_130-1" class="reference"><a href="#cite_note-Doherty-130"><span class="cite-bracket">[</span>130<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Topoisomerases_and_helicases">Topoisomerases and helicases</h4></div> <p><a href="/wiki/Topoisomerase" title="Topoisomerase">Topoisomerases</a> are enzymes with both nuclease and ligase activity. These proteins change the amount of <a href="/wiki/DNA_supercoil" title="DNA supercoil">supercoiling</a> in DNA. Some of these enzymes work by cutting the DNA helix and allowing one section to rotate, thereby reducing its level of supercoiling; the enzyme then seals the DNA break.<sup id="cite_ref-Champoux_44-1" class="reference"><a href="#cite_note-Champoux-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> Other types of these enzymes are capable of cutting one DNA helix and then passing a second strand of DNA through this break, before rejoining the helix.<sup id="cite_ref-131" class="reference"><a href="#cite_note-131"><span class="cite-bracket">[</span>131<span class="cite-bracket">]</span></a></sup> Topoisomerases are required for many processes involving DNA, such as DNA replication and transcription.<sup id="cite_ref-Wang_45-1" class="reference"><a href="#cite_note-Wang-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Helicase" title="Helicase">Helicases</a> are proteins that are a type of <a href="/wiki/Molecular_motor" title="Molecular motor">molecular motor</a>. They use the chemical energy in <a href="/wiki/Nucleoside_triphosphate" title="Nucleoside triphosphate">nucleoside triphosphates</a>, predominantly <a href="/wiki/Adenosine_triphosphate" title="Adenosine triphosphate">adenosine triphosphate</a> (ATP), to break hydrogen bonds between bases and unwind the DNA double helix into single strands.<sup id="cite_ref-132" class="reference"><a href="#cite_note-132"><span class="cite-bracket">[</span>132<span class="cite-bracket">]</span></a></sup> These enzymes are essential for most processes where enzymes need to access the DNA bases. </p> <div class="mw-heading mw-heading4"><h4 id="Polymerases">Polymerases</h4></div> <p><a href="/wiki/Polymerase" title="Polymerase">Polymerases</a> are <a href="/wiki/Enzyme" title="Enzyme">enzymes</a> that synthesize polynucleotide chains from <a href="/wiki/Nucleoside_triphosphate" title="Nucleoside triphosphate">nucleoside triphosphates</a>. The sequence of their products is created based on existing polynucleotide chains—which are called <i>templates</i>. These enzymes function by repeatedly adding a nucleotide to the 3′ <a href="/wiki/Hydroxyl" class="mw-redirect" title="Hydroxyl">hydroxyl</a> group at the end of the growing polynucleotide chain. As a consequence, all polymerases work in a 5′ to 3′ direction.<sup id="cite_ref-Joyce_133-0" class="reference"><a href="#cite_note-Joyce-133"><span class="cite-bracket">[</span>133<span class="cite-bracket">]</span></a></sup> In the <a href="/wiki/Active_site" title="Active site">active site</a> of these enzymes, the incoming nucleoside triphosphate base-pairs to the template: this allows polymerases to accurately synthesize the complementary strand of their template. Polymerases are classified according to the type of template that they use. </p><p>In DNA replication, DNA-dependent <a href="/wiki/DNA_polymerase" title="DNA polymerase">DNA polymerases</a> make copies of DNA polynucleotide chains. To preserve biological information, it is essential that the sequence of bases in each copy are precisely complementary to the sequence of bases in the template strand. Many DNA polymerases have a <a href="/wiki/Proofreading_(biology)" title="Proofreading (biology)">proofreading</a> activity. Here, the polymerase recognizes the occasional mistakes in the synthesis reaction by the lack of base pairing between the mismatched nucleotides. If a mismatch is detected, a 3′ to 5′ <a href="/wiki/Exonuclease" title="Exonuclease">exonuclease</a> activity is activated and the incorrect base removed.<sup id="cite_ref-134" class="reference"><a href="#cite_note-134"><span class="cite-bracket">[</span>134<span class="cite-bracket">]</span></a></sup> In most organisms, DNA polymerases function in a large complex called the <a href="/wiki/Replisome" title="Replisome">replisome</a> that contains multiple accessory subunits, such as the <a href="/wiki/DNA_clamp" title="DNA clamp">DNA clamp</a> or <a href="/wiki/Helicase" title="Helicase">helicases</a>.<sup id="cite_ref-135" class="reference"><a href="#cite_note-135"><span class="cite-bracket">[</span>135<span class="cite-bracket">]</span></a></sup> </p><p>RNA-dependent DNA polymerases are a specialized class of polymerases that copy the sequence of an RNA strand into DNA. They include <a href="/wiki/Reverse_transcriptase" title="Reverse transcriptase">reverse transcriptase</a>, which is a <a href="/wiki/Virus" title="Virus">viral</a> enzyme involved in the infection of cells by <a href="/wiki/Retrovirus" title="Retrovirus">retroviruses</a>, and <a href="/wiki/Telomerase" title="Telomerase">telomerase</a>, which is required for the replication of telomeres.<sup id="cite_ref-Greider_63-1" class="reference"><a href="#cite_note-Greider-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Tarrago-Litvak1994_136-0" class="reference"><a href="#cite_note-Tarrago-Litvak1994-136"><span class="cite-bracket">[</span>136<span class="cite-bracket">]</span></a></sup> For example, HIV reverse transcriptase is an enzyme for AIDS virus replication.<sup id="cite_ref-Tarrago-Litvak1994_136-1" class="reference"><a href="#cite_note-Tarrago-Litvak1994-136"><span class="cite-bracket">[</span>136<span class="cite-bracket">]</span></a></sup> Telomerase is an unusual polymerase because it contains its own RNA template as part of its structure. It synthesizes <a href="/wiki/Telomeres" class="mw-redirect" title="Telomeres">telomeres</a> at the ends of chromosomes. Telomeres prevent fusion of the ends of neighboring chromosomes and protect chromosome ends from damage.<sup id="cite_ref-Nugent_64-2" class="reference"><a href="#cite_note-Nugent-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> </p><p>Transcription is carried out by a DNA-dependent <a href="/wiki/RNA_polymerase" title="RNA polymerase">RNA polymerase</a> that copies the sequence of a DNA strand into RNA. To begin transcribing a gene, the RNA polymerase binds to a sequence of DNA called a promoter and separates the DNA strands. It then copies the gene sequence into a <a href="/wiki/Messenger_RNA" title="Messenger RNA">messenger RNA</a> transcript until it reaches a region of DNA called the <a href="/wiki/Terminator_(genetics)" title="Terminator (genetics)">terminator</a>, where it halts and detaches from the DNA. As with human DNA-dependent DNA polymerases, <a href="/wiki/RNA_polymerase_II" title="RNA polymerase II">RNA polymerase II</a>, the enzyme that transcribes most of the genes in the human genome, operates as part of a large <a href="/wiki/Protein_complex" title="Protein complex">protein complex</a> with multiple regulatory and accessory subunits.<sup id="cite_ref-137" class="reference"><a href="#cite_note-137"><span class="cite-bracket">[</span>137<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Genetic_recombination">Genetic recombination</h2></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Genetic_recombination" title="Genetic recombination">Genetic recombination</a></div> <div class="thumb tright" style="background:#f9f9f9; border:1px solid #ccc; margin:0.5em;"> <table border="0" cellpadding="0" cellspacing="0" style="width:250px; font-size:85%; border:1px solid #ccc; margin:0.3em;"> <tbody><tr> <td><span typeof="mw:File"><a href="/wiki/File:Holliday_Junction.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/83/Holliday_Junction.svg/250px-Holliday_Junction.svg.png" decoding="async" width="250" height="199" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/83/Holliday_Junction.svg/375px-Holliday_Junction.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/83/Holliday_Junction.svg/500px-Holliday_Junction.svg.png 2x" data-file-width="472" data-file-height="376" /></a></span> </td></tr> <tr> <td><span typeof="mw:File"><a href="/wiki/File:Holliday_junction_coloured.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/92/Holliday_junction_coloured.png/250px-Holliday_junction_coloured.png" decoding="async" width="250" height="250" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/92/Holliday_junction_coloured.png/375px-Holliday_junction_coloured.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/92/Holliday_junction_coloured.png/500px-Holliday_junction_coloured.png 2x" data-file-width="1620" data-file-height="1620" /></a></span> </td></tr></tbody></table> <div style="border: none; width:250px;"><div class="thumbcaption">Structure of the <a href="/wiki/Holliday_junction" title="Holliday junction">Holliday junction</a> intermediate in <a href="/wiki/Genetic_recombination" title="Genetic recombination">genetic recombination</a>. The four separate DNA strands are coloured red, blue, green and yellow.<sup id="cite_ref-138" class="reference"><a href="#cite_note-138"><span class="cite-bracket">[</span>138<span class="cite-bracket">]</span></a></sup></div></div></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Homologous_Recombination.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6d/Homologous_Recombination.jpg/300px-Homologous_Recombination.jpg" decoding="async" width="300" height="359" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/6/6d/Homologous_Recombination.jpg 1.5x" data-file-width="393" data-file-height="470" /></a><figcaption> A current model of meiotic recombination, initiated by a double-strand break or gap, followed by pairing with an homologous chromosome and strand invasion to initiate the recombinational repair process. Repair of the gap can lead to crossover (CO) or non-crossover (NCO) of the flanking regions. CO recombination is thought to occur by the Double Holliday Junction (DHJ) model, illustrated on the right, above. NCO recombinants are thought to occur primarily by the Synthesis Dependent Strand Annealing (SDSA) model, illustrated on the left, above. Most recombination events appear to be the SDSA type.</figcaption></figure> <p>A DNA helix usually does not interact with other segments of DNA, and in human cells, the different chromosomes even occupy separate areas in the nucleus called "<a href="/wiki/Chromosome_territories" title="Chromosome territories">chromosome territories</a>".<sup id="cite_ref-139" class="reference"><a href="#cite_note-139"><span class="cite-bracket">[</span>139<span class="cite-bracket">]</span></a></sup> This physical separation of different chromosomes is important for the ability of DNA to function as a stable repository for information, as one of the few times chromosomes interact is in <a href="/wiki/Chromosomal_crossover" title="Chromosomal crossover">chromosomal crossover</a> which occurs during <a href="/wiki/Sexual_reproduction" title="Sexual reproduction">sexual reproduction</a>, when <a href="/wiki/Genetic_recombination" title="Genetic recombination">genetic recombination</a> occurs. Chromosomal crossover is when two DNA helices break, swap a section and then rejoin. </p><p>Recombination allows chromosomes to exchange genetic information and produces new combinations of genes, which increases the efficiency of <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> and can be important in the rapid evolution of new proteins.<sup id="cite_ref-140" class="reference"><a href="#cite_note-140"><span class="cite-bracket">[</span>140<span class="cite-bracket">]</span></a></sup> Genetic recombination can also be involved in DNA repair, particularly in the cell's response to double-strand breaks.<sup id="cite_ref-141" class="reference"><a href="#cite_note-141"><span class="cite-bracket">[</span>141<span class="cite-bracket">]</span></a></sup> </p><p>The most common form of chromosomal crossover is <a href="/wiki/Homologous_recombination" title="Homologous recombination">homologous recombination</a>, where the two chromosomes involved share very similar sequences. <a href="/wiki/Non-homologous_recombination" class="mw-redirect" title="Non-homologous recombination">Non-homologous recombination</a> can be damaging to cells, as it can produce <a href="/wiki/Chromosomal_translocation" title="Chromosomal translocation">chromosomal translocations</a> and genetic abnormalities. The recombination reaction is catalyzed by enzymes known as <a href="/wiki/Recombinase" title="Recombinase">recombinases</a>, such as <a href="/wiki/RAD51" title="RAD51">RAD51</a>.<sup id="cite_ref-142" class="reference"><a href="#cite_note-142"><span class="cite-bracket">[</span>142<span class="cite-bracket">]</span></a></sup> The first step in recombination is a double-stranded break caused by either an <a href="/wiki/Endonuclease" title="Endonuclease">endonuclease</a> or damage to the DNA.<sup id="cite_ref-143" class="reference"><a href="#cite_note-143"><span class="cite-bracket">[</span>143<span class="cite-bracket">]</span></a></sup> A series of steps catalyzed in part by the recombinase then leads to joining of the two helices by at least one <a href="/wiki/Holliday_junction" title="Holliday junction">Holliday junction</a>, in which a segment of a single strand in each helix is annealed to the complementary strand in the other helix. The Holliday junction is a tetrahedral junction structure that can be moved along the pair of chromosomes, swapping one strand for another. The recombination reaction is then halted by cleavage of the junction and re-ligation of the released DNA.<sup id="cite_ref-144" class="reference"><a href="#cite_note-144"><span class="cite-bracket">[</span>144<span class="cite-bracket">]</span></a></sup> Only strands of like polarity exchange DNA during recombination. There are two types of cleavage: east-west cleavage and north–south cleavage. The north–south cleavage nicks both strands of DNA, while the east–west cleavage has one strand of DNA intact. The formation of a Holliday junction during recombination makes it possible for genetic diversity, genes to exchange on chromosomes, and expression of wild-type viral genomes. </p> <div class="mw-heading mw-heading2"><h2 id="Evolution">Evolution</h2></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/RNA_world_hypothesis" class="mw-redirect" title="RNA world hypothesis">RNA world hypothesis</a></div> <p>DNA contains the genetic information that allows all forms of life to function, grow and reproduce. However, it is unclear how long in the 4-billion-year <a href="/wiki/Timeline_of_evolution" class="mw-redirect" title="Timeline of evolution">history of life</a> DNA has performed this function, as it has been proposed that the earliest forms of life may have used RNA as their genetic material.<sup id="cite_ref-Joyce-2002_145-0" class="reference"><a href="#cite_note-Joyce-2002-145"><span class="cite-bracket">[</span>145<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-146" class="reference"><a href="#cite_note-146"><span class="cite-bracket">[</span>146<span class="cite-bracket">]</span></a></sup> RNA may have acted as the central part of early <a href="/wiki/Cell_metabolism" class="mw-redirect" title="Cell metabolism">cell metabolism</a> as it can both transmit genetic information and carry out <a href="/wiki/Catalysis" title="Catalysis">catalysis</a> as part of <a href="/wiki/Ribozyme" title="Ribozyme">ribozymes</a>.<sup id="cite_ref-147" class="reference"><a href="#cite_note-147"><span class="cite-bracket">[</span>147<span class="cite-bracket">]</span></a></sup> This ancient <a href="/wiki/RNA_world_hypothesis" class="mw-redirect" title="RNA world hypothesis">RNA world</a> where nucleic acid would have been used for both catalysis and genetics may have influenced the <a href="/wiki/Evolution" title="Evolution">evolution</a> of the current genetic code based on four nucleotide bases. This would occur, since the number of different bases in such an organism is a trade-off between a small number of bases increasing replication accuracy and a large number of bases increasing the catalytic efficiency of ribozymes.<sup id="cite_ref-148" class="reference"><a href="#cite_note-148"><span class="cite-bracket">[</span>148<span class="cite-bracket">]</span></a></sup> However, there is no direct evidence of ancient genetic systems, as recovery of DNA from most fossils is impossible because DNA survives in the environment for less than one million years, and slowly degrades into short fragments in solution.<sup id="cite_ref-149" class="reference"><a href="#cite_note-149"><span class="cite-bracket">[</span>149<span class="cite-bracket">]</span></a></sup> Claims for older DNA have been made, most notably a report of the isolation of a viable bacterium from a salt crystal 250 million years old,<sup id="cite_ref-150" class="reference"><a href="#cite_note-150"><span class="cite-bracket">[</span>150<span class="cite-bracket">]</span></a></sup> but these claims are controversial.<sup id="cite_ref-151" class="reference"><a href="#cite_note-151"><span class="cite-bracket">[</span>151<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-152" class="reference"><a href="#cite_note-152"><span class="cite-bracket">[</span>152<span class="cite-bracket">]</span></a></sup> </p><p>Building blocks of DNA (<a href="/wiki/Adenine" title="Adenine">adenine</a>, <a href="/wiki/Guanine" title="Guanine">guanine</a>, and related <a href="/wiki/Organic_molecules" class="mw-redirect" title="Organic molecules">organic molecules</a>) may have been formed extraterrestrially in <a href="/wiki/Outer_space" title="Outer space">outer space</a>.<sup id="cite_ref-Callahan_153-0" class="reference"><a href="#cite_note-Callahan-153"><span class="cite-bracket">[</span>153<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Steigerwald_154-0" class="reference"><a href="#cite_note-Steigerwald-154"><span class="cite-bracket">[</span>154<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-DNA_155-0" class="reference"><a href="#cite_note-DNA-155"><span class="cite-bracket">[</span>155<span class="cite-bracket">]</span></a></sup> Complex DNA and <a href="/wiki/RNA" title="RNA">RNA</a> <a href="/wiki/Organic_compound" title="Organic compound">organic compounds</a> of <a href="/wiki/Life" title="Life">life</a>, including <a href="/wiki/Uracil" title="Uracil">uracil</a>, <a href="/wiki/Cytosine" title="Cytosine">cytosine</a>, and <a href="/wiki/Thymine" title="Thymine">thymine</a>, have also been formed in the laboratory under conditions mimicking those found in <a href="/wiki/Outer_space" title="Outer space">outer space</a>, using starting chemicals, such as <a href="/wiki/Pyrimidine" title="Pyrimidine">pyrimidine</a>, found in <a href="/wiki/Meteorite" title="Meteorite">meteorites</a>. Pyrimidine, like <a href="/wiki/Polycyclic_aromatic_hydrocarbons" class="mw-redirect" title="Polycyclic aromatic hydrocarbons">polycyclic aromatic hydrocarbons</a> (PAHs), the most carbon-rich chemical found in the <a href="/wiki/Universe" title="Universe">universe</a>, may have been formed in <a href="/wiki/Red_giant" title="Red giant">red giants</a> or in interstellar <a href="/wiki/Cosmic_dust" title="Cosmic dust">cosmic dust</a> and gas clouds.<sup id="cite_ref-NASA-20150303_156-0" class="reference"><a href="#cite_note-NASA-20150303-156"><span class="cite-bracket">[</span>156<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Ancient_DNA" title="Ancient DNA">Ancient DNA</a> has been recovered from ancient organisms at a timescale where genome evolution can be directly observed, including from extinct organisms up to millions of years old, such as the <a href="/wiki/Woolly_mammoth" title="Woolly mammoth">woolly mammoth</a>.<sup id="cite_ref-CNN-20210217_157-0" class="reference"><a href="#cite_note-CNN-20210217-157"><span class="cite-bracket">[</span>157<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-NAT-20210217_158-0" class="reference"><a href="#cite_note-NAT-20210217-158"><span class="cite-bracket">[</span>158<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Uses_in_technology">Uses in technology</h2></div> <div class="mw-heading mw-heading3"><h3 id="Genetic_engineering">Genetic engineering</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Molecular_biology" title="Molecular biology">Molecular biology</a>, <a href="/wiki/Nucleic_acid_methods" title="Nucleic acid methods">Nucleic acid methods</a>, and <a href="/wiki/Genetic_engineering" title="Genetic engineering">Genetic engineering</a></div> <p>Methods have been developed to purify DNA from organisms, such as <a href="/wiki/Phenol-chloroform_extraction" class="mw-redirect" title="Phenol-chloroform extraction">phenol-chloroform extraction</a>, and to manipulate it in the laboratory, such as <a href="/wiki/Restriction_digest" title="Restriction digest">restriction digests</a> and the <a href="/wiki/Polymerase_chain_reaction" title="Polymerase chain reaction">polymerase chain reaction</a>. Modern <a href="/wiki/Biology" title="Biology">biology</a> and <a href="/wiki/Biochemistry" title="Biochemistry">biochemistry</a> make intensive use of these techniques in recombinant DNA technology. <a href="/wiki/Recombinant_DNA" title="Recombinant DNA">Recombinant DNA</a> is a man-made DNA sequence that has been assembled from other DNA sequences. They can be <a href="/wiki/Transformation_(genetics)" class="mw-redirect" title="Transformation (genetics)">transformed</a> into organisms in the form of <a href="/wiki/Plasmid" title="Plasmid">plasmids</a> or in the appropriate format, by using a <a href="/wiki/Viral_vector" title="Viral vector">viral vector</a>.<sup id="cite_ref-159" class="reference"><a href="#cite_note-159"><span class="cite-bracket">[</span>159<span class="cite-bracket">]</span></a></sup> The <a href="/wiki/Genetic_engineering" title="Genetic engineering">genetically modified</a> organisms produced can be used to produce products such as recombinant <a href="/wiki/Protein" title="Protein">proteins</a>, used in <a href="/wiki/Medical_research" title="Medical research">medical research</a>,<sup id="cite_ref-160" class="reference"><a href="#cite_note-160"><span class="cite-bracket">[</span>160<span class="cite-bracket">]</span></a></sup> or be grown in <a href="/wiki/Agriculture" title="Agriculture">agriculture</a>.<sup id="cite_ref-161" class="reference"><a href="#cite_note-161"><span class="cite-bracket">[</span>161<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-162" class="reference"><a href="#cite_note-162"><span class="cite-bracket">[</span>162<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="DNA_profiling">DNA profiling</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/DNA_profiling" title="DNA profiling">DNA profiling</a></div> <p><a href="/wiki/Forensic_science" title="Forensic science">Forensic scientists</a> can use DNA in <a href="/wiki/Blood" title="Blood">blood</a>, <a href="/wiki/Semen" title="Semen">semen</a>, <a href="/wiki/Skin" title="Skin">skin</a>, <a href="/wiki/Saliva" title="Saliva">saliva</a> or <a href="/wiki/Hair" title="Hair">hair</a> found at a <a href="/wiki/Crime_scene" title="Crime scene">crime scene</a> to identify a matching DNA of an individual, such as a perpetrator.<sup id="cite_ref-163" class="reference"><a href="#cite_note-163"><span class="cite-bracket">[</span>163<span class="cite-bracket">]</span></a></sup> This process is formally termed <a href="/wiki/DNA_profiling" title="DNA profiling">DNA profiling</a>, also called <i>DNA fingerprinting</i>. In DNA profiling, the lengths of variable sections of repetitive DNA, such as <a href="/wiki/Short_tandem_repeat" class="mw-redirect" title="Short tandem repeat">short tandem repeats</a> and <a href="/wiki/Minisatellite" title="Minisatellite">minisatellites</a>, are compared between people. This method is usually an extremely reliable technique for identifying a matching DNA.<sup id="cite_ref-164" class="reference"><a href="#cite_note-164"><span class="cite-bracket">[</span>164<span class="cite-bracket">]</span></a></sup> However, identification can be complicated if the scene is contaminated with DNA from several people.<sup id="cite_ref-165" class="reference"><a href="#cite_note-165"><span class="cite-bracket">[</span>165<span class="cite-bracket">]</span></a></sup> DNA profiling was developed in 1984 by British geneticist Sir <a href="/wiki/Alec_Jeffreys" title="Alec Jeffreys">Alec Jeffreys</a>,<sup id="cite_ref-166" class="reference"><a href="#cite_note-166"><span class="cite-bracket">[</span>166<span class="cite-bracket">]</span></a></sup> and first used in forensic science to convict Colin Pitchfork in the 1988 <a href="/wiki/Colin_Pitchfork" title="Colin Pitchfork">Enderby murders</a> case.<sup id="cite_ref-167" class="reference"><a href="#cite_note-167"><span class="cite-bracket">[</span>167<span class="cite-bracket">]</span></a></sup> </p><p>The development of forensic science and the ability to now obtain genetic matching on minute samples of blood, skin, saliva, or hair has led to re-examining many cases. Evidence can now be uncovered that was scientifically impossible at the time of the original examination. Combined with the removal of the <a href="/wiki/Double_jeopardy" title="Double jeopardy">double jeopardy</a> law in some places, this can allow cases to be reopened where prior trials have failed to produce sufficient evidence to convince a jury. People charged with serious crimes may be required to provide a sample of DNA for matching purposes. The most obvious defense to DNA matches obtained forensically is to claim that cross-contamination of evidence has occurred. This has resulted in meticulous strict handling procedures with new cases of serious crime. </p><p>DNA profiling is also used successfully to positively identify victims of mass casualty incidents,<sup id="cite_ref-168" class="reference"><a href="#cite_note-168"><span class="cite-bracket">[</span>168<span class="cite-bracket">]</span></a></sup> bodies or body parts in serious accidents, and individual victims in mass war graves, via matching to family members. </p><p>DNA profiling is also used in <a href="/wiki/DNA_paternity_testing" title="DNA paternity testing">DNA paternity testing</a> to determine if someone is the biological parent or grandparent of a child with the probability of parentage is typically 99.99% when the alleged parent is biologically related to the child. Normal <a href="/wiki/DNA_sequencing" title="DNA sequencing">DNA sequencing</a> methods happen after birth, but there are new methods to test paternity while a mother is still pregnant.<sup id="cite_ref-169" class="reference"><a href="#cite_note-169"><span class="cite-bracket">[</span>169<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="DNA_enzymes_or_catalytic_DNA">DNA enzymes or catalytic DNA</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Deoxyribozyme" title="Deoxyribozyme">Deoxyribozyme</a></div> <p><a href="/wiki/Deoxyribozyme" title="Deoxyribozyme">Deoxyribozymes</a>, also called DNAzymes or catalytic DNA, were first discovered in 1994.<sup id="cite_ref-Breaker_223–229_170-0" class="reference"><a href="#cite_note-Breaker_223–229-170"><span class="cite-bracket">[</span>170<span class="cite-bracket">]</span></a></sup> They are mostly single stranded DNA sequences isolated from a large pool of random DNA sequences through a combinatorial approach called <a href="/wiki/In_vitro" title="In vitro">in vitro</a> selection or <a href="/wiki/Systematic_evolution_of_ligands_by_exponential_enrichment" title="Systematic evolution of ligands by exponential enrichment">systematic evolution of ligands by exponential enrichment</a> (SELEX). DNAzymes catalyze variety of chemical reactions including RNA-DNA cleavage, RNA-DNA ligation, amino acids phosphorylation-dephosphorylation, carbon-carbon bond formation, etc. DNAzymes can enhance catalytic rate of chemical reactions up to 100,000,000,000-fold over the uncatalyzed reaction.<sup id="cite_ref-171" class="reference"><a href="#cite_note-171"><span class="cite-bracket">[</span>171<span class="cite-bracket">]</span></a></sup> The most extensively studied class of DNAzymes is RNA-cleaving types which have been used to detect different metal ions and designing therapeutic agents. Several metal-specific DNAzymes have been reported including the GR-5 DNAzyme (lead-specific),<sup id="cite_ref-Breaker_223–229_170-1" class="reference"><a href="#cite_note-Breaker_223–229-170"><span class="cite-bracket">[</span>170<span class="cite-bracket">]</span></a></sup> the CA1-3 DNAzymes (copper-specific),<sup id="cite_ref-172" class="reference"><a href="#cite_note-172"><span class="cite-bracket">[</span>172<span class="cite-bracket">]</span></a></sup> the 39E DNAzyme (uranyl-specific) and the NaA43 DNAzyme (sodium-specific).<sup id="cite_ref-173" class="reference"><a href="#cite_note-173"><span class="cite-bracket">[</span>173<span class="cite-bracket">]</span></a></sup> The NaA43 DNAzyme, which is reported to be more than 10,000-fold selective for sodium over other metal ions, was used to make a real-time sodium sensor in cells. </p> <div class="mw-heading mw-heading3"><h3 id="Bioinformatics">Bioinformatics</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Bioinformatics" title="Bioinformatics">Bioinformatics</a></div> <p><a href="/wiki/Bioinformatics" title="Bioinformatics">Bioinformatics</a> involves the development of techniques to store, <a href="/wiki/Data_mining" title="Data mining">data mine</a>, search and manipulate biological data, including DNA <a href="/wiki/Nucleic_acid_sequence" title="Nucleic acid sequence">nucleic acid sequence</a> data. These have led to widely applied advances in <a href="/wiki/Computer_science" title="Computer science">computer science</a>, especially <a href="/wiki/String_searching_algorithm" class="mw-redirect" title="String searching algorithm">string searching algorithms</a>, <a href="/wiki/Machine_learning" title="Machine learning">machine learning</a>, and <a href="/wiki/Database_theory" title="Database theory">database theory</a>.<sup id="cite_ref-174" class="reference"><a href="#cite_note-174"><span class="cite-bracket">[</span>174<span class="cite-bracket">]</span></a></sup> String searching or matching algorithms, which find an occurrence of a sequence of letters inside a larger sequence of letters, were developed to search for specific sequences of nucleotides.<sup id="cite_ref-175" class="reference"><a href="#cite_note-175"><span class="cite-bracket">[</span>175<span class="cite-bracket">]</span></a></sup> The DNA sequence may be <a href="/wiki/Sequence_alignment" title="Sequence alignment">aligned</a> with other DNA sequences to identify <a href="/wiki/Sequence_homology" title="Sequence homology">homologous sequences</a> and locate the specific <a href="/wiki/Mutation" title="Mutation">mutations</a> that make them distinct. These techniques, especially <a href="/wiki/Multiple_sequence_alignment" title="Multiple sequence alignment">multiple sequence alignment</a>, are used in studying <a href="/wiki/Phylogenetics" title="Phylogenetics">phylogenetic</a> relationships and protein function.<sup id="cite_ref-176" class="reference"><a href="#cite_note-176"><span class="cite-bracket">[</span>176<span class="cite-bracket">]</span></a></sup> Data sets representing entire genomes' worth of DNA sequences, such as those produced by the <a href="/wiki/Human_Genome_Project" title="Human Genome Project">Human Genome Project</a>, are difficult to use without the annotations that identify the locations of genes and regulatory elements on each chromosome. Regions of DNA sequence that have the characteristic patterns associated with protein- or RNA-coding genes can be identified by <a href="/wiki/Gene_prediction" title="Gene prediction">gene finding</a> algorithms, which allow researchers to predict the presence of particular <a href="/wiki/Gene_product" title="Gene product">gene products</a> and their possible functions in an organism even before they have been isolated experimentally.<sup id="cite_ref-Mount_177-0" class="reference"><a href="#cite_note-Mount-177"><span class="cite-bracket">[</span>177<span class="cite-bracket">]</span></a></sup> Entire genomes may also be compared, which can shed light on the evolutionary history of particular organism and permit the examination of complex evolutionary events. </p> <div class="mw-heading mw-heading3"><h3 id="DNA_nanotechnology">DNA nanotechnology</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/DNA_nanotechnology" title="DNA nanotechnology">DNA nanotechnology</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:DNA_nanostructures.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/55/DNA_nanostructures.png/400px-DNA_nanostructures.png" decoding="async" width="400" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/55/DNA_nanostructures.png/600px-DNA_nanostructures.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/55/DNA_nanostructures.png/800px-DNA_nanostructures.png 2x" data-file-width="1260" data-file-height="693" /></a><figcaption>The DNA structure at left (schematic shown) will self-assemble into the structure visualized by <a href="/wiki/Atomic_force_microscope" class="mw-redirect" title="Atomic force microscope">atomic force microscopy</a> at right. <a href="/wiki/DNA_nanotechnology" title="DNA nanotechnology">DNA nanotechnology</a> is the field that seeks to design nanoscale structures using the <a href="/wiki/Molecular_recognition" title="Molecular recognition">molecular recognition</a> properties of DNA molecules.<sup id="cite_ref-178" class="reference"><a href="#cite_note-178"><span class="cite-bracket">[</span>178<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>DNA nanotechnology uses the unique <a href="/wiki/Molecular_recognition" title="Molecular recognition">molecular recognition</a> properties of DNA and other nucleic acids to create self-assembling branched DNA complexes with useful properties.<sup id="cite_ref-179" class="reference"><a href="#cite_note-179"><span class="cite-bracket">[</span>179<span class="cite-bracket">]</span></a></sup> DNA is thus used as a structural material rather than as a carrier of biological information. This has led to the creation of two-dimensional periodic lattices (both tile-based and using the <i><a href="/wiki/DNA_origami" title="DNA origami">DNA origami</a></i> method) and three-dimensional structures in the shapes of <a href="/wiki/Polyhedron" title="Polyhedron">polyhedra</a>.<sup id="cite_ref-180" class="reference"><a href="#cite_note-180"><span class="cite-bracket">[</span>180<span class="cite-bracket">]</span></a></sup> <a href="/wiki/DNA_machine" title="DNA machine">Nanomechanical devices</a> and <a href="/wiki/DNA_computing" title="DNA computing">algorithmic self-assembly</a> have also been demonstrated,<sup id="cite_ref-181" class="reference"><a href="#cite_note-181"><span class="cite-bracket">[</span>181<span class="cite-bracket">]</span></a></sup> and these DNA structures have been used to template the arrangement of other molecules such as <a href="/wiki/Colloidal_gold" title="Colloidal gold">gold nanoparticles</a> and <a href="/wiki/Streptavidin" title="Streptavidin">streptavidin</a> proteins.<sup id="cite_ref-182" class="reference"><a href="#cite_note-182"><span class="cite-bracket">[</span>182<span class="cite-bracket">]</span></a></sup> DNA and other nucleic acids are the basis of <a href="/wiki/Aptamers" class="mw-redirect" title="Aptamers">aptamers</a>, synthetic oligonucleotide ligands for specific target molecules used in a range of biotechnology and biomedical applications.<sup id="cite_ref-183" class="reference"><a href="#cite_note-183"><span class="cite-bracket">[</span>183<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="History_and_anthropology">History and anthropology</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Phylogenetics" title="Phylogenetics">Phylogenetics</a> and <a href="/wiki/Genetic_genealogy" title="Genetic genealogy">Genetic genealogy</a></div> <p>Because DNA collects mutations over time, which are then inherited, it contains historical information, and, by comparing DNA sequences, geneticists can infer the evolutionary history of organisms, their <a href="/wiki/Phylogenetics" title="Phylogenetics">phylogeny</a>.<sup id="cite_ref-184" class="reference"><a href="#cite_note-184"><span class="cite-bracket">[</span>184<span class="cite-bracket">]</span></a></sup> This field of phylogenetics is a powerful tool in <a href="/wiki/Evolutionary_biology" title="Evolutionary biology">evolutionary biology</a>. If DNA sequences within a species are compared, <a href="/wiki/Population_genetics" title="Population genetics">population geneticists</a> can learn the history of particular populations. This can be used in studies ranging from <a href="/wiki/Ecological_genetics" title="Ecological genetics">ecological genetics</a> to <a href="/wiki/Anthropology" title="Anthropology">anthropology</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Information_storage">Information storage</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/DNA_digital_data_storage" title="DNA digital data storage">DNA digital data storage</a></div> <p>DNA as a <a href="/wiki/Data_storage" title="Data storage">storage device</a> for information has enormous potential since it has much higher <a href="/wiki/Storage_density" class="mw-redirect" title="Storage density">storage density</a> compared to electronic devices. However, high costs, slow read and write times (<a href="/wiki/Memory_latency" title="Memory latency">memory latency</a>), and insufficient <a href="/wiki/Data_corruption" title="Data corruption">reliability</a> has prevented its practical use.<sup id="cite_ref-pmid29744271_185-0" class="reference"><a href="#cite_note-pmid29744271-185"><span class="cite-bracket">[</span>185<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid30073589_186-0" class="reference"><a href="#cite_note-pmid30073589-186"><span class="cite-bracket">[</span>186<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="History">History</h2></div> <p><span class="anchor" id="History_of_DNA_research"></span> </p> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/History_of_molecular_biology" title="History of molecular biology">History of molecular biology</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Maclyn_McCarty_with_Francis_Crick_and_James_D_Watson_-_10.1371_journal.pbio.0030341.g001-O.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/ed/Maclyn_McCarty_with_Francis_Crick_and_James_D_Watson_-_10.1371_journal.pbio.0030341.g001-O.jpg/220px-Maclyn_McCarty_with_Francis_Crick_and_James_D_Watson_-_10.1371_journal.pbio.0030341.g001-O.jpg" decoding="async" width="220" height="173" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/ed/Maclyn_McCarty_with_Francis_Crick_and_James_D_Watson_-_10.1371_journal.pbio.0030341.g001-O.jpg/330px-Maclyn_McCarty_with_Francis_Crick_and_James_D_Watson_-_10.1371_journal.pbio.0030341.g001-O.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/ed/Maclyn_McCarty_with_Francis_Crick_and_James_D_Watson_-_10.1371_journal.pbio.0030341.g001-O.jpg/440px-Maclyn_McCarty_with_Francis_Crick_and_James_D_Watson_-_10.1371_journal.pbio.0030341.g001-O.jpg 2x" data-file-width="2740" data-file-height="2153" /></a><figcaption><a href="/wiki/Maclyn_McCarty" title="Maclyn McCarty">Maclyn McCarty</a> (left) shakes hands with <a href="/wiki/Francis_Crick" title="Francis Crick">Francis Crick</a> and <a href="/wiki/James_Watson" title="James Watson">James Watson</a>, co-originators of the double-helix model based on the X-ray diffraction data and insights of <a href="/wiki/Rosalind_Franklin" title="Rosalind Franklin">Rosalind Franklin</a> and <a href="/wiki/Raymond_Gosling" title="Raymond Gosling">Raymond Gosling</a>.</figcaption></figure> <p>DNA was first isolated by the Swiss physician <a href="/wiki/Friedrich_Miescher" title="Friedrich Miescher">Friedrich Miescher</a> who, in 1869, discovered a microscopic substance in the <a href="/wiki/Pus" title="Pus">pus</a> of discarded surgical bandages. As it resided in the nuclei of cells, he called it "nuclein".<sup id="cite_ref-187" class="reference"><a href="#cite_note-187"><span class="cite-bracket">[</span>187<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-188" class="reference"><a href="#cite_note-188"><span class="cite-bracket">[</span>188<span class="cite-bracket">]</span></a></sup> In 1878, <a href="/wiki/Albrecht_Kossel" title="Albrecht Kossel">Albrecht Kossel</a> isolated the non-protein component of "nuclein", nucleic acid, and later isolated its five primary <a href="/wiki/Nucleobase" class="mw-redirect" title="Nucleobase">nucleobases</a>.<sup id="cite_ref-189" class="reference"><a href="#cite_note-189"><span class="cite-bracket">[</span>189<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Yale_Jones_1953_190-0" class="reference"><a href="#cite_note-Yale_Jones_1953-190"><span class="cite-bracket">[</span>190<span class="cite-bracket">]</span></a></sup> </p><p>In 1909, <a href="/wiki/Phoebus_Levene" title="Phoebus Levene">Phoebus Levene</a> identified the base, sugar, and phosphate nucleotide unit of RNA (then named "yeast nucleic acid").<sup id="cite_ref-191" class="reference"><a href="#cite_note-191"><span class="cite-bracket">[</span>191<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-192" class="reference"><a href="#cite_note-192"><span class="cite-bracket">[</span>192<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-193" class="reference"><a href="#cite_note-193"><span class="cite-bracket">[</span>193<span class="cite-bracket">]</span></a></sup> In 1929, Levene identified deoxyribose sugar in "thymus nucleic acid" (DNA).<sup id="cite_ref-194" class="reference"><a href="#cite_note-194"><span class="cite-bracket">[</span>194<span class="cite-bracket">]</span></a></sup> Levene suggested that DNA consisted of a string of four nucleotide units linked together through the phosphate groups ("tetranucleotide hypothesis"). Levene thought the chain was short and the bases repeated in a fixed order. In 1927, <a href="/wiki/Nikolai_Koltsov" title="Nikolai Koltsov">Nikolai Koltsov</a> proposed that inherited traits would be inherited via a "giant hereditary molecule" made up of "two mirror strands that would replicate in a semi-conservative fashion using each strand as a template".<sup id="cite_ref-195" class="reference"><a href="#cite_note-195"><span class="cite-bracket">[</span>195<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Soyfer_196-0" class="reference"><a href="#cite_note-Soyfer-196"><span class="cite-bracket">[</span>196<span class="cite-bracket">]</span></a></sup> In 1928, <a href="/wiki/Frederick_Griffith" title="Frederick Griffith">Frederick Griffith</a> in his <a href="/wiki/Griffith%27s_experiment" title="Griffith's experiment">experiment</a> discovered that <a href="/wiki/Trait_(biology)" class="mw-redirect" title="Trait (biology)">traits</a> of the "smooth" form of <i>Pneumococcus</i> could be transferred to the "rough" form of the same bacteria by mixing killed "smooth" bacteria with the live "rough" form.<sup id="cite_ref-197" class="reference"><a href="#cite_note-197"><span class="cite-bracket">[</span>197<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-198" class="reference"><a href="#cite_note-198"><span class="cite-bracket">[</span>198<span class="cite-bracket">]</span></a></sup> This system provided the first clear suggestion that DNA carries genetic information. </p><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 DNA is found in the <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-199" class="reference"><a href="#cite_note-199"><span class="cite-bracket">[</span>199<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-200" class="reference"><a href="#cite_note-200"><span class="cite-bracket">[</span>200<span class="cite-bracket">]</span></a></sup> </p><p>In 1937, <a href="/wiki/William_Astbury" title="William Astbury">William Astbury</a> produced the first X-ray diffraction patterns that showed that DNA had a regular structure.<sup id="cite_ref-201" class="reference"><a href="#cite_note-201"><span class="cite-bracket">[</span>201<span class="cite-bracket">]</span></a></sup> </p><p>In 1943, <a href="/wiki/Oswald_Avery" title="Oswald Avery">Oswald Avery</a>, along with co-workers <a href="/wiki/Colin_Munro_MacLeod" title="Colin Munro MacLeod">Colin MacLeod</a> and <a href="/wiki/Maclyn_McCarty" title="Maclyn McCarty">Maclyn McCarty</a>, identified DNA as the <a href="/wiki/Griffith%27s_experiment" title="Griffith's experiment">transforming principle</a>, supporting Griffith's suggestion (<a href="/wiki/Avery%E2%80%93MacLeod%E2%80%93McCarty_experiment" title="Avery–MacLeod–McCarty experiment">Avery–MacLeod–McCarty experiment</a>).<sup id="cite_ref-202" class="reference"><a href="#cite_note-202"><span class="cite-bracket">[</span>202<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Erwin_Chargaff" title="Erwin Chargaff">Erwin Chargaff</a> developed and published observations now known as <a href="/wiki/Chargaff%27s_rules" title="Chargaff's rules">Chargaff's rules</a>, stating that in DNA from any species of any organism, the amount of <a href="/wiki/Guanine" title="Guanine">guanine</a> should be equal to <a href="/wiki/Cytosine" title="Cytosine">cytosine</a> and the amount of <a href="/wiki/Adenine" title="Adenine">adenine</a> should be equal to <a href="/wiki/Thymine" title="Thymine">thymine</a>.<sup id="cite_ref-chargaff_1950_203-0" class="reference"><a href="#cite_note-chargaff_1950-203"><span class="cite-bracket">[</span>203<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-kresge_2005_204-0" class="reference"><a href="#cite_note-kresge_2005-204"><span class="cite-bracket">[</span>204<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:TheEaglePub-Cambridge-BluePlaque.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f1/TheEaglePub-Cambridge-BluePlaque.jpg/220px-TheEaglePub-Cambridge-BluePlaque.jpg" decoding="async" width="220" height="170" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f1/TheEaglePub-Cambridge-BluePlaque.jpg/330px-TheEaglePub-Cambridge-BluePlaque.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f1/TheEaglePub-Cambridge-BluePlaque.jpg/440px-TheEaglePub-Cambridge-BluePlaque.jpg 2x" data-file-width="1000" data-file-height="773" /></a><figcaption>A <a href="/wiki/Blue_plaque" title="Blue plaque">blue plaque</a> outside <a href="/wiki/The_Eagle,_Cambridge" title="The Eagle, Cambridge">The Eagle</a> <a href="/wiki/Pub" title="Pub">pub</a> in Cambridge, England commemorating Crick and Watson</figcaption></figure> <p>Late in 1951, <a href="/wiki/Francis_Crick" title="Francis Crick">Francis Crick</a> started working with <a href="/wiki/James_Watson" title="James Watson">James Watson</a> at the <a href="/wiki/Cavendish_Laboratory" title="Cavendish Laboratory">Cavendish Laboratory</a> within the <a href="/wiki/University_of_Cambridge" title="University of Cambridge">University of Cambridge</a>. DNA's role in <a href="/wiki/Heredity" title="Heredity">heredity</a> was confirmed in 1952 when <a href="/wiki/Alfred_Hershey" title="Alfred Hershey">Alfred Hershey</a> and <a href="/wiki/Martha_Chase" title="Martha Chase">Martha Chase</a> in the <a href="/wiki/Hershey%E2%80%93Chase_experiment" title="Hershey–Chase experiment">Hershey–Chase experiment</a> showed that DNA is the <a href="/wiki/Genetic_material" class="mw-redirect" title="Genetic material">genetic material</a> of the <a href="/wiki/Enterobacteria_phage_T2" title="Enterobacteria phage T2">enterobacteria phage T2</a>.<sup id="cite_ref-205" class="reference"><a href="#cite_note-205"><span class="cite-bracket">[</span>205<span class="cite-bracket">]</span></a></sup> </p><p>In May 1952, <a href="/wiki/Raymond_Gosling" title="Raymond Gosling">Raymond Gosling</a>, a graduate student working under the supervision of <a href="/wiki/Rosalind_Franklin" title="Rosalind Franklin">Rosalind Franklin</a>, took an <a href="/wiki/X-ray_diffraction" title="X-ray diffraction">X-ray diffraction</a> image, labeled as "<a href="/wiki/Photo_51" title="Photo 51">Photo 51</a>",<sup id="cite_ref-206" class="reference"><a href="#cite_note-206"><span class="cite-bracket">[</span>206<span class="cite-bracket">]</span></a></sup> at high hydration levels of DNA. This photo was given to Watson and Crick by <a href="/wiki/Maurice_Wilkins" title="Maurice Wilkins">Maurice Wilkins</a> and was critical to their obtaining the correct structure of DNA. Franklin told Crick and Watson that the backbones had to be on the outside. Before then, Linus Pauling, and Watson and Crick, had erroneous models with the chains inside and the bases pointing outwards. Franklin's identification of the <a href="/wiki/Space_group" title="Space group">space group</a> for DNA crystals revealed to Crick that the two DNA strands were <a href="/wiki/Antiparallel_(biochemistry)" title="Antiparallel (biochemistry)">antiparallel</a>.<sup id="cite_ref-ReferenceA_207-0" class="reference"><a href="#cite_note-ReferenceA-207"><span class="cite-bracket">[</span>207<span class="cite-bracket">]</span></a></sup> In February 1953, <a href="/wiki/Linus_Pauling" title="Linus Pauling">Linus Pauling</a> and <a href="/wiki/Robert_Corey" title="Robert Corey">Robert Corey</a> proposed a model for nucleic acids containing three intertwined chains, with the phosphates near the axis, and the bases on the outside.<sup id="cite_ref-pmid16578429_208-0" class="reference"><a href="#cite_note-pmid16578429-208"><span class="cite-bracket">[</span>208<span class="cite-bracket">]</span></a></sup> Watson and Crick completed their model, which is now accepted as the first correct model of the double helix of <a href="/wiki/Molecular_structure_of_Nucleic_Acids" class="mw-redirect" title="Molecular structure of Nucleic Acids">DNA</a>. On 28 February 1953 Crick interrupted patrons' lunchtime at <a href="/wiki/The_Eagle,_Cambridge" title="The Eagle, Cambridge">The Eagle</a> <a href="/wiki/Pub" title="Pub">pub</a> in Cambridge, England to announce that he and Watson had "discovered the secret of life".<sup id="cite_ref-209" class="reference"><a href="#cite_note-209"><span class="cite-bracket">[</span>209<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Pencil_sketch_of_the_DNA_double_helix_by_Francis_Crick_Wellcome_L0051225.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/ab/Pencil_sketch_of_the_DNA_double_helix_by_Francis_Crick_Wellcome_L0051225.jpg/170px-Pencil_sketch_of_the_DNA_double_helix_by_Francis_Crick_Wellcome_L0051225.jpg" decoding="async" width="170" height="217" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/ab/Pencil_sketch_of_the_DNA_double_helix_by_Francis_Crick_Wellcome_L0051225.jpg/255px-Pencil_sketch_of_the_DNA_double_helix_by_Francis_Crick_Wellcome_L0051225.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/ab/Pencil_sketch_of_the_DNA_double_helix_by_Francis_Crick_Wellcome_L0051225.jpg/340px-Pencil_sketch_of_the_DNA_double_helix_by_Francis_Crick_Wellcome_L0051225.jpg 2x" data-file-width="4900" data-file-height="6254" /></a><figcaption>Pencil sketch of the DNA double helix by Francis Crick in 1953</figcaption></figure> <p>The 25 April 1953 issue of the journal <i><a href="/wiki/Nature_(journal)" title="Nature (journal)">Nature</a></i> published a series of five articles giving the Watson and Crick double-helix structure DNA and evidence supporting it.<sup id="cite_ref-NatureDNA50_210-0" class="reference"><a href="#cite_note-NatureDNA50-210"><span class="cite-bracket">[</span>210<span class="cite-bracket">]</span></a></sup> The structure was reported in a letter titled "<i>MOLECULAR STRUCTURE OF NUCLEIC ACIDS A Structure for Deoxyribose Nucleic Acid</i><span style="padding-left:.15em;">"</span>, in which they said, "It has not escaped our notice that the specific pairing we have postulated immediately suggests a possible copying mechanism for the genetic material."<sup id="cite_ref-Watson-1953_9-3" class="reference"><a href="#cite_note-Watson-1953-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> This letter was followed by a letter from Franklin and Gosling, which was the first publication of their own X-ray diffraction data and of their original analysis method.<sup id="cite_ref-NatFranGos_48-1" class="reference"><a href="#cite_note-NatFranGos-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-211" class="reference"><a href="#cite_note-211"><span class="cite-bracket">[</span>211<span class="cite-bracket">]</span></a></sup> Then followed a letter by Wilkins and two of his colleagues, which contained an analysis of <i>in vivo</i> B-DNA X-ray patterns, and which supported the presence <i>in vivo</i> of the Watson and Crick structure.<sup id="cite_ref-NatWilk_49-1" class="reference"><a href="#cite_note-NatWilk-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> </p><p>In April 2023, scientists, based on new evidence, concluded that Rosalind Franklin was a contributor and "equal player" in the discovery process of DNA, rather than otherwise, as may have been presented subsequently after the time of the discovery.<sup id="cite_ref-AP-20230425_212-0" class="reference"><a href="#cite_note-AP-20230425-212"><span class="cite-bracket">[</span>212<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-NYT-20230425_213-0" class="reference"><a href="#cite_note-NYT-20230425-213"><span class="cite-bracket">[</span>213<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-NAT-202304254_214-0" class="reference"><a href="#cite_note-NAT-202304254-214"><span class="cite-bracket">[</span>214<span class="cite-bracket">]</span></a></sup> </p><p>In 1962, after Franklin's death, Watson, Crick, and Wilkins jointly received the <a href="/wiki/Nobel_Prize_in_Physiology_or_Medicine" title="Nobel Prize in Physiology or Medicine">Nobel Prize in Physiology or Medicine</a>.<sup id="cite_ref-215" class="reference"><a href="#cite_note-215"><span class="cite-bracket">[</span>215<span class="cite-bracket">]</span></a></sup> Nobel Prizes are awarded only to living recipients. A debate continues about who should receive credit for the discovery.<sup id="cite_ref-216" class="reference"><a href="#cite_note-216"><span class="cite-bracket">[</span>216<span class="cite-bracket">]</span></a></sup> </p><p>In an influential presentation in 1957, Crick laid out the <a href="/wiki/Central_dogma_of_molecular_biology" title="Central dogma of molecular biology">central dogma of molecular biology</a>, which foretold the relationship between DNA, RNA, and proteins, and articulated the "adaptor hypothesis".<sup id="cite_ref-217" class="reference"><a href="#cite_note-217"><span class="cite-bracket">[</span>217<span class="cite-bracket">]</span></a></sup> Final confirmation of the replication mechanism that was implied by the double-helical structure followed in 1958 through the <a href="/wiki/Meselson%E2%80%93Stahl_experiment" title="Meselson–Stahl experiment">Meselson–Stahl experiment</a>.<sup id="cite_ref-218" class="reference"><a href="#cite_note-218"><span class="cite-bracket">[</span>218<span class="cite-bracket">]</span></a></sup> Further work by Crick and co-workers showed that the genetic code was based on non-overlapping triplets of bases, called <a href="/wiki/Genetic_code#Codons" title="Genetic code">codons</a>, allowing <a href="/wiki/Har_Gobind_Khorana" title="Har Gobind Khorana">Har Gobind Khorana</a>, <a href="/wiki/Robert_W._Holley" title="Robert W. Holley">Robert W. Holley</a>, and <a href="/wiki/Marshall_Warren_Nirenberg" title="Marshall Warren Nirenberg">Marshall Warren Nirenberg</a> to decipher the genetic code.<sup id="cite_ref-219" class="reference"><a href="#cite_note-219"><span class="cite-bracket">[</span>219<span class="cite-bracket">]</span></a></sup> These findings represent the birth of <a href="/wiki/Molecular_biology" title="Molecular biology">molecular biology</a>.<sup id="cite_ref-220" class="reference"><a href="#cite_note-220"><span class="cite-bracket">[</span>220<span class="cite-bracket">]</span></a></sup> </p> <div style="clear:both;" class=""></div> <p>In 1986, DNA analysis was first used for criminal investigative purposes when police in the UK requested <a href="/wiki/Alec_Jeffreys" title="Alec Jeffreys">Alec Jeffreys</a> of the University of Leicester to verify or disprove a suspect's rape-murder "confession". In this particular case, the suspect had confessed to two rape-murders, but had later retracted his confession. DNA testing at the university labs soon disproved the veracity of the suspect's original "confession", and the suspect was exonerated from the murder-rape charges.<sup id="cite_ref-221" class="reference"><a href="#cite_note-221"><span class="cite-bracket">[</span>221<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2></div> <style data-mw-deduplicate="TemplateStyles:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col"> <ul><li><a href="/wiki/Autosome" title="Autosome">Autosome</a> – Any chromosome other than a sex chromosome</li> <li><a href="/wiki/Crystallography" title="Crystallography">Crystallography</a> – Scientific study of crystal structures</li> <li><a href="/wiki/DNA_Day" title="DNA Day">DNA Day</a> – Holiday celebrated on April 25</li> <li><a href="/wiki/DNA_microarray" title="DNA microarray">DNA microarray</a> – Collection of microscopic DNA spots attached to a solid surface</li> <li><a href="/wiki/DNA_sequencing" title="DNA sequencing">DNA sequencing</a> – Process of determining the nucleic acid sequence</li> <li><a href="/wiki/Genetic_disorder" title="Genetic disorder">Genetic disorder</a> – Health problem caused by one or more abnormalities in the genome</li> <li><a href="/wiki/Genetic_genealogy" title="Genetic genealogy">Genetic genealogy</a> – DNA testing to infer relationships</li> <li><a href="/wiki/Haplotype" title="Haplotype">Haplotype</a> – Group of genes from one parent</li> <li><a href="/wiki/Meiosis" title="Meiosis">Meiosis</a> – Cell division producing haploid gametes</li> <li><a href="/wiki/Nucleic_acid_notation" title="Nucleic acid notation">Nucleic acid notation</a> – Universal notation using the Roman characters A, C, G, and T to call the four DNA nucleotides</li> <li><a href="/wiki/Nucleic_acid_sequence" title="Nucleic acid sequence">Nucleic acid sequence</a> – Succession of nucleotides in a nucleic acid</li> <li><a href="/wiki/Ribosomal_DNA" title="Ribosomal DNA">Ribosomal DNA</a> – Genes coding for ribosomal RNA</li> <li><a href="/wiki/Southern_blot" title="Southern blot">Southern blot</a> – DNA analysis technique</li> <li><a href="/wiki/X-ray_scattering_techniques" title="X-ray scattering techniques">X-ray scattering techniques</a> – family of non-destructive analytical techniques<span style="display:none" class="category-wikidata-fallback-annotation">Pages displaying wikidata descriptions as a fallback</span></li> <li><a href="/wiki/Xeno_nucleic_acid" title="Xeno nucleic acid">Xeno nucleic acid</a> – Synthetic nucleic acid analogues</li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2></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"> <div class="mw-references-wrap mw-references-columns"><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"><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 class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.merriam-webster.com/dictionary/deoxyribonucleic%20acid">"deoxyribonucleic acid"</a>. <i><a href="/wiki/Merriam-Webster" title="Merriam-Webster">Merriam-Webster.com Dictionary</a></i>. Merriam-Webster.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Merriam-Webster.com+Dictionary&rft.atitle=deoxyribonucleic+acid&rft_id=https%3A%2F%2Fwww.merriam-webster.com%2Fdictionary%2Fdeoxyribonucleic%2520acid&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></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"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAlbertsJohnsonLewisRaff2014" class="citation book cs1">Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2014). <a rel="nofollow" class="external text" href="http://www.garlandscience.com/product/isbn/9780815344322"><i>Molecular Biology of the Cell</i></a> (6th ed.). Garland. p. Chapter 4: DNA, Chromosomes and Genomes. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-8153-4432-2" title="Special:BookSources/978-0-8153-4432-2"><bdi>978-0-8153-4432-2</bdi></a>. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20140714210549/http://www.garlandscience.com/product/isbn/9780815344322">Archived</a> from the original on 14 July 2014.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Molecular+Biology+of+the+Cell&rft.pages=Chapter+4%3A+DNA%2C+Chromosomes+and+Genomes&rft.edition=6th&rft.pub=Garland&rft.date=2014&rft.isbn=978-0-8153-4432-2&rft.aulast=Alberts&rft.aufirst=B&rft.au=Johnson%2C+A&rft.au=Lewis%2C+J&rft.au=Raff%2C+M&rft.au=Roberts%2C+K&rft.au=Walter%2C+P&rft_id=http%3A%2F%2Fwww.garlandscience.com%2Fproduct%2Fisbn%2F9780815344322&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" 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="CITEREFPurcell" class="citation web cs1">Purcell A. <a rel="nofollow" class="external text" href="http://basicbiology.net/micro/genetics/dna">"DNA"</a>. <i>Basic Biology</i>. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20170105045651/http://basicbiology.net/micro/genetics/dna/">Archived</a> from the original on 5 January 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Basic+Biology&rft.atitle=DNA&rft.aulast=Purcell&rft.aufirst=A&rft_id=http%3A%2F%2Fbasicbiology.net%2Fmicro%2Fgenetics%2Fdna&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" 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 class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.genome.gov/genetics-glossary/Uracil">"Uracil"</a>. <i>Genome.gov</i><span class="reference-accessdate">. Retrieved <span class="nowrap">21 November</span> 2019</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Genome.gov&rft.atitle=Uracil&rft_id=https%3A%2F%2Fwww.genome.gov%2Fgenetics-glossary%2FUracil&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></span> </li> <li id="cite_note-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-5">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRussell2001" class="citation book cs1">Russell P (2001). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/igenetics0000russ_v6o1"><i>iGenetics</i></a></span>. 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class="Z3988"></span></span> </li> <li id="cite_note-Weinberg-87"><span class="mw-cite-backlink"><b><a href="#cite_ref-Weinberg_87-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFJohnson2010" class="citation news cs1">Johnson G (28 December 2010). <a rel="nofollow" class="external text" href="https://www.nytimes.com/2010/12/28/health/28cancer.html">"Unearthing Prehistoric Tumors, and Debate"</a>. <i><a href="/wiki/The_New_York_Times" title="The New York Times">The New York Times</a></i>. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20170624233156/http://www.nytimes.com/2010/12/28/health/28cancer.html">Archived</a> from the original on 24 June 2017. <q>If we lived long enough, sooner or later we all would get cancer.</q></cite><span 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"Discovering DNA: Friedrich Miescher and the early years of nucleic acid research". <i>Human Genetics</i>. <b>122</b> (6): 565–81. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fs00439-007-0433-0">10.1007/s00439-007-0433-0</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/17901982">17901982</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:915930">915930</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Human+Genetics&rft.atitle=Discovering+DNA%3A+Friedrich+Miescher+and+the+early+years+of+nucleic+acid+research&rft.volume=122&rft.issue=6&rft.pages=565-81&rft.date=2008-01&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A915930%23id-name%3DS2CID&rft_id=info%3Apmid%2F17901982&rft_id=info%3Adoi%2F10.1007%2Fs00439-007-0433-0&rft.aulast=Dahm&rft.aufirst=R&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></span> </li> <li id="cite_note-189"><span class="mw-cite-backlink"><b><a href="#cite_ref-189">^</a></b></span> <span class="reference-text">See: <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKossel1879" class="citation journal cs1 cs1-prop-foreign-lang-source">Kossel A (1879). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=4H5NAAAAYAAJ&pg=PA284">"Ueber Nucleïn der Hefe"</a> [On nuclein in yeast]. <i>Zeitschrift für physiologische Chemie</i> (in German). <b>3</b>: 284–91.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Zeitschrift+f%C3%BCr+physiologische+Chemie&rft.atitle=Ueber+Nucle%C3%AFn+der+Hefe&rft.volume=3&rft.pages=284-91&rft.date=1879&rft.aulast=Kossel&rft.aufirst=A&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3D4H5NAAAAYAAJ%26pg%3DPA284&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKossel1880" class="citation journal cs1 cs1-prop-foreign-lang-source">Kossel A (1880). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=u4s1AQAAMAAJ&pg=PA290">"Ueber Nucleïn der Hefe II"</a> [On nuclein in yeast, Part 2]. <i>Zeitschrift für physiologische Chemie</i> (in German). <b>4</b>: 290–95.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Zeitschrift+f%C3%BCr+physiologische+Chemie&rft.atitle=Ueber+Nucle%C3%AFn+der+Hefe+II&rft.volume=4&rft.pages=290-95&rft.date=1880&rft.aulast=Kossel&rft.aufirst=A&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3Du4s1AQAAMAAJ%26pg%3DPA290&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKossel1881" class="citation journal cs1 cs1-prop-foreign-lang-source">Kossel A (1881). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=xYs1AQAAMAAJ&pg=PA267">"Ueber die Verbreitung des Hypoxanthins im Thier- und Pflanzenreich"</a> [On the distribution of hypoxanthins in the animal and plant kingdoms]. <i>Zeitschrift für physiologische Chemie</i> (in German). <b>5</b>: 267–71.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Zeitschrift+f%C3%BCr+physiologische+Chemie&rft.atitle=Ueber+die+Verbreitung+des+Hypoxanthins+im+Thier-+und+Pflanzenreich&rft.volume=5&rft.pages=267-71&rft.date=1881&rft.aulast=Kossel&rft.aufirst=A&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DxYs1AQAAMAAJ%26pg%3DPA267&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKossel1881" class="citation book cs1 cs1-prop-foreign-lang-source">Kossel A (1881). <i>Untersuchungen über die Nucleine und ihre Spaltungsprodukte</i> [<i>Investigations into nuclein and its cleavage products</i>] (in German). Strassburg, Germany: K.J. Trübner. p. 19.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Untersuchungen+%C3%BCber+die+Nucleine+und+ihre+Spaltungsprodukte&rft.place=Strassburg%2C+Germany&rft.pages=19&rft.pub=K.J.+Tr%C3%BCbner&rft.date=1881&rft.aulast=Kossel&rft.aufirst=A&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKossel1882" class="citation journal cs1">Kossel A (1882). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=z4s1AQAAMAAJ&pg=PA422">"Ueber Xanthin und Hypoxanthin"</a> [On xanthin and hypoxanthin]. <i>Zeitschrift für physiologische Chemie</i>. <b>6</b>: 422–31.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Zeitschrift+f%C3%BCr+physiologische+Chemie&rft.atitle=Ueber+Xanthin+und+Hypoxanthin&rft.volume=6&rft.pages=422-31&rft.date=1882&rft.aulast=Kossel&rft.aufirst=A&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3Dz4s1AQAAMAAJ%26pg%3DPA422&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li> <li>Albrect Kossel (1883) <a rel="nofollow" class="external text" href="https://books.google.com/books?id=2os1AQAAMAAJ&pg=PA7">"Zur Chemie des Zellkerns"</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20171117235430/https://books.google.com/books?id=2os1AQAAMAAJ&pg=PA7">Archived</a> 17 November 2017 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a> (On the chemistry of the cell nucleus), <i>Zeitschrift für physiologische Chemie</i>, <b>7</b>: 7–22.</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKossel1886" class="citation journal cs1 cs1-prop-foreign-lang-source">Kossel A (1886). <a rel="nofollow" class="external text" href="http://vlp.mpiwg-berlin.mpg.de/library/data/lit16615/index_html?pn=1&ws=1.5">"Weitere Beiträge zur Chemie des Zellkerns"</a> [Further contributions to the chemistry of the cell nucleus]. <i>Zeitschrift für Physiologische Chemie</i> (in German). <b>10</b>: 248–64. <q>On p. 264, Kossel remarked presciently: Der Erforschung der quantitativen Verhältnisse der vier stickstoffreichen Basen, der Abhängigkeit ihrer Menge von den physiologischen Zuständen der Zelle, verspricht wichtige Aufschlüsse über die elementaren physiologisch-chemischen Vorgänge. (The study of the quantitative relations of the four nitrogenous bases—[and] of the dependence of their quantity on the physiological states of the cell—promises important insights into the fundamental physiological-chemical processes.)</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Zeitschrift+f%C3%BCr+Physiologische+Chemie&rft.atitle=Weitere+Beitr%C3%A4ge+zur+Chemie+des+Zellkerns&rft.volume=10&rft.pages=248-64&rft.date=1886&rft.aulast=Kossel&rft.aufirst=A&rft_id=http%3A%2F%2Fvlp.mpiwg-berlin.mpg.de%2Flibrary%2Fdata%2Flit16615%2Findex_html%3Fpn%3D1%26ws%3D1.5&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li></ul> </span></li> <li id="cite_note-Yale_Jones_1953-190"><span class="mw-cite-backlink"><b><a href="#cite_ref-Yale_Jones_1953_190-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFJones1953" class="citation journal cs1">Jones ME (September 1953). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2599350">"Albrecht Kossel, a biographical sketch"</a>. <i>The Yale Journal of Biology and Medicine</i>. <b>26</b> (1): 80–97. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2599350">2599350</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/13103145">13103145</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Yale+Journal+of+Biology+and+Medicine&rft.atitle=Albrecht+Kossel%2C+a+biographical+sketch&rft.volume=26&rft.issue=1&rft.pages=80-97&rft.date=1953-09&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2599350%23id-name%3DPMC&rft_id=info%3Apmid%2F13103145&rft.aulast=Jones&rft.aufirst=ME&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2599350&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></span> </li> <li id="cite_note-191"><span class="mw-cite-backlink"><b><a href="#cite_ref-191">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLeveneJacobs1909" class="citation journal cs1 cs1-prop-foreign-lang-source">Levene PA, Jacobs WA (1909). <a rel="nofollow" class="external text" href="https://babel.hathitrust.org/cgi/pt?id=iau.31858002459620&view=1up&seq=1054">"Über Inosinsäure"</a>. <i>Berichte der Deutschen Chemischen Gesellschaft</i> (in German). <b>42</b>: 1198–203. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1002%2Fcber.190904201196">10.1002/cber.190904201196</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Berichte+der+Deutschen+Chemischen+Gesellschaft&rft.atitle=%C3%9Cber+Inosins%C3%A4ure&rft.volume=42&rft.pages=1198-203&rft.date=1909&rft_id=info%3Adoi%2F10.1002%2Fcber.190904201196&rft.aulast=Levene&rft.aufirst=PA&rft.au=Jacobs%2C+WA&rft_id=https%3A%2F%2Fbabel.hathitrust.org%2Fcgi%2Fpt%3Fid%3Diau.31858002459620%26view%3D1up%26seq%3D1054&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></span> </li> <li id="cite_note-192"><span class="mw-cite-backlink"><b><a href="#cite_ref-192">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLeveneJacobs1909" class="citation journal cs1 cs1-prop-foreign-lang-source">Levene PA, Jacobs WA (1909). <a rel="nofollow" class="external text" href="https://zenodo.org/record/2175598">"Über die Hefe-Nucleinsäure"</a>. <i>Berichte der Deutschen Chemischen Gesellschaft</i> (in German). <b>42</b> (2): 2474–78. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1002%2Fcber.190904202148">10.1002/cber.190904202148</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Berichte+der+Deutschen+Chemischen+Gesellschaft&rft.atitle=%C3%9Cber+die+Hefe-Nucleins%C3%A4ure&rft.volume=42&rft.issue=2&rft.pages=2474-78&rft.date=1909&rft_id=info%3Adoi%2F10.1002%2Fcber.190904202148&rft.aulast=Levene&rft.aufirst=PA&rft.au=Jacobs%2C+WA&rft_id=https%3A%2F%2Fzenodo.org%2Frecord%2F2175598&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></span> </li> <li id="cite_note-193"><span class="mw-cite-backlink"><b><a href="#cite_ref-193">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLevene1919" class="citation journal cs1">Levene P (1919). <a rel="nofollow" class="external text" href="https://doi.org/10.1016%2FS0021-9258%2818%2987254-4">"The structure of yeast nucleic acid"</a>. <i>J Biol Chem</i>. <b>40</b> (2): 415–24. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1016%2FS0021-9258%2818%2987254-4">10.1016/S0021-9258(18)87254-4</a></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=J+Biol+Chem&rft.atitle=The+structure+of+yeast+nucleic+acid&rft.volume=40&rft.issue=2&rft.pages=415-24&rft.date=1919&rft_id=info%3Adoi%2F10.1016%2FS0021-9258%2818%2987254-4&rft.aulast=Levene&rft.aufirst=P&rft_id=https%3A%2F%2Fdoi.org%2F10.1016%252FS0021-9258%252818%252987254-4&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></span> </li> <li id="cite_note-194"><span class="mw-cite-backlink"><b><a href="#cite_ref-194">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCohenPortugal1974" class="citation journal cs1">Cohen JS, Portugal FH (1974). <a rel="nofollow" class="external text" href="https://profiles.nlm.nih.gov/ps/access/CCAAHW.pdf">"The search for the chemical structure of DNA"</a> <span class="cs1-format">(PDF)</span>. <i>Connecticut Medicine</i>. <b>38</b> (10): 551–52, 554–57. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/4609088">4609088</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Connecticut+Medicine&rft.atitle=The+search+for+the+chemical+structure+of+DNA&rft.volume=38&rft.issue=10&rft.pages=551-52%2C+554-57&rft.date=1974&rft_id=info%3Apmid%2F4609088&rft.aulast=Cohen&rft.aufirst=JS&rft.au=Portugal%2C+FH&rft_id=https%3A%2F%2Fprofiles.nlm.nih.gov%2Fps%2Faccess%2FCCAAHW.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></span> </li> <li id="cite_note-195"><span class="mw-cite-backlink"><b><a href="#cite_ref-195">^</a></b></span> <span class="reference-text">Koltsov proposed that a cell's genetic information was encoded in a long chain of amino acids. See: <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKoltsov1927" class="citation speech cs1 cs1-prop-foreign-lang-source">Koltsov HK (12 December 1927). <i>Физико-химические основы морфологии</i> [<i>The physical-chemical basis of morphology</i>] (Speech). 3rd All-Union Meeting of Zoologist, Anatomists, and Histologists (in Russian). Leningrad, U.S.S.R.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=%D0%A4%D0%B8%D0%B7%D0%B8%D0%BA%D0%BE-%D1%85%D0%B8%D0%BC%D0%B8%D1%87%D0%B5%D1%81%D0%BA%D0%B8%D0%B5+%D0%BE%D1%81%D0%BD%D0%BE%D0%B2%D1%8B+%D0%BC%D0%BE%D1%80%D1%84%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D0%B8&rft.place=Leningrad%2C+U.S.S.R.&rft.date=1927-12-12&rft.aulast=Koltsov&rft.aufirst=HK&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li> <li>Reprinted in: <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKoltsov1928" class="citation journal cs1 cs1-prop-foreign-lang-source">Koltsov HK (1928). "Физико-химические основы морфологии" [The physical-chemical basis of morphology]. <i>Успехи экспериментальной биологии (Advances in Experimental Biology) series B</i> (in Russian). <b>7</b> (1): ?.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=%D0%A3%D1%81%D0%BF%D0%B5%D1%85%D0%B8+%D1%8D%D0%BA%D1%81%D0%BF%D0%B5%D1%80%D0%B8%D0%BC%D0%B5%D0%BD%D1%82%D0%B0%D0%BB%D1%8C%D0%BD%D0%BE%D0%B9+%D0%B1%D0%B8%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D0%B8+%28Advances+in+Experimental+Biology%29+series+B&rft.atitle=%D0%A4%D0%B8%D0%B7%D0%B8%D0%BA%D0%BE-%D1%85%D0%B8%D0%BC%D0%B8%D1%87%D0%B5%D1%81%D0%BA%D0%B8%D0%B5+%D0%BE%D1%81%D0%BD%D0%BE%D0%B2%D1%8B+%D0%BC%D0%BE%D1%80%D1%84%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D0%B8&rft.volume=7&rft.issue=1&rft.pages=%3F&rft.date=1928&rft.aulast=Koltsov&rft.aufirst=HK&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li> <li>Reprinted in German as: <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKoltzoff1928" class="citation journal cs1 cs1-prop-foreign-lang-source">Koltzoff NK (1928). "Physikalisch-chemische Grundlagen der Morphologie" [The physical-chemical basis of morphology]. <i>Biologisches Zentralblatt</i> (in German). <b>48</b> (6): 345–69.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Biologisches+Zentralblatt&rft.atitle=Physikalisch-chemische+Grundlagen+der+Morphologie&rft.volume=48&rft.issue=6&rft.pages=345-69&rft.date=1928&rft.aulast=Koltzoff&rft.aufirst=NK&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li> <li>In 1934, Koltsov contended that the proteins that contain a cell's genetic information replicate. See: <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKoltzoff1934" class="citation journal cs1">Koltzoff N (October 1934). "The structure of the chromosomes in the salivary glands of Drosophila". <i>Science</i>. <b>80</b> (2075): 312–13. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1934Sci....80..312K">1934Sci....80..312K</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscience.80.2075.312">10.1126/science.80.2075.312</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/17769043">17769043</a>. <q>From page 313: "I think that the size of the chromosomes in the salivary glands [of Drosophila] is determined through the multiplication of <i>genonemes</i>. By this term I designate the axial thread of the chromosome, in which the geneticists locate the linear combination of genes; … In the normal chromosome there is usually only one genoneme; before cell-division this genoneme has become divided into two strands."<span class="cs1-kern-right"></span></q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Science&rft.atitle=The+structure+of+the+chromosomes+in+the+salivary+glands+of+Drosophila&rft.volume=80&rft.issue=2075&rft.pages=312-13&rft.date=1934-10&rft_id=info%3Apmid%2F17769043&rft_id=info%3Adoi%2F10.1126%2Fscience.80.2075.312&rft_id=info%3Abibcode%2F1934Sci....80..312K&rft.aulast=Koltzoff&rft.aufirst=N&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li></ul> </span></li> <li id="cite_note-Soyfer-196"><span class="mw-cite-backlink"><b><a href="#cite_ref-Soyfer_196-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSoyfer2001" class="citation journal cs1">Soyfer VN (September 2001). "The consequences of political dictatorship for Russian science". <i>Nature Reviews Genetics</i>. <b>2</b> (9): 723–29. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2F35088598">10.1038/35088598</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11533721">11533721</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:46277758">46277758</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature+Reviews+Genetics&rft.atitle=The+consequences+of+political+dictatorship+for+Russian+science&rft.volume=2&rft.issue=9&rft.pages=723-29&rft.date=2001-09&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A46277758%23id-name%3DS2CID&rft_id=info%3Apmid%2F11533721&rft_id=info%3Adoi%2F10.1038%2F35088598&rft.aulast=Soyfer&rft.aufirst=VN&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></span> </li> <li id="cite_note-197"><span class="mw-cite-backlink"><b><a href="#cite_ref-197">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGriffith1928" class="citation journal cs1">Griffith F (January 1928). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2167760">"The Significance of Pneumococcal Types"</a>. <i>The Journal of Hygiene</i>. <b>27</b> (2): 113–59. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1017%2FS0022172400031879">10.1017/S0022172400031879</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2167760">2167760</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/20474956">20474956</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Journal+of+Hygiene&rft.atitle=The+Significance+of+Pneumococcal+Types&rft.volume=27&rft.issue=2&rft.pages=113-59&rft.date=1928-01&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2167760%23id-name%3DPMC&rft_id=info%3Apmid%2F20474956&rft_id=info%3Adoi%2F10.1017%2FS0022172400031879&rft.aulast=Griffith&rft.aufirst=F&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2167760&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></span> </li> <li id="cite_note-198"><span class="mw-cite-backlink"><b><a href="#cite_ref-198">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLorenzWackernagel1994" class="citation journal cs1">Lorenz MG, Wackernagel W (September 1994). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC372978">"Bacterial gene transfer by natural genetic transformation in the environment"</a>. <i>Microbiological Reviews</i>. <b>58</b> (3): 563–602. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1128%2FMMBR.58.3.563-602.1994">10.1128/MMBR.58.3.563-602.1994</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC372978">372978</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/7968924">7968924</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Microbiological+Reviews&rft.atitle=Bacterial+gene+transfer+by+natural+genetic+transformation+in+the+environment&rft.volume=58&rft.issue=3&rft.pages=563-602&rft.date=1994-09&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC372978%23id-name%3DPMC&rft_id=info%3Apmid%2F7968924&rft_id=info%3Adoi%2F10.1128%2FMMBR.58.3.563-602.1994&rft.aulast=Lorenz&rft.aufirst=MG&rft.au=Wackernagel%2C+W&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC372978&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></span> </li> <li id="cite_note-199"><span class="mw-cite-backlink"><b><a href="#cite_ref-199">^</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". <i>Archives de Biologie</i> (in Italian). <b>44</b>: 519–76.</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=519-76&rft.date=1933&rft.aulast=Brachet&rft.aufirst=J&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></span> </li> <li id="cite_note-200"><span class="mw-cite-backlink"><b><a href="#cite_ref-200">^</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 JF (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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Retrieved <span class="nowrap">26 April</span> 2023</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+New+York+Times&rft.atitle=Untangling+Rosalind+Franklin%27s+Role+in+DNA+Discovery%2C+70+Years+On+%E2%80%93+Historians+have+long+debated+the+role+that+Dr.+Franklin+played+in+identifying+the+double+helix.+A+new+opinion+essay+argues+that+she+was+an+%22equal+contributor.%22&rft.date=2023-04-25&rft.aulast=Anthes&rft.aufirst=E&rft_id=https%3A%2F%2Fwww.nytimes.com%2F2023%2F04%2F25%2Fscience%2Frosalind-franklin-dna.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></span> </li> <li id="cite_note-NAT-202304254-214"><span class="mw-cite-backlink"><b><a href="#cite_ref-NAT-202304254_214-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCobbComfort2023" class="citation journal cs1">Cobb M, Comfort N (25 April 2023). <a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fd41586-023-01313-5">"What Rosalind Franklin truly contributed to the discovery of DNA's structure – Franklin was no victim in how the DNA double helix was solved. An overlooked letter and an unpublished news article, both written in 1953, reveal that she was an equal player"</a>. <i><a href="/wiki/Nature_(journal)" title="Nature (journal)">Nature</a></i>. <b>616</b> (7958): 657–660. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2023Natur.616..657C">2023Natur.616..657C</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fd41586-023-01313-5">10.1038/d41586-023-01313-5</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/37100935">37100935</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID 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class="Z3988"></span></span> </li> <li id="cite_note-215"><span class="mw-cite-backlink"><b><a href="#cite_ref-215">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://nobelprize.org/nobel_prizes/medicine/laureates/1962/">"The Nobel Prize in Physiology or Medicine 1962"</a>. <i>Nobelprize.org</i>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Nobelprize.org&rft.atitle=The+Nobel+Prize+in+Physiology+or+Medicine+1962&rft_id=http%3A%2F%2Fnobelprize.org%2Fnobel_prizes%2Fmedicine%2Flaureates%2F1962%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></span> </li> <li id="cite_note-216"><span class="mw-cite-backlink"><b><a href="#cite_ref-216">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" 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class="external text" href="https://pubmed.ncbi.nlm.nih.gov/12540909">12540909</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:4428347">4428347</a>. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20161017011403/http://www.biomath.nyu.edu/index/course/hw_articles/nature4.pdf">Archived</a> <span class="cs1-format">(PDF)</span> from the original on 17 October 2016.</cite><span 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href="https://web.archive.org/web/20081001223217/http://genome.wellcome.ac.uk/assets/wtx030893.pdf"><i>A Note for the RNA Tie Club</i></a> <span class="cs1-format">(PDF)</span> (Speech). 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"Discovery of DNA structure and function: Watson and Crick". <i>Nature Education</i>. <b>1</b> (1): 100.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature+Education&rft.atitle=Discovery+of+DNA+structure+and+function%3A+Watson+and+Crick.&rft.volume=1&rft.issue=1&rft.pages=100&rft.date=2008&rft.aulast=Pray&rft.aufirst=L&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></span> </li> <li id="cite_note-221"><span class="mw-cite-backlink"><b><a href="#cite_ref-221">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPanneerchelvamNorazmi2003" class="citation journal cs1">Panneerchelvam S, Norazmi MN (2003). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3561883">"Forensic DNA Profiling and Database"</a>. <i>The Malaysian Journal of Medical Sciences</i>. <b>10</b> (2): 20–26. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3561883">3561883</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/23386793">23386793</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Malaysian+Journal+of+Medical+Sciences&rft.atitle=Forensic+DNA+Profiling+and+Database&rft.volume=10&rft.issue=2&rft.pages=20-26&rft.date=2003&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3561883%23id-name%3DPMC&rft_id=info%3Apmid%2F23386793&rft.aulast=Panneerchelvam&rft.aufirst=S&rft.au=Norazmi%2C+MN&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3561883&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></span> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2></div> <style data-mw-deduplicate="TemplateStyles:r1239549316">.mw-parser-output .refbegin{margin-bottom:0.5em}.mw-parser-output .refbegin-hanging-indents>ul{margin-left:0}.mw-parser-output .refbegin-hanging-indents>ul>li{margin-left:0;padding-left:3.2em;text-indent:-3.2em}.mw-parser-output .refbegin-hanging-indents ul,.mw-parser-output .refbegin-hanging-indents ul li{list-style:none}@media(max-width:720px){.mw-parser-output .refbegin-hanging-indents>ul>li{padding-left:1.6em;text-indent:-1.6em}}.mw-parser-output .refbegin-columns{margin-top:0.3em}.mw-parser-output .refbegin-columns ul{margin-top:0}.mw-parser-output .refbegin-columns li{page-break-inside:avoid;break-inside:avoid-column}@media screen{.mw-parser-output .refbegin{font-size:90%}}</style><div class="refbegin" style=""> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBerryWatson2003" class="citation book cs1">Berry A, <a href="/wiki/James_Watson" title="James Watson">Watson J</a> (2003). <a rel="nofollow" class="external text" href="https://archive.org/details/dnasecretoflife00wats"><i>DNA: the secret of life</i></a>. New York: Alfred A. Knopf. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-375-41546-7" title="Special:BookSources/0-375-41546-7"><bdi>0-375-41546-7</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=DNA%3A+the+secret+of+life&rft.place=New+York&rft.pub=Alfred+A.+Knopf&rft.date=2003&rft.isbn=0-375-41546-7&rft.aulast=Berry&rft.aufirst=A&rft.au=Watson%2C+J&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fdnasecretoflife00wats&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCalladineDrewLuisiTravers2003" class="citation book cs1">Calladine CR, Drew HR, Luisi BF, Travers AA (2003). <i>Understanding DNA: the molecule & how it works</i>. Amsterdam: Elsevier Academic Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-12-155089-3" title="Special:BookSources/0-12-155089-3"><bdi>0-12-155089-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Understanding+DNA%3A+the+molecule+%26+how+it+works&rft.place=Amsterdam&rft.pub=Elsevier+Academic+Press&rft.date=2003&rft.isbn=0-12-155089-3&rft.aulast=Calladine&rft.aufirst=CR&rft.au=Drew%2C+HR&rft.au=Luisi%2C+BF&rft.au=Travers%2C+AA&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCarinaClayton2003" class="citation book cs1">Carina D, Clayton J (2003). <a rel="nofollow" class="external text" href="https://archive.org/details/50yearsofdna00clay"><i>50 years of DNA</i></a>. Basingstoke: Palgrave Macmillan. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/1-4039-1479-6" title="Special:BookSources/1-4039-1479-6"><bdi>1-4039-1479-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=50+years+of+DNA&rft.place=Basingstoke&rft.pub=Palgrave+Macmillan&rft.date=2003&rft.isbn=1-4039-1479-6&rft.aulast=Carina&rft.aufirst=D&rft.au=Clayton%2C+J&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2F50yearsofdna00clay&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFJudson1979" class="citation book cs1"><a href="/wiki/Horace_Freeland_Judson" title="Horace Freeland Judson">Judson HF</a> (1979). <i>The Eighth Day of Creation: Makers of the Revolution in Biology</i> (2nd ed.). Cold Spring Harbor Laboratory Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-671-22540-5" title="Special:BookSources/0-671-22540-5"><bdi>0-671-22540-5</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Eighth+Day+of+Creation%3A+Makers+of+the+Revolution+in+Biology&rft.edition=2nd&rft.pub=Cold+Spring+Harbor+Laboratory+Press&rft.date=1979&rft.isbn=0-671-22540-5&rft.aulast=Judson&rft.aufirst=HF&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFOlby1994" class="citation book cs1"><a href="/wiki/Robert_Olby" title="Robert Olby">Olby RC</a> (1994). <i>The path to the double helix: the discovery of DNA</i>. New York: Dover Publications. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-486-68117-3" title="Special:BookSources/0-486-68117-3"><bdi>0-486-68117-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+path+to+the+double+helix%3A+the+discovery+of+DNA&rft.place=New+York&rft.pub=Dover+Publications&rft.date=1994&rft.isbn=0-486-68117-3&rft.aulast=Olby&rft.aufirst=RC&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span> First published in October 1974 by MacMillan, with foreword by Francis Crick; the definitive DNA textbook, revised in 1994 with a nine-page postscript.</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFOlby2003" class="citation journal cs1"><a href="/wiki/Robert_Olby" title="Robert Olby">Olby R</a> (January 2003). <a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fnature01397">"Quiet debut for the double helix"</a>. <i>Nature</i>. <b>421</b> (6921): 402–05. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2003Natur.421..402O">2003Natur.421..402O</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fnature01397">10.1038/nature01397</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/12540907">12540907</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=Quiet+debut+for+the+double+helix&rft.volume=421&rft.issue=6921&rft.pages=402-05&rft.date=2003-01&rft_id=info%3Apmid%2F12540907&rft_id=info%3Adoi%2F10.1038%2Fnature01397&rft_id=info%3Abibcode%2F2003Natur.421..402O&rft.aulast=Olby&rft.aufirst=R&rft_id=https%3A%2F%2Fdoi.org%2F10.1038%252Fnature01397&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFOlby2009" class="citation book cs1">Olby RC (2009). <i>Francis Crick: A Biography</i>. Plainview, N.Y: Cold Spring Harbor Laboratory Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-87969-798-3" title="Special:BookSources/978-0-87969-798-3"><bdi>978-0-87969-798-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Francis+Crick%3A+A+Biography&rft.place=Plainview%2C+N.Y&rft.pub=Cold+Spring+Harbor+Laboratory+Press&rft.date=2009&rft.isbn=978-0-87969-798-3&rft.aulast=Olby&rft.aufirst=RC&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMicklas2003" class="citation book cs1">Micklas D (2003). <i>DNA Science: A First Course</i>. Cold Spring Harbor Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-87969-636-8" title="Special:BookSources/978-0-87969-636-8"><bdi>978-0-87969-636-8</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=DNA+Science%3A+A+First+Course&rft.pub=Cold+Spring+Harbor+Press&rft.date=2003&rft.isbn=978-0-87969-636-8&rft.aulast=Micklas&rft.aufirst=D&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRidley2006" class="citation book cs1"><a href="/wiki/Matt_Ridley" title="Matt Ridley">Ridley M</a> (2006). <i>Francis Crick: discoverer of the genetic code</i>. Ashland, OH: Eminent Lives, Atlas Books. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-06-082333-X" title="Special:BookSources/0-06-082333-X"><bdi>0-06-082333-X</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Francis+Crick%3A+discoverer+of+the+genetic+code&rft.place=Ashland%2C+OH&rft.pub=Eminent+Lives%2C+Atlas+Books&rft.date=2006&rft.isbn=0-06-082333-X&rft.aulast=Ridley&rft.aufirst=M&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRosenfeld2010" class="citation book cs1">Rosenfeld I (2010). <i>DNA: A Graphic Guide to the Molecule that Shook the World</i>. Columbia University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-231-14271-7" title="Special:BookSources/978-0-231-14271-7"><bdi>978-0-231-14271-7</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=DNA%3A+A+Graphic+Guide+to+the+Molecule+that+Shook+the+World&rft.pub=Columbia+University+Press&rft.date=2010&rft.isbn=978-0-231-14271-7&rft.aulast=Rosenfeld&rft.aufirst=I&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSchultzCannon2009" class="citation book cs1">Schultz M, Cannon Z (2009). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/stuffoflifegraph00schu"><i>The Stuff of Life: A Graphic Guide to Genetics and DNA</i></a></span>. Hill and Wang. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-8090-8947-5" title="Special:BookSources/978-0-8090-8947-5"><bdi>978-0-8090-8947-5</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Stuff+of+Life%3A+A+Graphic+Guide+to+Genetics+and+DNA&rft.pub=Hill+and+Wang&rft.date=2009&rft.isbn=978-0-8090-8947-5&rft.aulast=Schultz&rft.aufirst=M&rft.au=Cannon%2C+Z&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fstuffoflifegraph00schu&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFStentWatson1980" class="citation book cs1"><a href="/wiki/Gunther_Stent" title="Gunther Stent">Stent GS</a>, Watson J (1980). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/doublehelixpers00wats_0"><i>The Double Helix: A Personal Account of the Discovery of the Structure of DNA</i></a></span>. New York: Norton. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-393-95075-1" title="Special:BookSources/0-393-95075-1"><bdi>0-393-95075-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Double+Helix%3A+A+Personal+Account+of+the+Discovery+of+the+Structure+of+DNA&rft.place=New+York&rft.pub=Norton&rft.date=1980&rft.isbn=0-393-95075-1&rft.aulast=Stent&rft.aufirst=GS&rft.au=Watson%2C+J&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fdoublehelixpers00wats_0&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWatson2004" class="citation book cs1">Watson J (2004). <i>DNA: The Secret of Life</i>. Random House. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-09-945184-6" title="Special:BookSources/978-0-09-945184-6"><bdi>978-0-09-945184-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=DNA%3A+The+Secret+of+Life&rft.pub=Random+House&rft.date=2004&rft.isbn=978-0-09-945184-6&rft.aulast=Watson&rft.aufirst=J&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWilkins2003" class="citation book cs1"><a href="/wiki/Maurice_Wilkins" title="Maurice Wilkins">Wilkins M</a> (2003). <i>The third man of the double helix the autobiography of Maurice Wilkins</i>. Cambridge, England: University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-19-860665-6" title="Special:BookSources/0-19-860665-6"><bdi>0-19-860665-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+third+man+of+the+double+helix+the+autobiography+of+Maurice+Wilkins&rft.place=Cambridge%2C+England&rft.pub=University+Press&rft.date=2003&rft.isbn=0-19-860665-6&rft.aulast=Wilkins&rft.aufirst=M&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADNA" class="Z3988"></span></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2></div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output .side-box-text{flex:1;min-width:0}}@media(min-width:720px){.mw-parser-output .side-box{width:238px}.mw-parser-output .side-box-right{clear:right;float:right;margin-left:1em}.mw-parser-output .side-box-left{margin-right:1em}}</style><div class="side-box metadata side-box-right"><style data-mw-deduplicate="TemplateStyles:r1126788409">.mw-parser-output .plainlist ol,.mw-parser-output .plainlist ul{line-height:inherit;list-style:none;margin:0;padding:0}.mw-parser-output .plainlist ol li,.mw-parser-output .plainlist ul li{margin-bottom:0}</style> <div class="side-box-abovebelow"> <a href="/wiki/Wikipedia:The_Wikipedia_Library" title="Wikipedia:The Wikipedia Library">Library resources</a> about <br /> <b>DNA</b> <hr /></div> <div class="side-box-flex"> <div class="side-box-text plainlist"><ul><li><a class="external text" href="https://ftl.toolforge.org/cgi-bin/ftl?st=&su=DNA&library=OLBP">Online books</a></li> <li><a class="external text" href="https://ftl.toolforge.org/cgi-bin/ftl?st=&su=DNA">Resources in your library</a></li> <li><a class="external text" href="https://ftl.toolforge.org/cgi-bin/ftl?st=&su=DNA&library=0CHOOSE0">Resources in other libraries</a></li> </ul></div></div> </div> <style data-mw-deduplicate="TemplateStyles:r1235611614">.mw-parser-output .spoken-wikipedia{border:1px solid #a2a9b1;background-color:var(--background-color-interactive-subtle,#f8f9fa);margin:0.5em 0;padding:0.2em;line-height:1.5em;font-size:90%}.mw-parser-output .spoken-wikipedia-header{text-align:center}.mw-parser-output .spoken-wikipedia-listen-to{font-weight:bold}.mw-parser-output .spoken-wikipedia-files{text-align:center;margin-top:10px;margin-bottom:0.4em}.mw-parser-output .spoken-wikipedia-icon{float:left;margin-left:5px;margin-top:10px}.mw-parser-output .spoken-wikipedia-disclaimer{margin-left:60px;margin-top:10px;font-size:95%;line-height:1.4em}.mw-parser-output .spoken-wikipedia-footer{margin-top:10px;text-align:center}@media(min-width:720px){.mw-parser-output .spoken-wikipedia{width:20em;float:right;clear:right;margin-left:1em}}</style><div class="spoken-wikipedia noprint haudio"><div class="spoken-wikipedia-header"><span class="spoken-wikipedia-listen-to">Listen to this article</span> (<span class="duration"><span class="min">34</span> minutes</span>)</div><div class="spoken-wikipedia-files"><figure class="mw-halign-center" typeof="mw:File"><span><audio id="mwe_player_0" controls="" preload="none" data-mw-tmh="" class="mw-file-element" width="200" style="width:200px;" data-durationhint="2032" data-mwtitle="Dna.ogg" data-mwprovider="wikimediacommons"><source src="//upload.wikimedia.org/wikipedia/commons/b/bc/Dna.ogg" type="audio/ogg; codecs="vorbis"" data-width="0" data-height="0" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/b/bc/Dna.ogg/Dna.ogg.mp3" type="audio/mpeg" data-transcodekey="mp3" data-width="0" data-height="0" /></audio></span><figcaption></figcaption></figure> </div><div class="spoken-wikipedia-icon"><span typeof="mw:File"><span title="Spoken Wikipedia"><img alt="Spoken Wikipedia icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/4/47/Sound-icon.svg/45px-Sound-icon.svg.png" decoding="async" width="45" height="34" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/47/Sound-icon.svg/68px-Sound-icon.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/47/Sound-icon.svg/90px-Sound-icon.svg.png 2x" data-file-width="128" data-file-height="96" /></span></span></div><div class="spoken-wikipedia-disclaimer"><a href="/wiki/File:Dna.ogg" title="File:Dna.ogg">This audio file</a> was created from a revision of this article dated 12 February 2007<span style="display:none"> (<span class="bday dtstart published updated itvstart">2007-02-12</span>)</span>, and does not reflect subsequent edits.</div><div class="spoken-wikipedia-footer">(<a href="/wiki/Wikipedia:Media_help" class="mw-redirect" title="Wikipedia:Media help">Audio help</a> · <a href="/wiki/Wikipedia:Spoken_articles" title="Wikipedia:Spoken articles">More spoken articles</a>)</div></div> <ul><li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20070306082905/http://pipe.scs.fsu.edu/displar.html">DNA binding site prediction on protein</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20100223035803/http://nobelprize.org/educational_games/medicine/dna_double_helix/">DNA the Double Helix Game</a> From the official Nobel Prize web site</li> <li><a rel="nofollow" class="external text" href="http://www.fidelitysystems.com/Unlinked_DNA.html">DNA under electron microscope</a></li> <li><a rel="nofollow" class="external text" href="http://www.dnalc.org/">Dolan DNA Learning Center</a></li> <li><a rel="nofollow" class="external text" href="http://www.nature.com/nature/dna50/archive.html">Double Helix: 50 years of DNA</a>, <i><a href="/wiki/Nature_(journal)" title="Nature (journal)">Nature</a></i></li> <li><i><a href="/wiki/Proteopedia" title="Proteopedia">Proteopedia</a> <a rel="nofollow" class="external text" href="http://www.proteopedia.org/wiki/index.php/DNA">DNA</a></i></li> <li><i><a href="/wiki/Proteopedia" title="Proteopedia">Proteopedia</a> <a rel="nofollow" class="external text" href="http://www.proteopedia.org/wiki/index.php/Forms_of_DNA">Forms_of_DNA</a></i></li> <li><a rel="nofollow" class="external text" href="http://www.nature.com/encode/">ENCODE threads explorer</a> ENCODE home page at <a href="/wiki/Nature_(journal)" title="Nature (journal)">Nature</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20070213030135/http://www.ncbe.reading.ac.uk/DNA50/">Double Helix 1953–2003</a> National Centre for Biotechnology Education</li> <li><a rel="nofollow" class="external text" href="http://www.genome.gov/10506718">Genetic Education Modules for Teachers</a> – <i>DNA from the Beginning</i> Study Guide</li> <li><a href="/wiki/Protein_Data_Bank" title="Protein Data Bank">PDB</a> Molecule of the Month <i><a rel="nofollow" class="external text" href="https://pdb101.rcsb.org/motm/23">DNA</a></i></li> <li><a rel="nofollow" class="external text" href="https://www.nytimes.com/packages/pdf/science/dna-article.pdf">"Clue to chemistry of heredity found"</a>. <i><a href="/wiki/The_New_York_Times" title="The New York Times">The New York Times</a></i>, June 1953. First American newspaper coverage of the discovery of the DNA structure</li> <li><a rel="nofollow" class="external text" href="http://www.dnaftb.org/">DNA from the Beginning</a> Another DNA Learning Center site on DNA, genes, and heredity from Mendel to the human genome project.</li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20070825101712/http://orpheus.ucsd.edu/speccoll/testing/html/mss0660a.html#abstract">The Register of Francis Crick Personal Papers 1938 – 2007</a> at Mandeville Special Collections Library, <a href="/wiki/University_of_California,_San_Diego" title="University of California, San Diego">University of California, San Diego</a></li> <li><a rel="nofollow" class="external text" href="http://www.nature.com/polopoly_fs/7.9746!/file/Crick%20letter%20to%20Michael.pdf">Seven-page, handwritten letter that Crick sent to his 12-year-old son Michael in 1953 describing the structure of DNA.</a> See <a rel="nofollow" class="external text" href="http://www.nature.com/news/crick-s-medal-goes-under-the-hammer-1.12705">Crick's medal goes under the hammer</a>, Nature, 5 April 2013.</li></ul> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><style data-mw-deduplicate="TemplateStyles:r1236075235">.mw-parser-output .navbox{box-sizing:border-box;border:1px solid #a2a9b1;width:100%;clear:both;font-size:88%;text-align:center;padding:1px;margin:1em auto 0}.mw-parser-output .navbox .navbox{margin-top:0}.mw-parser-output .navbox+.navbox,.mw-parser-output .navbox+.navbox-styles+.navbox{margin-top:-1px}.mw-parser-output .navbox-inner,.mw-parser-output .navbox-subgroup{width:100%}.mw-parser-output .navbox-group,.mw-parser-output .navbox-title,.mw-parser-output .navbox-abovebelow{padding:0.25em 1em;line-height:1.5em;text-align:center}.mw-parser-output .navbox-group{white-space:nowrap;text-align:right}.mw-parser-output .navbox,.mw-parser-output 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dl,.mw-parser-output .navbox td.hlist ol,.mw-parser-output .navbox td.hlist ul{padding:0.125em 0}.mw-parser-output .navbox .navbar{display:block;font-size:100%}.mw-parser-output .navbox-title .navbar{float:left;text-align:left;margin-right:0.5em}body.skin--responsive .mw-parser-output .navbox-image img{max-width:none!important}@media print{body.ns-0 .mw-parser-output .navbox{display:none!important}}</style></div><div role="navigation" class="navbox" aria-labelledby="Genetics" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Genetics" title="Template:Genetics"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Genetics" title="Template talk:Genetics"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Genetics" title="Special:EditPage/Template:Genetics"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Genetics" style="font-size:114%;margin:0 4em"><a href="/wiki/Genetics" title="Genetics">Genetics</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/Introduction_to_genetics" title="Introduction to genetics">Introduction</a></li> <li><a href="/wiki/Outline_of_genetics" title="Outline of genetics">Outline</a></li> <li><a href="/wiki/History_of_genetics" title="History of genetics">History</a></li> <li><a href="/wiki/Timeline_of_the_history_of_genetics" title="Timeline of the history of genetics">Timeline</a></li> <li><a href="/wiki/Index_of_genetics_articles" title="Index of genetics articles">Index</a></li> <li><a href="/wiki/Glossary_of_genetics" class="mw-redirect" title="Glossary of genetics">Glossary</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Key components</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Chromosome" title="Chromosome">Chromosome</a></li> <li><a class="mw-selflink selflink">DNA</a></li> <li><a href="/wiki/RNA" title="RNA">RNA</a></li> <li><a href="/wiki/Genome" title="Genome">Genome</a></li> <li><a href="/wiki/Heredity" title="Heredity">Heredity</a></li> <li><a href="/wiki/Nucleotide" title="Nucleotide">Nucleotide</a></li> <li><a href="/wiki/Mutation" title="Mutation">Mutation</a></li> <li><a href="/wiki/Genetic_variation" title="Genetic variation">Genetic variation</a></li> <li><a href="/wiki/Allele" title="Allele">Allele</a></li> <li><a href="/wiki/Amino_acid" title="Amino acid">Amino acid</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Fields</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Classical_genetics" title="Classical genetics">Classical</a></li> <li><a href="/wiki/Conservation_genetics" title="Conservation genetics">Conservation</a></li> <li><a href="/wiki/Cytogenetics" title="Cytogenetics">Cytogenetics</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological</a></li> <li><a href="/wiki/Immunogenetics" title="Immunogenetics">Immunogenetics</a></li> <li><a href="/wiki/Microbial_genetics" title="Microbial genetics">Microbial</a></li> <li><a href="/wiki/Molecular_genetics" title="Molecular genetics">Molecular</a></li> <li><a href="/wiki/Population_genetics" title="Population genetics">Population</a></li> <li><a href="/wiki/Quantitative_genetics" title="Quantitative genetics">Quantitative</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Archaeogenetics" title="Archaeogenetics">Archaeogenetics</a> of</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Genetic_history_of_Africa" title="Genetic history of Africa">Africa</a></li> <li><a href="/wiki/Genetic_history_of_indigenous_peoples_of_the_Americas" class="mw-redirect" title="Genetic history of indigenous peoples of the Americas">the Americas</a></li> <li><a href="/wiki/Genetic_history_of_the_British_Isles" title="Genetic history of the British Isles">the British Isles</a></li> <li><a href="/wiki/Genetic_history_of_Europe" title="Genetic history of Europe">Europe</a></li> <li><a href="/wiki/Genetic_history_of_Italy" title="Genetic history of Italy">Italy</a></li> <li><a href="/wiki/Genetic_history_of_the_Middle_East" title="Genetic history of the Middle East">the Middle East</a></li> <li><a href="/wiki/Genetics_and_archaeogenetics_of_South_Asia" title="Genetics and archaeogenetics of South Asia">South Asia</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related topics</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Behavioural_genetics" title="Behavioural genetics">Behavioural genetics</a></li> <li><a href="/wiki/Epigenetics" title="Epigenetics">Epigenetics</a></li> <li><a href="/wiki/Geneticist" title="Geneticist">Geneticist</a></li> <li><a href="/wiki/Genome_editing" title="Genome editing">Genome editing</a></li> <li><a href="/wiki/Genomics" title="Genomics">Genomics</a></li> <li><a href="/wiki/Genetic_code" title="Genetic code">Genetic code</a></li> <li><a href="/wiki/Genetic_engineering" title="Genetic engineering">Genetic engineering</a></li> <li><a href="/wiki/Genetic_diversity" title="Genetic diversity">Genetic diversity</a></li> <li><a href="/wiki/Genetic_monitoring" title="Genetic monitoring">Genetic monitoring</a></li> <li><a href="/wiki/Genetic_genealogy" title="Genetic genealogy">Genetic genealogy</a></li> <li><a href="/wiki/Heredity" title="Heredity">Heredity</a></li> <li><a href="/wiki/He_Jiankui_affair" title="He Jiankui affair">He Jiankui affair</a></li> <li><a href="/wiki/Medical_genetics" title="Medical genetics">Medical genetics</a></li> <li><a href="/wiki/Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li> <li><a href="/wiki/Molecular_evolution" title="Molecular evolution">Molecular evolution</a></li> <li><a href="/wiki/Plant_genetics" title="Plant genetics">Plant genetics</a></li> <li><a href="/wiki/Population_genomics" title="Population genomics">Population genomics</a></li> <li><a href="/wiki/Reverse_genetics" title="Reverse genetics">Reverse genetics</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Lists</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/List_of_genetic_codes" title="List of genetic codes">List of genetic codes</a></li> <li><a href="/wiki/List_of_genetics_research_organizations" title="List of genetics research organizations">List of genetics research organizations</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Genetics" title="Category:Genetics">Category</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Gene_expression" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Gene_expression" title="Template:Gene expression"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Gene_expression" title="Template talk:Gene expression"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Gene_expression" title="Special:EditPage/Template:Gene expression"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Gene_expression" style="font-size:114%;margin:0 4em"><a href="/wiki/Gene_expression" title="Gene expression">Gene expression</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Introduction_to_genetics" title="Introduction to genetics">Introduction<br />to genetics</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Genetic_code" title="Genetic code">Genetic code</a></li> <li><a href="/wiki/Central_dogma_of_molecular_biology" title="Central dogma of molecular biology">Central dogma</a> <ul><li><a class="mw-selflink selflink">DNA</a> → <a href="/wiki/Messenger_RNA" title="Messenger RNA">RNA</a> → <a href="/wiki/Protein" title="Protein">Protein</a></li></ul></li> <li><a href="/wiki/Central_dogma_of_molecular_biology#Special_transfers_of_biological_sequential_information" title="Central dogma of molecular biology">Special transfers</a> <ul><li><a href="/wiki/RNA-dependent_RNA_polymerase" title="RNA-dependent RNA polymerase">RNA→RNA</a></li> <li><a href="/wiki/Reverse_transcription" class="mw-redirect" title="Reverse transcription">RNA→DNA</a></li> <li><a href="/wiki/Prion" title="Prion">Protein→Protein</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Transcription_(biology)" title="Transcription (biology)">Transcription</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;">Types</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bacterial_transcription" title="Bacterial transcription">Bacterial</a></li> <li><a href="/wiki/Archaeal_transcription" title="Archaeal transcription">Archaeal</a></li> <li><a href="/wiki/Eukaryotic_transcription" title="Eukaryotic transcription">Eukaryotic</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;">Key elements</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Transcription_factor" title="Transcription factor">Transcription factor</a></li> <li><a href="/wiki/RNA_polymerase" title="RNA polymerase">RNA polymerase</a></li> <li><a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">Promoter</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;"><a href="/wiki/Post-transcriptional_modification" title="Post-transcriptional modification">Post-transcription</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Primary_transcript" title="Primary transcript">Precursor mRNA (pre-mRNA / hnRNA)</a></li> <li><a href="/wiki/Five-prime_cap" title="Five-prime cap">5' capping</a></li> <li><a href="/wiki/RNA_splicing" title="RNA splicing">Splicing</a></li> <li><a href="/wiki/Polyadenylation" title="Polyadenylation">Polyadenylation</a></li> <li><a href="/wiki/Histone_acetylation_and_deacetylation" title="Histone acetylation and deacetylation">Histone acetylation and deacetylation</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Translation_(biology)" title="Translation (biology)">Translation</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;">Types</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bacterial_translation" title="Bacterial translation">Bacterial</a></li> <li><a href="/wiki/Archaeal_translation" title="Archaeal translation">Archaeal</a></li> <li><a href="/wiki/Eukaryotic_translation" title="Eukaryotic translation">Eukaryotic</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;">Key elements</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0;white-space:nowrap;"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Ribosome" title="Ribosome">Ribosome</a></li> <li><a href="/wiki/Transfer_RNA" title="Transfer RNA">Transfer RNA (tRNA)</a></li> <li><a href="/wiki/Ribosome-nascent_chain_complex" title="Ribosome-nascent chain complex">Ribosome-nascent chain complex (RNC)</a></li> <li><a href="/wiki/Post-translational_modification" title="Post-translational modification">Post-translational modification</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Regulation_of_gene_expression" title="Regulation of gene expression">Regulation</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Epigenetics" title="Epigenetics">Epigenetic</a> <ul><li><a href="/wiki/Genomic_imprinting" title="Genomic imprinting">imprinting</a></li></ul></li> <li><a href="/wiki/Transcriptional_regulation" title="Transcriptional regulation">Transcriptional</a> <ul><li><a href="/wiki/Gene_regulatory_network" title="Gene regulatory network">Gene regulatory network</a></li> <li><a href="/wiki/Cis-regulatory_element" title="Cis-regulatory element">cis-regulatory element</a></li></ul></li> <li><a href="/wiki/Lac_operon" title="Lac operon">lac operon</a></li> <li><a href="/wiki/Post-transcriptional_regulation" title="Post-transcriptional regulation">Post-transcriptional</a> <ul><li><a href="/wiki/P-bodies" title="P-bodies">sequestration (P-bodies)</a></li> <li><a href="/wiki/Alternative_splicing" title="Alternative splicing">alternative splicing</a></li> <li><a href="/wiki/MicroRNA" title="MicroRNA">microRNA</a></li></ul></li> <li><a href="/wiki/Translational_regulation" title="Translational regulation">Translational</a></li> <li><a href="/wiki/Post-translational_regulation" title="Post-translational regulation">Post-translational</a> <ul><li><a href="/wiki/Phosphorylation" title="Phosphorylation">reversible</a></li> <li><a href="/wiki/Proteolysis" title="Proteolysis">irreversible</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Influential people</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Fran%C3%A7ois_Jacob" title="François Jacob">François Jacob</a></li> <li><a href="/wiki/Jacques_Monod" title="Jacques Monod">Jacques Monod</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"><style data-mw-deduplicate="TemplateStyles:r886047488">.mw-parser-output .nobold{font-weight:normal}</style></div><div role="navigation" class="navbox" aria-labelledby="Types_of_nucleic_acids" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2" style="text-align: center;"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Nucleic_acids" title="Template:Nucleic acids"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Nucleic_acids" title="Template talk:Nucleic acids"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Nucleic_acids" title="Special:EditPage/Template:Nucleic acids"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Types_of_nucleic_acids" style="font-size:114%;margin:0 4em">Types of <a href="/wiki/Nucleic_acid" title="Nucleic acid">nucleic acids </a></div></th></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%">Constituents</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Nucleobase" class="mw-redirect" title="Nucleobase">Nucleobases</a></li> <li><a href="/wiki/Nucleoside" title="Nucleoside">Nucleosides</a></li> <li><a href="/wiki/Nucleotide" title="Nucleotide">Nucleotides</a></li> <li><a href="/wiki/Deoxyribonucleotide" title="Deoxyribonucleotide">Deoxynucleotides</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/RNA" title="RNA">Ribonucleic acids </a><br /><span class="nobold">(coding, <a href="/wiki/Non-coding_RNA" title="Non-coding RNA">non-coding</a>)</span></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;"><a href="/wiki/RNA#In_translation" title="RNA">Translational</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Messenger_RNA" title="Messenger RNA">Messenger </a> <ul><li><a href="/wiki/Primary_transcript" title="Primary transcript">precursor, heterogenous nuclear</a></li></ul></li> <li><a href="/wiki/Nucleoside-modified_messenger_RNA" title="Nucleoside-modified messenger RNA">modified Messenger</a></li> <li><a href="/wiki/Transfer_RNA" title="Transfer RNA">Transfer </a></li> <li><a href="/wiki/Ribosomal_RNA" title="Ribosomal RNA">Ribosomal </a></li> <li><a href="/wiki/Transfer-messenger_RNA" title="Transfer-messenger RNA">Transfer-messenger </a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;"><a href="/wiki/RNA#Regulatory_RNAs" title="RNA">Regulatory</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/RNA_interference" title="RNA interference">Interferential </a> <ul><li><a href="/wiki/MicroRNA" title="MicroRNA">Micro </a></li> <li><a href="/wiki/Small_interfering_RNA" title="Small interfering RNA">Small interfering </a></li> <li><a href="/wiki/Piwi-interacting_RNA" title="Piwi-interacting RNA">Piwi-interacting</a></li></ul></li> <li><a href="/wiki/Antisense_RNA" title="Antisense RNA">Antisense </a></li> <li><a href="/wiki/RNA#In_RNA_processing" title="RNA">Processual</a> <ul><li><a href="/wiki/Small_nuclear_RNA" title="Small nuclear RNA">Small nuclear </a></li> <li><a href="/wiki/Small_nucleolar_RNA" title="Small nucleolar RNA">Small nucleolar </a></li> <li><a href="/wiki/Small_Cajal_body-specific_RNA" title="Small Cajal body-specific RNA">Small Cajal Body RNAs</a></li> <li><a href="/wiki/Y_RNA" title="Y RNA">Y RNA</a></li></ul></li> <li><a href="/wiki/Enhancer_RNA" title="Enhancer RNA">Enhancer RNAs </a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;">Others</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Guide_RNA" title="Guide RNA">Guide </a></li> <li><a href="/wiki/Ribozyme" title="Ribozyme">Ribozyme</a></li> <li><a href="/wiki/Small_hairpin_RNA" class="mw-redirect" title="Small hairpin RNA">Small hairpin </a></li> <li><a href="/wiki/Small_temporal_RNA" title="Small temporal RNA">Small temporal </a></li> <li><a href="/wiki/Trans-acting_siRNA" title="Trans-acting siRNA">Trans-acting small interfering</a></li> <li><a href="/wiki/Subgenomic_mRNA" title="Subgenomic mRNA">Subgenomic messenger</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a class="mw-selflink selflink">Deoxyribonucleic <br />acids </a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Organellar_DNA" title="Organellar DNA">Organellar</a> <ul><li><a href="/wiki/Chloroplast_DNA" title="Chloroplast DNA">Chloroplast </a></li> <li><a href="/wiki/Mitochondrial_DNA" title="Mitochondrial DNA">Mitochondrial</a></li></ul></li> <li><a href="/wiki/Complementary_DNA" title="Complementary DNA">Complementary </a></li> <li><a href="/wiki/Deoxyribozyme" title="Deoxyribozyme">Deoxyribozyme</a></li> <li><a href="/wiki/Genomic_DNA" title="Genomic DNA">Genomic </a></li> <li><a href="/wiki/Hachimoji_DNA" title="Hachimoji DNA">Hachimoji</a></li> <li><a href="/wiki/Multicopy_single-stranded_DNA" title="Multicopy single-stranded DNA">Multicopy single-stranded </a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Nucleic_acid_analogue" title="Nucleic acid analogue">Analogues</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Xeno_nucleic_acid" title="Xeno nucleic acid">Xeno </a> <ul><li><a href="/wiki/Glycol_nucleic_acid" title="Glycol nucleic acid">Glycol </a></li> <li><a href="/wiki/Threose_nucleic_acid" title="Threose nucleic acid">Threose </a></li> <li><a href="/wiki/Hexose" title="Hexose">Hexose</a></li></ul></li> <li><a href="/wiki/Locked_nucleic_acid" title="Locked nucleic acid">Locked </a></li> <li><a href="/wiki/Peptide_nucleic_acid" title="Peptide nucleic acid">Peptide </a></li> <li><a href="/wiki/Morpholino" title="Morpholino">Morpholino</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Cloning_vector" title="Cloning vector">Cloning vectors</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Phagemid" title="Phagemid">Phagemid</a></li> <li><a href="/wiki/Plasmid" title="Plasmid">Plasmid</a></li> <li><a href="/wiki/Lambda_phage" title="Lambda phage">Lambda phage</a></li> <li><a href="/wiki/Cosmid" title="Cosmid">Cosmid</a></li> <li><a href="/wiki/Fosmid" title="Fosmid">Fosmid</a></li> <li><a href="/wiki/Human_artificial_chromosome" title="Human artificial chromosome">Artificial chromosomes</a> <ul><li><a href="/wiki/P1-derived_artificial_chromosome" title="P1-derived artificial chromosome">P1-derived </a></li> <li><a href="/wiki/Bacterial_artificial_chromosome" title="Bacterial artificial chromosome">Bacterial </a></li> <li><a href="/wiki/Yeast_artificial_chromosome" title="Yeast artificial chromosome">Yeast </a></li> <li><a href="/wiki/Human_artificial_chromosome" title="Human artificial chromosome">Human</a></li></ul></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="text-align: center;"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Nucleic_acids" title="Category:Nucleic acids">Category</a></li></ul> </div></td></tr></tbody></table></div> <style data-mw-deduplicate="TemplateStyles:r1130092004">.mw-parser-output .portal-bar{font-size:88%;font-weight:bold;display:flex;justify-content:center;align-items:baseline}.mw-parser-output .portal-bar-bordered{padding:0 2em;background-color:#fdfdfd;border:1px solid #a2a9b1;clear:both;margin:1em auto 0}.mw-parser-output .portal-bar-related{font-size:100%;justify-content:flex-start}.mw-parser-output .portal-bar-unbordered{padding:0 1.7em;margin-left:0}.mw-parser-output .portal-bar-header{margin:0 1em 0 0.5em;flex:0 0 auto;min-height:24px}.mw-parser-output .portal-bar-content{display:flex;flex-flow:row wrap;flex:0 1 auto;padding:0.15em 0;column-gap:1em;align-items:baseline;margin:0;list-style:none}.mw-parser-output .portal-bar-content-related{margin:0;list-style:none}.mw-parser-output .portal-bar-item{display:inline-block;margin:0.15em 0.2em;min-height:24px;line-height:24px}@media screen and (max-width:768px){.mw-parser-output .portal-bar{font-size:88%;font-weight:bold;display:flex;flex-flow:column wrap;align-items:baseline}.mw-parser-output .portal-bar-header{text-align:center;flex:0;padding-left:0.5em;margin:0 auto}.mw-parser-output .portal-bar-related{font-size:100%;align-items:flex-start}.mw-parser-output .portal-bar-content{display:flex;flex-flow:row wrap;align-items:center;flex:0;column-gap:1em;border-top:1px solid #a2a9b1;margin:0 auto;list-style:none}.mw-parser-output .portal-bar-content-related{border-top:none;margin:0;list-style:none}}.mw-parser-output .navbox+link+.portal-bar,.mw-parser-output .navbox+style+.portal-bar,.mw-parser-output .navbox+link+.portal-bar-bordered,.mw-parser-output .navbox+style+.portal-bar-bordered,.mw-parser-output .sister-bar+link+.portal-bar,.mw-parser-output .sister-bar+style+.portal-bar,.mw-parser-output .portal-bar+.navbox-styles+.navbox,.mw-parser-output .portal-bar+.navbox-styles+.sister-bar{margin-top:-1px}</style><div class="portal-bar noprint metadata noviewer portal-bar-bordered" role="navigation" aria-label="Portals"><span class="portal-bar-header"><a href="/wiki/Wikipedia:Contents/Portals" title="Wikipedia:Contents/Portals">Portal</a>:</span><ul class="portal-bar-content"><li class="portal-bar-item"><span class="nowrap"><span typeof="mw:File"><a href="/wiki/File:Issoria_lathonia.jpg" class="mw-file-description"><img alt="icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Issoria_lathonia.jpg/21px-Issoria_lathonia.jpg" decoding="async" width="21" height="15" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Issoria_lathonia.jpg/32px-Issoria_lathonia.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Issoria_lathonia.jpg/42px-Issoria_lathonia.jpg 2x" data-file-width="629" data-file-height="445" /></a></span> </span><a href="/wiki/Portal:Biology" title="Portal:Biology">Biology</a></li></ul></div><style data-mw-deduplicate="TemplateStyles:r1236088147">.mw-parser-output .sister-bar{display:flex;justify-content:center;align-items:baseline;font-size:88%;background-color:#fdfdfd;border:1px solid #a2a9b1;clear:both;margin:1em 0 0;padding:0 2em}.mw-parser-output .sister-bar-header{margin:0 1em 0 0.5em;padding:0.2em 0;flex:0 0 auto;min-height:24px;line-height:22px}.mw-parser-output .sister-bar-content{display:flex;flex-flow:row wrap;flex:0 1 auto;align-items:baseline;padding:0.2em 0;column-gap:1em;margin:0;list-style:none}.mw-parser-output .sister-bar-item{display:flex;align-items:baseline;margin:0.15em 0;min-height:24px;text-align:left}.mw-parser-output .sister-bar-logo{width:22px;line-height:22px;margin:0 0.2em;text-align:right}.mw-parser-output .sister-bar-link{margin:0 0.2em;text-align:left}@media screen and (max-width:960px){.mw-parser-output .sister-bar{flex-flow:column wrap;margin:1em auto 0}.mw-parser-output .sister-bar-header{flex:0 1}.mw-parser-output .sister-bar-content{flex:1;border-top:1px solid #a2a9b1;margin:0;list-style:none}.mw-parser-output .sister-bar-item{flex:0 0 20em;min-width:20em}}.mw-parser-output .navbox+link+.sister-bar,.mw-parser-output .navbox+style+.sister-bar,.mw-parser-output .portal-bar+link+.sister-bar,.mw-parser-output .portal-bar+style+.sister-bar,.mw-parser-output .sister-bar+.navbox-styles+.navbox,.mw-parser-output .sister-bar+.navbox-styles+.portal-bar{margin-top:-1px}@media print{body.ns-0 .mw-parser-output .sister-bar{display:none!important}}</style><div class="noprint metadata sister-bar" role="navigation" aria-label="sister-projects"><div class="sister-bar-header"><b>DNA</b> at Wikipedia's <a href="/wiki/Wikipedia:Wikimedia_sister_projects" title="Wikipedia:Wikimedia sister projects"><span id="sister-projects" style="white-space:nowrap;">sister projects</span></a>:</div><ul class="sister-bar-content"><li class="sister-bar-item"><span class="sister-bar-logo"><span typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/14px-Commons-logo.svg.png" decoding="async" width="14" height="19" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/21px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/28px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></span></span></span><span class="sister-bar-link"><b><a href="https://commons.wikimedia.org/wiki/DNA" class="extiw" title="c:DNA">Media</a></b> from Commons</span></li><li class="sister-bar-item"><span class="sister-bar-logo"><span typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Wikiquote-logo.svg/16px-Wikiquote-logo.svg.png" decoding="async" width="16" height="19" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Wikiquote-logo.svg/24px-Wikiquote-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Wikiquote-logo.svg/32px-Wikiquote-logo.svg.png 2x" data-file-width="300" data-file-height="355" /></span></span></span><span class="sister-bar-link"><b><a href="https://en.wikiquote.org/wiki/DNA" class="extiw" title="q:DNA">Quotations</a></b> from Wikiquote</span></li><li class="sister-bar-item"><span class="sister-bar-logo"><span typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0b/Wikiversity_logo_2017.svg/21px-Wikiversity_logo_2017.svg.png" decoding="async" width="21" height="17" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0b/Wikiversity_logo_2017.svg/32px-Wikiversity_logo_2017.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0b/Wikiversity_logo_2017.svg/42px-Wikiversity_logo_2017.svg.png 2x" data-file-width="626" data-file-height="512" /></span></span></span><span class="sister-bar-link"><b><a href="https://en.wikiversity.org/wiki/DNA" class="extiw" title="v:DNA">Resources</a></b> from Wikiversity</span></li><li class="sister-bar-item"><span class="sister-bar-logo"><span typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/f/ff/Wikidata-logo.svg/21px-Wikidata-logo.svg.png" decoding="async" width="21" height="12" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/ff/Wikidata-logo.svg/32px-Wikidata-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/ff/Wikidata-logo.svg/42px-Wikidata-logo.svg.png 2x" data-file-width="1050" data-file-height="590" /></span></span></span><span class="sister-bar-link"><b><a href="https://www.wikidata.org/wiki/Q7430" class="extiw" title="d:Q7430">Data</a></b> from Wikidata</span></li></ul></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"><style data-mw-deduplicate="TemplateStyles:r1038841319">.mw-parser-output .tooltip-dotted{border-bottom:1px dotted;cursor:help}</style></div><div role="navigation" class="navbox authority-control" aria-labelledby="Authority_control_databases_frameless&#124;text-top&#124;10px&#124;alt=Edit_this_at_Wikidata&#124;link=https&#58;//www.wikidata.org/wiki/Q7430#identifiers&#124;class=noprint&#124;Edit_this_at_Wikidata" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Authority_control_databases_frameless&#124;text-top&#124;10px&#124;alt=Edit_this_at_Wikidata&#124;link=https&#58;//www.wikidata.org/wiki/Q7430#identifiers&#124;class=noprint&#124;Edit_this_at_Wikidata" style="font-size:114%;margin:0 4em"><a href="/wiki/Help:Authority_control" title="Help:Authority control">Authority control databases</a> <span class="mw-valign-text-top noprint" typeof="mw:File/Frameless"><a href="https://www.wikidata.org/wiki/Q7430#identifiers" title="Edit this at Wikidata"><img alt="Edit this at Wikidata" src="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/10px-OOjs_UI_icon_edit-ltr-progressive.svg.png" decoding="async" width="10" height="10" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/15px-OOjs_UI_icon_edit-ltr-progressive.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/20px-OOjs_UI_icon_edit-ltr-progressive.svg.png 2x" data-file-width="20" data-file-height="20" /></a></span></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">International</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"><ul><li><span class="uid"><a rel="nofollow" class="external text" href="http://id.worldcat.org/fast/886555/">FAST</a></span></li></ul></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">National</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"><ul><li><span class="uid"><a rel="nofollow" class="external text" href="https://d-nb.info/gnd/4070512-2">Germany</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://id.loc.gov/authorities/sh85037008">United States</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://catalogue.bnf.fr/ark:/12148/cb119649178">France</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://data.bnf.fr/ark:/12148/cb119649178">BnF data</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://id.ndl.go.jp/auth/ndlna/00561484">Japan</a></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="DNA (nukleová kyselina)"><a rel="nofollow" class="external text" href="https://aleph.nkp.cz/F/?func=find-c&local_base=aut&ccl_term=ica=ph116977&CON_LNG=ENG">Czech Republic</a></span></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="http://catalogo.bne.es/uhtbin/authoritybrowse.cgi?action=display&authority_id=XX527602">Spain</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="http://olduli.nli.org.il/F/?func=find-b&local_base=NLX10&find_code=UID&request=987007548275805171">Israel</a></span></li></ul></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"><ul><li><span class="uid"><a rel="nofollow" class="external text" href="http://esu.com.ua/search_articles.php?id=21681">Encyclopedia of Modern Ukraine</a></span></li></ul></div></td></tr></tbody></table></div> <!-- NewPP limit report Parsed by mw‐web.codfw.main‐f69cdc8f6‐rmdbt Cached time: 20241122140340 Cache expiry: 2592000 Reduced expiry: false Complications: [vary‐revision‐sha1, show‐toc] CPU time usage: 3.195 seconds Real time usage: 3.692 seconds Preprocessor visited node count: 16971/1000000 Post‐expand include size: 685589/2097152 bytes Template argument size: 6155/2097152 bytes Highest expansion depth: 16/100 Expensive parser function count: 50/500 Unstrip recursion depth: 1/20 Unstrip post‐expand size: 998520/5000000 bytes Lua time usage: 2.142/10.000 seconds Lua memory usage: 24351482/52428800 bytes Lua Profile: ? 240 ms 11.2% MediaWiki\Extension\Scribunto\Engines\LuaSandbox\LuaSandboxCallback::callParserFunction 220 ms 10.3% MediaWiki\Extension\Scribunto\Engines\LuaSandbox\LuaSandboxCallback::find 180 ms 8.4% recursiveClone <mwInit.lua:45> 160 ms 7.5% <Module:Citation/CS1:813> 140 ms 6.5% 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