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<p>Genes often contain regions that do not encode products, but <a class="mw-redirect" title="Gene regulation" href="http://en.wikipedia.org/wiki/Gene_regulation">regulate gene expression</a>. The genes of <a title="Eukaryote" href="http://en.wikipedia.org/wiki/Eukaryote">eukaryotic</a> organisms can contain regions called <a title="Intron" href="http://en.wikipedia.org/wiki/Intron">introns</a> that are removed from the messenger RNA in a process called <a title="Splicing (genetics)" href="http://en.wikipedia.org/wiki/Splicing_%28genetics%29">splicing</a>. The regions encoding gene products are called <a title="Exon" href="http://en.wikipedia.org/wiki/Exon">exons</a>. In eukaryotes, a single gene can encode multiple proteins, which are produced through the creation of different arrangements of exons through <em>alternative splicing</em>. In <a title="Prokaryote" href="http://en.wikipedia.org/wiki/Prokaryote">prokaryotes</a> (<a title="Bacteria" href="http://en.wikipedia.org/wiki/Bacteria">bacteria</a> and <a title="Archaea" href="http://en.wikipedia.org/wiki/Archaea">archaea</a>), introns are less common and genes often contain a single uninterrupted stretch of DNA, called a <em>cistron</em>, that codes for a product. Prokaryotic genes are often arranged in groups called <a class="mw-redirect" title="Operons" href="http://en.wikipedia.org/wiki/Operons">operons</a> with <a title="Promoter" href="http://en.wikipedia.org/wiki/Promoter">promoter</a> and <a title="Operator" href="http://en.wikipedia.org/wiki/Operator">operator</a> sequences that regulate <a title="Transcription (genetics)" href="http://en.wikipedia.org/wiki/Transcription_%28genetics%29">transcription</a> of a single long <a title="RNA" href="http://en.wikipedia.org/wiki/RNA">RNA</a>. This RNA contains multiple coding sequences. Each coding sequence is preceded by a <a title="Shine-Dalgarno sequence" href="http://en.wikipedia.org/wiki/Shine-Dalgarno_sequence">Shine-Dalgarno sequence</a> that ribosomes recognize.</p>
<p>The total set of genes in an organism is known as its <a title="Genome" href="http://en.wikipedia.org/wiki/Genome">genome</a>. An organism's <a title="Genome size" href="http://en.wikipedia.org/wiki/Genome_size">genome size</a> is generally lower in <a title="Prokaryote" href="http://en.wikipedia.org/wiki/Prokaryote">prokaryotes</a>, both in number of <a title="Base pair" href="http://en.wikipedia.org/wiki/Base_pair">base pairs</a> and number of genes, than even single-celled <a title="Eukaryote" href="http://en.wikipedia.org/wiki/Eukaryote">eukaryotes</a>. However, there is no clear relationship between genome sizes and complexity in eukaryotic organisms. One of the largest known genomes belongs to the single-celled <a title="Amoeba" href="http://en.wikipedia.org/wiki/Amoeba">amoeba</a> <em>Amoeba dubia</em>, with over 670 billion base pairs, some 200 times larger than the human genome.<sup class="reference" id="_ref-Cavalier-Smith_0"><a title="" href="http://en.wikipedia.org/wiki/Gene#_note-Cavalier-Smith">[5]</a></sup> The estimated number of genes in the <a title="Human genome" href="http://en.wikipedia.org/wiki/Human_genome">human genome</a> has been repeatedly revised downward since the completion of the <a title="Human Genome Project" href="http://en.wikipedia.org/wiki/Human_Genome_Project">Human Genome Project</a>; current estimates place the human genome at just under 3 billion base pairs and about 20,000&ndash;25,000 genes.<sup class="reference" id="_ref-IHSGC2004_0"><a title="" href="http://en.wikipedia.org/wiki/Gene#_note-IHSGC2004">[6]</a></sup> A recent <em><a title="Science (journal)" href="http://en.wikipedia.org/wiki/Science_%28journal%29">Science</a></em> article gives a final number of 20,488, with perhaps 100 more yet to be discovered .<sup class="reference" id="_ref-gene_count2007_0"><a title="" href="http://en.wikipedia.org/wiki/Gene#_note-gene_count2007">[7]</a></sup> The gene density of a genome is a measure of the number of genes per million base pairs (called a megabase, Mb); prokaryotic genomes have much higher gene densities than eukaryotes. The gene density of the human genome is roughly 12&ndash;15 genes/Mb.<sup class="reference" id="_ref-Watson_2004_0"><a title="" href="http://en.wikipedia.org/wiki/Gene#_note-Watson_2004">[8]</a></sup></p>
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<h2>Contents</h2>
<span class="toctoggle">[<a class="internal" id="togglelink" href="javascript:toggleToc()">hide</a>]</span></div>
<ul>
<li class="toclevel-1"><a href="http://en.wikipedia.org/wiki/Gene#History"><span class="tocnumber">1</span> <span class="toctext">History</span></a> </li>
<li class="toclevel-1"><a href="http://en.wikipedia.org/wiki/Gene#Mendelian_inheritance_and_classical_genetics"><span class="tocnumber">2</span> <span class="toctext">Mendelian inheritance and classical genetics</span></a> </li>
<li class="toclevel-1"><a href="http://en.wikipedia.org/wiki/Gene#Physical_definitions"><span class="tocnumber">3</span> <span class="toctext">Physical definitions</span></a>
<ul>
<li class="toclevel-2"><a href="http://en.wikipedia.org/wiki/Gene#RNA_genes"><span class="tocnumber">3.1</span> <span class="toctext">RNA genes</span></a> </li>
<li class="toclevel-2"><a href="http://en.wikipedia.org/wiki/Gene#Functional_structure_of_a_gene"><span class="tocnumber">3.2</span> <span class="toctext">Functional structure of a gene</span></a> </li>
<li class="toclevel-2"><a href="http://en.wikipedia.org/wiki/Gene#Chromosomes"><span class="tocnumber">3.3</span> <span class="toctext">Chromosomes</span></a> </li>
</ul>
</li>
<li class="toclevel-1"><a href="http://en.wikipedia.org/wiki/Gene#Gene_expression"><span class="tocnumber">4</span> <span class="toctext">Gene expression</span></a>
<ul>
<li class="toclevel-2"><a href="http://en.wikipedia.org/wiki/Gene#Genetic_code"><span class="tocnumber">4.1</span> <span class="toctext">Genetic code</span></a> </li>
<li class="toclevel-2"><a href="http://en.wikipedia.org/wiki/Gene#Transcription"><span class="tocnumber">4.2</span> <span class="toctext">Transcription</span></a> </li>
<li class="toclevel-2"><a href="http://en.wikipedia.org/wiki/Gene#Translation"><span class="tocnumber">4.3</span> <span class="toctext">Translation</span></a> </li>
</ul>
</li>
<li class="toclevel-1"><a href="http://en.wikipedia.org/wiki/Gene#DNA_replication_and_inheritance"><span class="tocnumber">5</span> <span class="toctext">DNA replication and inheritance</span></a>
<ul>
<li class="toclevel-2"><a href="http://en.wikipedia.org/wiki/Gene#Molecular_inheritance"><span class="tocnumber">5.1</span> <span class="toctext">Molecular inheritance</span></a> </li>
<li class="toclevel-2"><a href="http://en.wikipedia.org/wiki/Gene#Mutation"><span class="tocnumber">5.2</span> <span class="toctext">Mutation</span></a> </li>
</ul>
</li>
<li class="toclevel-1"><a href="http://en.wikipedia.org/wiki/Gene#The_genome"><span class="tocnumber">6</span> <span class="toctext">The genome</span></a>
<ul>
<li class="toclevel-2"><a href="http://en.wikipedia.org/wiki/Gene#Chromosomal_organization"><span class="tocnumber">6.1</span> <span class="toctext">Chromosomal organization</span></a> </li>
<li class="toclevel-2"><a href="http://en.wikipedia.org/wiki/Gene#Composition_of_the_genome"><span class="tocnumber">6.2</span> <span class="toctext">Composition of the genome</span></a> </li>
<li class="toclevel-2"><a href="http://en.wikipedia.org/wiki/Gene#Genetic_and_genomic_nomenclature"><span class="tocnumber">6.3</span> <span class="toctext">Genetic and genomic nomenclature</span></a> </li>
</ul>
</li>
<li class="toclevel-1"><a href="http://en.wikipedia.org/wiki/Gene#Evolutionary_concept_of_a_gene"><span class="tocnumber">7</span> <span class="toctext">Evolutionary concept of a gene</span></a> </li>
<li class="toclevel-1"><a href="http://en.wikipedia.org/wiki/Gene#Gene_targeting_and_implications"><span class="tocnumber">8</span> <span class="toctext">Gene targeting and implications</span></a> </li>
<li class="toclevel-1"><a href="http://en.wikipedia.org/wiki/Gene#The_gene_concept_is_still_changing"><span class="tocnumber">9</span> <span class="toctext">The gene concept is still changing</span></a> </li>
<li class="toclevel-1"><a href="http://en.wikipedia.org/wiki/Gene#See_also"><span class="tocnumber">10</span> <span class="toctext">See also</span></a> </li>
<li class="toclevel-1"><a href="http://en.wikipedia.org/wiki/Gene#References"><span class="tocnumber">11</span> <span class="toctext">References</span></a> </li>
<li class="toclevel-1"><a href="http://en.wikipedia.org/wiki/Gene#Further_reading"><span class="tocnumber">12</span> <span class="toctext">Further reading</span></a> </li>
<li class="toclevel-1"><a href="http://en.wikipedia.org/wiki/Gene#External_links"><span class="tocnumber">13</span> <span class="toctext">External links</span></a>
<ul>
<li class="toclevel-2"><a href="http://en.wikipedia.org/wiki/Gene#Tutorial_and_news"><span class="tocnumber">13.1</span> <span class="toctext">Tutorial and news</span></a> </li>
<li class="toclevel-2"><a href="http://en.wikipedia.org/wiki/Gene#References_and_databases"><span class="tocnumber">13.2</span> <span class="toctext">References and databases</span></a> </li>
</ul>
</li>
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<p><a id="History" name="History"></a></p>
<h2><span class="editsection">[<a title="Edit section: History" href="http://en.wikipedia.org/w/index.php?title=Gene&amp;action=edit&amp;section=1">edit</a>]</span> <span class="mw-headline">History</span></h2>
<h2><span class="editsection">[<a title="Edit section: Physical definitions" href="http://en.wikipedia.org/w/index.php?title=Gene&amp;action=edit&amp;section=3">edit</a>]</span> <span class="mw-headline">Physical definitions</span></h2>
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<div class="thumbinner" style="WIDTH: 302px"><a class="image" title="The chemical structure of a four-base fragment of a DNA double helix." href="http://en.wikipedia.org/wiki/Image:DNA_chemical_structure.svg"><img class="thumbimage" height="350" alt="The chemical structure of a four-base fragment of a DNA double helix." width="300" border="0" src="http://upload.wikimedia.org/wikipedia/commons/thumb/e/e4/DNA_chemical_structure.svg/300px-DNA_chemical_structure.svg.png" width="300" border="0" /></a>
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<div class="magnify"><a class="internal" title="Enlarge" href="http://en.wikipedia.org/wiki/Image:DNA_chemical_structure.svg"><img height="11" alt="" width="15" src="http://en.wikipedia.org/skins-1.5/common/images/magnify-clip.png" width="15" /></a></div>
The chemical structure of a four-base fragment of a DNA double helix.</div>
</div>
<h3><span class="editsection">[<a title="Edit section: Functional structure of a gene" href="http://en.wikipedia.org/w/index.php?title=Gene&amp;action=edit&amp;section=5">edit</a>]</span> <span class="mw-headline">Functional structure of a gene</span></h3>
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<div class="thumbinner" style="WIDTH: 402px"><a class="image" title="Diagram of the &quot;typical&quot; eukaryotic protein-coding gene. Promoters and enhancers determine what portions of the DNA will be transcribed into the precursor mRNA (pre-mRNA). The pre-mRNA is then spliced into messenger RNA (mRNA) which is later translated into protein." href="http://en.wikipedia.org/wiki/Image:Gene2-plain.svg"><img class="thumbimage" height="339" alt="Diagram of the &quot;typical&quot; eukaryotic protein-coding gene. Promoters and enhancers determine what portions of the DNA will be transcribed into the precursor mRNA (pre-mRNA). The pre-mRNA is then spliced into messenger RNA (mRNA) which is later translated into protein." width="400" border="0" src="http://upload.wikimedia.org/wikipedia/commons/thumb/a/a7/Gene2-plain.svg/400px-Gene2-plain.svg.png" width="400" border="0" /></a>
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<div class="magnify"><a class="internal" title="Enlarge" href="http://en.wikipedia.org/wiki/Image:Gene2-plain.svg"><img height="11" alt="" width="15" src="http://en.wikipedia.org/skins-1.5/common/images/magnify-clip.png" width="15" /></a></div>
Diagram of the &quot;typical&quot; <a title="Eukaryote" href="http://en.wikipedia.org/wiki/Eukaryote">eukaryotic</a> protein-coding <strong>gene</strong>. <a title="Promoter" href="http://en.wikipedia.org/wiki/Promoter">Promoters</a> and <a title="Enhancer" href="http://en.wikipedia.org/wiki/Enhancer">enhancers</a> determine what portions of the <a title="DNA" href="http://en.wikipedia.org/wiki/DNA">DNA</a> will be <a title="Transcription (genetics)" href="http://en.wikipedia.org/wiki/Transcription_%28genetics%29">transcribed</a> into the <a title="Precursor mRNA" href="http://en.wikipedia.org/wiki/Precursor_mRNA">precursor mRNA</a> (pre-mRNA). The pre-mRNA is then spliced into <a title="Messenger RNA" href="http://en.wikipedia.org/wiki/Messenger_RNA">messenger RNA</a> (mRNA) which is later <a title="Translation (biology)" href="http://en.wikipedia.org/wiki/Translation_%28biology%29">translated</a> into <a title="Protein" href="http://en.wikipedia.org/wiki/Protein">protein</a>.</div>
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<h3><span class="editsection">[<a title="Edit section: Genetic code" href="http://en.wikipedia.org/w/index.php?title=Gene&amp;action=edit&amp;section=8">edit</a>]</span> <span class="mw-headline">Genetic code</span></h3>
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<div class="thumbinner" style="WIDTH: 202px"><a class="image" title="Schematic diagram of a single-stranded RNA molecule illustrating the position of three-base codons." href="http://en.wikipedia.org/wiki/Image:Rna-codons-protein.png"><img class="thumbimage" height="259" alt="Schematic diagram of a single-stranded RNA molecule illustrating the position of three-base codons." width="200" border="0" src="http://upload.wikimedia.org/wikipedia/en/thumb/1/1d/Rna-codons-protein.png/200px-Rna-codons-protein.png" width="200" border="0" /></a>
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<div class="magnify"><a class="internal" title="Enlarge" href="http://en.wikipedia.org/wiki/Image:Rna-codons-protein.png"><img height="11" alt="" width="15" src="http://en.wikipedia.org/skins-1.5/common/images/magnify-clip.png" width="15" /></a></div>
Schematic diagram of a single-stranded RNA molecule illustrating the position of three-base codons.</div>
</div>
<p><a id="Genetic_and_genomic_nomenclature" name="Genetic_and_genomic_nomenclature"></a></p>
<h3><span class="editsection">[<a title="Edit section: Genetic and genomic nomenclature" href="http://en.wikipedia.org/w/index.php?title=Gene&amp;action=edit&amp;section=17">edit</a>]</span> <span class="mw-headline">Genetic and genomic nomenclature</span></h3>
<p><a title="Gene nomenclature" href="http://en.wikipedia.org/wiki/Gene_nomenclature">Gene nomenclature</a> has been established by the <a class="mw-redirect" title="HUGO" href="http://en.wikipedia.org/wiki/HUGO">HUGO</a> Gene Nomenclature Committee (HGNC) for each known human gene in the form of an approved gene name and <a title="Symbol" href="http://en.wikipedia.org/wiki/Symbol">symbol</a> (short-form <a title="Abbreviation" href="http://en.wikipedia.org/wiki/Abbreviation">abbreviation</a>). All approved symbols are stored in the <a class="external text" title="http://www.genenames.org/cgi-bin/hgnc_search.pl" rel="nofollow" href="http://www.genenames.org/cgi-bin/hgnc_search.pl" rel="nofollow">HGNC Database</a>. Each symbol is unique and each gene is only given one approved gene symbol. It is necessary to provide a unique symbol for each gene so that people can talk about them. This also facilitates <a title="Electronics" href="http://en.wikipedia.org/wiki/Electronics">electronic</a> <a title="Data" href="http://en.wikipedia.org/wiki/Data">data</a> retrieval from publications. In preference each symbol maintains parallel construction in different members of a <a title="Gene family" href="http://en.wikipedia.org/wiki/Gene_family">gene family</a> and can be used in other <a title="Species" href="http://en.wikipedia.org/wiki/Species"><font color="#810081">species</font></a>, especially the <a title="Mouse" href="http://en.wikipedia.org/wiki/Mouse">mouse</a>.</p>
<p><a id="Evolutionary_concept_of_a_gene" name="Evolutionary_concept_of_a_gene"></a></p>
<h2><span class="editsection">[<a title="Edit section: Evolutionary concept of a gene" href="http://en.wikipedia.org/w/index.php?title=Gene&amp;action=edit&amp;section=18">edit</a>]</span> <span class="mw-headline">Evolutionary concept of a gene</span></h2>
<li id="_note-Gerstein_2007"><strong><a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-Gerstein_2007_0">^</a></strong> <cite style="FONT-STYLE: normal">Gerstein MB, Bruce C, Rozowsky JS, Zheng D, Du J, Korbel JO, Emanuelsson O, Zhang ZD, Weissman S, Snyder M (2007). &quot;What is a gene, post-ENCODE? History and updated definition&quot;. <em>Genome Research</em> 17 (6): 669-681. <a class="external" title="http://www.ncbi.nlm.nih.gov/pubmed/17567988" href="http://www.ncbi.nlm.nih.gov/pubmed/17567988">PMID 17567988</a>.</cite><span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.atitle=What+is+a+gene%2C+post-ENCODE%3F+History+and+updated+definition&amp;rft.jtitle=Genome+Research&amp;rft.date=2007&amp;rft.volume=17&amp;rft.issue=6&amp;rft.au=Gerstein+MB%2C+Bruce+C%2C+Rozowsky+JS%2C+Zheng+D%2C+Du+J%2C+Korbel+JO%2C+Emanuelsson+O%2C+Zhang+ZD%2C+Weissman+S%2C+Snyder+M&amp;rft.pages=669-681&amp;rft_id=info:pmid/17567988"> </span> </li>
<li id="_note-Cavalier-Smith"><strong><a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-Cavalier-Smith_0">^</a></strong> Cavalier-Smith T. (1985). Eukaryotic gene numbers, non-coding DNA, and genome size. In Cavalier-Smith T, ed. <em>The Evolution of Genome Size</em> Chichester: John Wiley. </li>
<li id="_note-IHSGC2004">^ <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-IHSGC2004_0"><sup><em><strong>a</strong></em></sup></a> <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-IHSGC2004_1"><sup><em><strong>b</strong></em></sup></a> <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-IHSGC2004_2"><sup><em><strong>c</strong></em></sup></a> <cite style="FONT-STYLE: normal">International Human Genome Sequencing Consortium (2004). &quot;<a class="external text" title="http://www.nature.com/nature/journal/v431/n7011/full/nature03001.html" rel="nofollow" href="http://www.nature.com/nature/journal/v431/n7011/full/nature03001.html" rel="nofollow">Finishing the euchromatic sequence of the human genome.</a>&quot;. <em>Nature</em> 431 (7011): 931-45. <a class="external" title="http://www.ncbi.nlm.nih.gov/pubmed/15496913" href="http://www.ncbi.nlm.nih.gov/pubmed/15496913">PMID 15496913</a>.</cite><span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.atitle=Finishing+the+euchromatic+sequence+of+the+human+genome.&amp;rft.jtitle=Nature&amp;rft.date=2004&amp;rft.volume=431&amp;rft.issue=7011&amp;rft.au=International+Human+Genome+Sequencing+Consortium&amp;rft.pages=931-45&amp;rft_id=http%3A%2F%2Fwww.nature.com%2Fnature%2Fjournal%2Fv431%2Fn7011%2Ffull%2Fnature03001.html"> </span> </li> <li id="_note-gene_count2007">^ <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-gene_count2007_0"><sup><em><strong>a</strong></em></sup></a> <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-gene_count2007_1"><sup><em><strong>b</strong></em></sup></a> <cite style="FONT-STYLE: normal">Pennisi, Elizabeth (2007). &quot;<a class="external text" title="http://www.sciencemag.org/cgi/content/full/316/5828/1113a?maxtoshow=&amp;HITS=10&amp;hits=10&amp;RESULTFORMAT=&amp;fulltext=gene+count&amp;searchid=1&amp;FIRSTINDEX=0&amp;resourcetype=HWCIT" rel="nofollow" href="http://www.sciencemag.org/cgi/content/full/316/5828/1113a?maxtoshow=&amp;HITS=10&amp;hits=10&amp;RESULTFORMAT=&amp;fulltext=gene+count&amp;searchid=1&amp;FIRSTINDEX=0&amp;resourcetype=HWCIT" rel="nofollow">Working the (Gene Count) Numbers_ Finally, a Firm Answer</a>&quot;. <em>Science</em> 316 (5828): 1113.</cite><span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.atitle=Working+the+%28Gene+Count%29+Numbers_+Finally%2C+a+Firm+Answer&amp;rft.jtitle=Science&amp;rft.date=2007&amp;rft.volume=316&amp;rft.issue=5828&amp;rft.au=Pennisi%2C+Elizabeth&amp;rft.pages=1113&amp;rft_id=http%3A%2F%2Fwww.sciencemag.org%2Fcgi%2Fcontent%2Ffull%2F316%2F5828%2F1113a%3Fmaxtoshow%3D%26HITS%3D10%26hits%3D10%26RESULTFORMAT%3D%26fulltext%3Dgene%2Bcount%26searchid%3D1%26FIRSTINDEX%3D0%26resourcetype%3DHWCIT"> </span> </li>
<li id="_note-Watson_2004">^ <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-Watson_2004_0"><sup><em><strong>a</strong></em></sup></a> <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-Watson_2004_1"><sup><em><strong>b</strong></em></sup></a> <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-Watson_2004_2"><sup><em><strong>c</strong></em></sup></a> <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-Watson_2004_3"><sup><em><strong>d</strong></em></sup></a> <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-Watson_2004_4"><sup><em><strong>e</strong></em></sup></a> <cite class="book" style="FONT-STYLE: normal">Watson JD, Baker TA, Bell SP, Gann A, Levine M, Losick R (2004). <em>Molecular Biology of the Gene</em>, 5th ed., Peason Benjamin Cummings (Cold Spring Harbor Laboratory Press). <a class="internal" href="http://en.wikipedia.org/w/index.php?title=Special:Booksources&amp;isbn=080534635X">ISBN 080534635X</a>.</cite><span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=book&amp;rft.btitle=Molecular+Biology+of+the+Gene&amp;rft.au=Watson+JD%2C+Baker+TA%2C+Bell+SP%2C+Gann+A%2C+Levine+M%2C+Losick+R&amp;rft.edition=5th+ed.&amp;rft.pub=Peason+Benjamin+Cummings+%28Cold+Spring+Harbor+Laboratory+Press%29">&nbsp;</span> </li>
<li id="_note-pangen">^ <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-pangen_0"><sup><em><strong>a</strong></em></sup></a> <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-pangen_1"><sup><em><strong>b</strong></em></sup></a> Vries, H. de (1889) <em>Intracellular Pangenesis</em> <a class="external autonumber" title="http://www.esp.org/books/devries/pangenesis/facsimile/" rel="nofollow" href="http://www.esp.org/books/devries/pangenesis/facsimile/" rel="nofollow">[1]</a> (&quot;pangen&quot; definition on page 7 and 40 of this 1910 translation in English) </li>
<li id="_note-Gerstein">^ <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-Gerstein_0"><sup><em><strong>a</strong></em></sup></a> <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-Gerstein_1"><sup><em><strong>b</strong></em></sup></a> <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-Gerstein_2"><sup><em><strong>c</strong></em></sup></a> <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-Gerstein_3"><sup><em><strong>d</strong></em></sup></a> <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-Gerstein_4"><sup><em><strong>e</strong></em></sup></a> Mark B. Gerstein <em>et al.</em>, &quot;What is a gene, post-ENCODE? History and updated definition,&quot; <em>Genome Research</em> 17(6) (2007): 669-681 </li>
<li id="_note-Min_1972"><strong><a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-Min_1972_0">^</a></strong> <cite style="FONT-STYLE: normal">Min Jou W, Haegeman G, Ysebaert M, Fiers W (1972). &quot;Nucleotide sequence of the gene coding for the bacteriophage MS2 coat protein&quot;. <em>Nature</em> 237 (5350): 82-8. <a class="external" title="http://www.ncbi.nlm.nih.gov/pubmed/4555447" href="http://www.ncbi.nlm.nih.gov/pubmed/4555447">PMID 4555447</a>.</cite><span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.atitle=Nucleotide+sequence+of+the+gene+coding+for+the+bacteriophage+MS2+coat+protein&amp;rft.jtitle=Nature&amp;rft.date=1972&amp;rft.volume=237&amp;rft.issue=5350&amp;rft.au=Min+Jou+W%2C+Haegeman+G%2C+Ysebaert+M%2C+Fiers+W&amp;rft.pages=82-8&amp;rft_id=info:pmid/4555447"> </span> </li>
<li id="_note-genome"><strong><a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-genome_0">^</a></strong> <a class="external text" title="http://www.genome.gov/Pages/Education/Kit/main.cfm?pageid=24" rel="nofollow" href="http://www.genome.gov/Pages/Education/Kit/main.cfm?pageid=24" rel="nofollow">The Human Genome Project Timeline</a>. Retrieved on <a title="2006" href="http://en.wikipedia.org/wiki/2006">2006</a>-<a title="September 13" href="http://en.wikipedia.org/wiki/September_13">09-13</a>. </li>
<li id="_note-Darwin"><strong><a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-Darwin_0">^</a></strong> Darwin C. (1868). Animals and Plants under Domestication (1868). </li>
<li id="_note-rass">^ <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-rass_0"><sup><em><strong>a</strong></em></sup></a> <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-rass_1"><sup><em><strong>b</strong></em></sup></a> <cite style="FONT-STYLE: normal">Rassoulzadegan M, Grandjean V, Gounon P, Vincent S, Gillot I, Cuzin F (2006). &quot;RNA-mediated non-mendelian inheritance of an epigenetic change in the mouse&quot;. <em>Nature</em> 441 (7092): 469-74. <a class="external" title="http://www.ncbi.nlm.nih.gov/pubmed/16724059" href="http://www.ncbi.nlm.nih.gov/pubmed/16724059">PMID 16724059</a>.</cite><span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.atitle=RNA-mediated+non-mendelian+inheritance+of+an+epigenetic+change+in+the+mouse&amp;rft.jtitle=Nature&amp;rft.date=2006&amp;rft.volume=441&amp;rft.issue=7092&amp;rft.au=Rassoulzadegan+M%2C+Grandjean+V%2C+Gounon+P%2C+Vincent+S%2C+Gillot+I%2C+Cuzin+F&amp;rft.pages=469-74&amp;rft_id=info:pmid/16724059"> </span> </li>
<li id="_note-1"><strong><a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-1">^</a></strong> <cite style="FONT-STYLE: normal">Woodson SA (1998). &quot;<a class="external text" title="http://www.genesdev.org/cgi/content/full/12/9/1243" rel="nofollow" href="http://www.genesdev.org/cgi/content/full/12/9/1243" rel="nofollow">Ironing out the kinks: splicing and translation in bacteria</a>&quot;. <em>Genes Dev.</em> 12 (9): 1243&ndash;7. <a class="external" title="http://www.ncbi.nlm.nih.gov/pubmed/9573040" href="http://www.ncbi.nlm.nih.gov/pubmed/9573040">PMID 9573040</a>.</cite><span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.atitle=Ironing+out+the+kinks%3A+splicing+and+translation+in+bacteria&amp;rft.jtitle=Genes+Dev.&amp;rft.date=1998&amp;rft.volume=12&amp;rft.issue=9&amp;rft.au=Woodson+SA&amp;rft.pages=1243%E2%80%937&amp;rft_id=info:pmid/9573040&amp;rft_id=http%3A%2F%2Fwww.genesdev.org%2Fcgi%2Fcontent%2Ffull%2F12%2F9%2F1243"> </span> </li>
<li id="_note-Braig"><strong><a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-Braig_0">^</a></strong> <cite style="FONT-STYLE: normal">Braig M, Schmitt C (2006). &quot;Oncogene-induced senescence: putting the brakes on tumor development&quot;. <em>Cancer Res</em> 66 (6): 2881-4. <a class="external" title="http://www.ncbi.nlm.nih.gov/pubmed/16540631" href="http://www.ncbi.nlm.nih.gov/pubmed/16540631">PMID 16540631</a>.</cite><span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.atitle=Oncogene-induced+senescence%3A+putting+the+brakes+on+tumor+development&amp;rft.jtitle=Cancer+Res&amp;rft.date=2006&amp;rft.volume=66&amp;rft.issue=6&amp;rft.au=Braig+M%2C+Schmitt+C&amp;rft.pages=2881-4&amp;rft_id=info:pmid/16540631"> </span> </li>
<li id="_note-Mount_2004"><strong><a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-Mount_2004_0">^</a></strong> <cite class="book" id="Reference-Mount-2004" style="FONT-STYLE: normal">Mount, DW (2004). <em>Bioinformatics: Sequence and genome analysis</em>, 2nd ed., Cold Spring Harbor Laboratory Press: Cold Spring Harbor, New York. <a class="internal" href="http://en.wikipedia.org/w/index.php?title=Special:Booksources&amp;isbn=0879697121">ISBN 0879697121</a>.</cite><span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=book&amp;rft.btitle=Bioinformatics%3A+Sequence+and+genome+analysis&amp;rft.aulast=Mount&amp;rft.aufirst=DW&amp;rft.edition=2nd+ed.&amp;rft.pub=Cold+Spring+Harbor+Laboratory+Press%3A+Cold+Spring+Harbor%2C+New+York">&nbsp;</span> </li>
<li id="_note-Lodish"><strong><a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-Lodish_0">^</a></strong> Lodish, H, Berk A, Matsudaira P, Kaiser CA, Krieger M, Scott MP, Zipursky SL, Darnell J. (2004). <em>Molecular Cell Biology</em>, 5th, New York: WH Freeman. </li>
<li id="_note-2"><strong><a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-2">^</a></strong> Thomas KR, Capecchi MR. Site-directed mutagenesis by gene targeting in mouse embryo-derived stem cells. Cell. 1987;51:503-12 </li>
<li id="_note-3"><strong><a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-3">^</a></strong> <a class="external free" title="http://nobelprize.org/nobel_prizes/medicine/laureates/2007/press.html" rel="nofollow" href="http://nobelprize.org/nobel_prizes/medicine/laureates/2007/press.html" rel="nofollow">http://nobelprize.org/nobel_prizes/medicine/laureates/2007/press.html</a> </li> <li id="_note-Deng_2007">^ <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-Deng_2007_0"><sup><em><strong>a</strong></em></sup></a> <a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-Deng_2007_1"><sup><em><strong>b</strong></em></sup></a> Deng C. In Celebration of Dr. Mario R. Capecchi's Nobel Prize. Int J Biol Sci 2007; 3:417-419. <a class="external free" title="http://www.biolsci.org/v03p0417.htm" rel="nofollow" href="http://www.biolsci.org/v03p0417.htm" rel="nofollow">http://www.biolsci.org/v03p0417.htm</a> </li> <li id="_note-4"><strong><a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-4">^</a></strong> <a class="external free" title="http://www.hhmi.org/research/investigators/capecchi.html" rel="nofollow" href="http://www.hhmi.org/research/investigators/capecchi.html" rel="nofollow">http://www.hhmi.org/research/investigators/capecchi.html</a> </li>
<li id="_note-5"><strong><a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-5">^</a></strong> Lolle &amp; colleagues (2005) Genome-wide non-mendelian inheritance of extra-genomic information in Arabidopsis. <a class="external" title="http://www.ncbi.nlm.nih.gov/pubmed/15785770" href="http://www.ncbi.nlm.nih.gov/pubmed/15785770">PMID 15785770</a> </li>
<li id="_note-6"><strong><a title="" href="http://en.wikipedia.org/wiki/Gene#_ref-6">^</a></strong> Spilianakis &amp; colleagues (2005) Interchromosomal associations between alternatively expressed loci. <a class="external" title="http://www.ncbi.nlm.nih.gov/pubmed/15880101" href="http://www.ncbi.nlm.nih.gov/pubmed/15880101">PMID 15880101</a> </li>
<h2><span class="editsection">[<a title="Edit section: Further reading" href="http://en.wikipedia.org/w/index.php?title=Gene&amp;action=edit&amp;section=23">edit</a>]</span> <span class="mw-headline">Further reading</span></h2>
<ul>
<li><cite class="book" id="Reference-Dawkins-1990" style="FONT-STYLE: normal"><a title="Richard Dawkins" href="http://en.wikipedia.org/wiki/Richard_Dawkins">Dawkins, Richard</a> (1990). <em><a title="The Selfish Gene" href="http://en.wikipedia.org/wiki/The_Selfish_Gene">The Selfish Gene</a></em>. Oxford University Press. <a class="internal" href="http://en.wikipedia.org/w/index.php?title=Special:Booksources&amp;isbn=0192860925">ISBN 0-19-286092-5</a>.</cite><span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=book&amp;rft.btitle=%5B%5BThe+Selfish+Gene%5D%5D&amp;rft.aulast=Dawkins&amp;rft.aufirst=Richard&amp;rft.pub=Oxford+University+Press">&nbsp;</span> <a class="external text" title="http://print.google.com/print?id=WkHO9HI7koEC" rel="nofollow" href="http://print.google.com/print?id=WkHO9HI7koEC" rel="nofollow">Google Book Search</a>; first published 1976. </li>
<li><cite class="book" id="Reference-Dawkins-1995" style="FONT-STYLE: normal"><a title="Richard Dawkins" href="http://en.wikipedia.org/wiki/Richard_Dawkins">Dawkins, Richard</a> (1995). <em><a class="mw-redirect" title="River Out of Eden" href="http://en.wikipedia.org/wiki/River_Out_of_Eden">River Out of Eden</a></em>. Basic Books. <a class="internal" href="http://en.wikipedia.org/w/index.php?title=Special:Booksources&amp;isbn=0465069908">ISBN 0-465-06990-8</a>.</cite><span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=book&amp;rft.btitle=%5B%5BRiver+Out+of+Eden%5D%5D&amp;rft.aulast=Dawkins&amp;rft.aufirst=Richard&amp;rft.pub=Basic+Books">&nbsp;</span> </li>
</ul>
<h2><span class="editsection">[<a title="Edit section: External links" href="http://en.wikipedia.org/w/index.php?title=Gene&amp;action=edit&amp;section=24">edit</a>]</span> <span class="mw-headline">External links</span></h2>
<ul>
<li><a class="external text" title="http://www.dnalc.org/" rel="nofollow" href="http://www.dnalc.org/" rel="nofollow">The Dolan DNA Learning Center</a> </li> <li><a class="external text" title="http://www.dnai.org/" rel="nofollow" href="http://www.dnai.org/" rel="nofollow">DNA Interactive</a> </li> <li><a class="external text" title="http://www.dnaftb.org/" rel="nofollow" href="http://www.dnaftb.org/" rel="nofollow">DNA From The Beginning</a> </li>
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<h3><span class="editsection">[<a title="Edit section: Tutorial and news" href="http://en.wikipedia.org/w/index.php?title=Gene&amp;action=edit&amp;section=25">edit</a>]</span> <span class="mw-headline">Tutorial and news</span></h3>
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<li><a class="external text" title="http://www.scienceaid.co.uk/biology/genetics" rel="nofollow" href="http://www.scienceaid.co.uk/biology/genetics" rel="nofollow">Science aid: Genetics</a> for beginners </li> <li><a class="external text" title="http://www.newscientist.com/news/news.jsp?id=ns99996561" rel="nofollow" href="http://www.newscientist.com/news/news.jsp?id=ns99996561" rel="nofollow">Recount slashes number of human genes</a> (from <a title="New Scientist" href="http://en.wikipedia.org/wiki/New_Scientist">New Scientist</a> magazine) </li> <li><a class="external text" title="http://www.genome.gov/12513430" rel="nofollow" href="http://www.genome.gov/12513430" rel="nofollow">National Human Genome Research Institute &mdash; News Release</a> </li>
<li><cite style="FONT-STYLE: normal">(2004) &quot;Finishing the euchromatic sequence of the human genome&quot;. <em>Nature</em> 431 (7011): 931-45. <a class="external" title="http://www.ncbi.nlm.nih.gov/pubmed/15496913" href="http://www.ncbi.nlm.nih.gov/pubmed/15496913">PMID 15496913</a>.</cite><span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.atitle=Finishing+the+euchromatic+sequence+of+the+human+genome&amp;rft.jtitle=Nature&amp;rft.date=2004&amp;rft.volume=431&amp;rft.issue=7011&amp;rft.pages=931-45&amp;rft_id=info:pmid/15496913"> </span> </li>
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<h3><span class="editsection">[<a title="Edit section: References and databases" href="http://en.wikipedia.org/w/index.php?title=Gene&amp;action=edit&amp;section=26">edit</a>]</span> <span class="mw-headline">References and databases</span></h3>
<ul>
<li><a class="external text" title="http://www.gene.ucl.ac.uk/nomenclature" rel="nofollow" href="http://www.gene.ucl.ac.uk/nomenclature" rel="nofollow">HUGO Gene Nomenclature Committee, HGNC</a> </li> <li><a class="external text" title="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=OMIM" rel="nofollow" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=OMIM" rel="nofollow">OMIM</a> NIH's National Library of Medicine NCBI website link to Online Mendelian Inheritance in Man. </li> <li><a class="external text" title="http://www.hugo-international.org" rel="nofollow" href="http://www.hugo-international.org/" rel="nofollow">Human Genome Organisation, HUGO</a> </li> <li><a class="external text" title="http://www.ihop-net.org/UniPub/iHOP/" rel="nofollow" href="http://www.ihop-net.org/UniPub/iHOP/" rel="nofollow">iHOP - Information Hyperlinked over Proteins</a> </li> <li><a class="external text" title="http://www.pir.uniprot.org/" rel="nofollow" href="http://www.pir.uniprot.org/" rel="nofollow">UniProt</a> </li> <li><a class="external text" title="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=gene" rel="nofollow" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=gene" rel="nofollow">Entrez Gene - A searchable database of genes</a> </li> <li><a class="external text" title="http://idconverter.bioinfo.cnio.es" rel="nofollow" href="http://idconverter.bioinfo.cnio.es/" rel="nofollow">IDconverter - Map your ids to other known public DBs</a> </li> <li><a class="external text" title="http://www.genecards.org" rel="nofollow" href="http://www.genecards.org/" rel="nofollow">GeneCards - the Human Gene Compendium</a> </li> <li><a class="external text" title="http://www.genetherapynet.com" rel="nofollow" href="http://www.genetherapynet.com/" rel="nofollow">Gene Therapy Net</a> </li>
</ul>
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