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class="vector-toc-numb">3.1</span> <span>Enhanced phenotypic evolution</span> </div> </a> <ul id="toc-Enhanced_phenotypic_evolution-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Genome_diversity" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genome_diversity"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Genome diversity</span> </div> </a> <ul id="toc-Genome_diversity-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Hybrid_vigor" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Hybrid_vigor"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.3</span> <span>Hybrid vigor</span> </div> </a> <ul id="toc-Hybrid_vigor-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Speciation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Speciation"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.4</span> <span>Speciation</span> </div> </a> <ul id="toc-Speciation-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Allopolyploidy_and_autopolyploidy" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Allopolyploidy_and_autopolyploidy"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Allopolyploidy and autopolyploidy</span> </div> </a> <ul id="toc-Allopolyploidy_and_autopolyploidy-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Vertebrates_as_paleopolyploid" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Vertebrates_as_paleopolyploid"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Vertebrates as paleopolyploid</span> </div> </a> <ul id="toc-Vertebrates_as_paleopolyploid-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item 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dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">State of having undergone whole genome duplication in deep evolutionary time</div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Paleopolyploidy.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/7f/Paleopolyploidy.jpg/280px-Paleopolyploidy.jpg" decoding="async" width="280" height="184" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/7f/Paleopolyploidy.jpg/420px-Paleopolyploidy.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/7/7f/Paleopolyploidy.jpg 2x" data-file-width="448" data-file-height="295" /></a><figcaption>Overview of paleopolyploidy process. Many higher eukaryotes were paleopolyploids at some point during their evolutionary history.</figcaption></figure> <p><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>Paleopolyploidy</b> is the result of <a href="/wiki/Genome_duplications" class="mw-redirect" title="Genome duplications">genome duplications</a> which occurred at least several million years ago (MYA). Such an event could either double the genome of a single species (<a href="/wiki/Autopolyploidy" class="mw-redirect" title="Autopolyploidy">autopolyploidy</a>) or combine those of two species (<a href="/wiki/Allopolyploidy" class="mw-redirect" title="Allopolyploidy">allopolyploidy</a>). Because of functional <a href="/wiki/Gene_redundancy" title="Gene redundancy">redundancy</a>, genes are rapidly silenced or lost from the duplicated genomes. Most paleopolyploids, through evolutionary time, have lost their <a href="/wiki/Polyploid" class="mw-redirect" title="Polyploid">polyploid</a> status through a process called <b><a href="/wiki/Diploidization" title="Diploidization">diploidization</a></b>, and are currently considered <a href="/wiki/Diploid" class="mw-redirect" title="Diploid">diploids</a>, e.g., <a href="/wiki/Baker%27s_yeast" title="Baker's yeast">baker's yeast</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> <i><a href="/wiki/Arabidopsis_thaliana" title="Arabidopsis thaliana">Arabidopsis thaliana</a></i>,<sup id="cite_ref-bowers_3-0" class="reference"><a href="#cite_note-bowers-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> and perhaps <a href="/wiki/Human" title="Human">humans</a>.<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><sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-paleopolyploidy_6-0" class="reference"><a href="#cite_note-paleopolyploidy-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Blanc2004b_7-0" class="reference"><a href="#cite_note-Blanc2004b-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p><p>Paleopolyploidy is extensively studied in plant lineages. It has been found that almost all flowering plants have undergone at least one round of genome duplication at some point during their evolutionary history. Ancient genome duplications are also found in the early ancestor of vertebrates (which includes the human lineage) near the origin of the <a href="/wiki/Bony_fish" class="mw-redirect" title="Bony fish">bony fishes</a>, and another in the stem lineage of <a href="/wiki/Teleost_fish" class="mw-redirect" title="Teleost fish">teleost fishes</a>.<sup id="cite_ref-:0_8-0" class="reference"><a href="#cite_note-:0-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Evidence suggests that baker's yeast (<i><a href="/wiki/Saccharomyces_cerevisiae" title="Saccharomyces cerevisiae">Saccharomyces cerevisiae</a></i>), which has a compact genome, experienced polyploidization during its evolutionary history. </p><p>The term <i>mesopolyploid</i> is sometimes used for species that have undergone whole genome multiplication events (whole genome duplication, whole genome triplification, etc.) in more recent history, such as within the last 17 million years.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Eukaryotes">Eukaryotes</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paleopolyploidy&action=edit&section=1" title="Edit section: Eukaryotes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:PaleopolyploidyTree.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/35/PaleopolyploidyTree.jpg/300px-PaleopolyploidyTree.jpg" decoding="async" width="300" height="305" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/35/PaleopolyploidyTree.jpg/450px-PaleopolyploidyTree.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/35/PaleopolyploidyTree.jpg/600px-PaleopolyploidyTree.jpg 2x" data-file-width="847" data-file-height="861" /></a><figcaption>A diagram that summarizes all well-known paleopolyploidization events.</figcaption></figure> <p>Ancient genome duplications are widespread throughout <a href="/wiki/Eukaryotic" class="mw-redirect" title="Eukaryotic">eukaryotic</a> lineages, particularly in plants. Studies suggest that the common ancestor of <a href="/wiki/Poaceae" title="Poaceae">Poaceae</a>, the grass family which includes important crop species such as maize, rice, wheat, and sugar cane, shared a whole genome duplication about <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=70">70</a> million years ago</span>.<sup id="cite_ref-paterson_10-0" class="reference"><a href="#cite_note-paterson-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> In more ancient monocot lineages one or likely multiple rounds of additional whole genome duplications had occurred, which were however not shared with the ancestral <a href="/wiki/Eudicots" title="Eudicots">eudicots</a>.<sup id="cite_ref-Tang_2010_11-0" class="reference"><a href="#cite_note-Tang_2010-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Further independent more recent whole genome duplications have occurred in the lineages leading to maize, sugar cane and wheat, but not rice, sorghum or foxtail millet.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2021)">citation needed</span></a></i>]</sup> </p><p>A polyploidy event <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=160">160</a> million years ago</span> is theorized to have created the ancestral line that led to all modern flowering plants.<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> That paleopolyploidy event was studied by sequencing the genome of an ancient flowering plant, <i><a href="/wiki/Amborella_trichopoda" class="mw-redirect" title="Amborella trichopoda">Amborella trichopoda</a></i>.<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p>The core eudicots also shared a common whole genome triplication (paleo-hexaploidy), which was estimated to have occurred after <a href="/wiki/Monocotyledon" title="Monocotyledon">monocot</a>-<a href="/wiki/Eudicots" title="Eudicots">eudicot</a> divergence but before the divergence of <a href="/wiki/Rosids" title="Rosids">rosids</a> and <a href="/wiki/Asterids" title="Asterids">asterids</a>.<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-grape_15-0" class="reference"><a href="#cite_note-grape-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> Many eudicot species have experienced additional whole genome duplications or triplications. For example, the model plant <i><a href="/wiki/Arabidopsis_thaliana" title="Arabidopsis thaliana">Arabidopsis thaliana</a></i>, the first plant to have its entire genome sequenced, has experienced at least two additional rounds of whole genome duplication since the duplication shared by the core eudicots.<sup id="cite_ref-bowers_3-1" class="reference"><a href="#cite_note-bowers-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> The most recent event took place before the divergence of the <i>Arabidopsis</i> and <i><a href="/wiki/Brassica" title="Brassica">Brassica</a></i> lineages, about <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=20">20</a> million years ago</span> to <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=45">45</a> million years ago</span>. Other examples include the sequenced eudicot genomes of apple, soybean, tomato, cotton, etc.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2021)">citation needed</span></a></i>]</sup> </p><p>Compared with plants, paleopolyploidy is much rarer in the animal kingdom. It has been identified mainly in amphibians and bony fishes. Although some studies suggested one or more common genome duplications are shared by all vertebrates (including humans), the evidence is not as strong as in the other cases because the duplications, if they exist, happened so long ago (about 400-500 Ma compared to less than 200 Ma in plants), and the matter is still under debate. The idea that vertebrates share a common whole genome duplication is known as the <a href="/wiki/2R_Hypothesis" class="mw-redirect" title="2R Hypothesis">2R Hypothesis</a>. Many researchers are interested in the reason why animal lineages, particularly mammals, have had so many fewer whole genome duplications than plant lineages.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2021)">citation needed</span></a></i>]</sup> </p><p>A well-supported paleopolyploidy has been found in baker's yeast (<i>Saccharomyces cerevisiae</i>), despite its small, compact genome (~13Mbp), after the divergence from <i><a href="/wiki/Kluyveromyces_lactis" title="Kluyveromyces lactis">Kluyveromyces lactis</a></i> and <i><a href="/wiki/Kluyveromyces_marxianus" title="Kluyveromyces marxianus">K. marxianus</a></i>.<sup id="cite_ref-wong_17-0" class="reference"><a href="#cite_note-wong-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Through genome streamlining, yeast has lost 90% of the duplicated genome over evolutionary time and is now recognized as a diploid organism.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2021)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading2"><h2 id="Detection_method">Detection method</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paleopolyploidy&action=edit&section=2" title="Edit section: Detection method"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1251242444">.mw-parser-output .ambox{border:1px solid #a2a9b1;border-left:10px solid #36c;background-color:#fbfbfb;box-sizing:border-box}.mw-parser-output .ambox+link+.ambox,.mw-parser-output .ambox+link+style+.ambox,.mw-parser-output .ambox+link+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+style+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+link+.ambox{margin-top:-1px}html body.mediawiki .mw-parser-output .ambox.mbox-small-left{margin:4px 1em 4px 0;overflow:hidden;width:238px;border-collapse:collapse;font-size:88%;line-height:1.25em}.mw-parser-output .ambox-speedy{border-left:10px solid #b32424;background-color:#fee7e6}.mw-parser-output .ambox-delete{border-left:10px solid #b32424}.mw-parser-output .ambox-content{border-left:10px solid #f28500}.mw-parser-output .ambox-style{border-left:10px solid #fc3}.mw-parser-output .ambox-move{border-left:10px solid #9932cc}.mw-parser-output .ambox-protection{border-left:10px solid #a2a9b1}.mw-parser-output .ambox .mbox-text{border:none;padding:0.25em 0.5em;width:100%}.mw-parser-output .ambox .mbox-image{border:none;padding:2px 0 2px 0.5em;text-align:center}.mw-parser-output .ambox .mbox-imageright{border:none;padding:2px 0.5em 2px 0;text-align:center}.mw-parser-output .ambox .mbox-empty-cell{border:none;padding:0;width:1px}.mw-parser-output .ambox .mbox-image-div{width:52px}@media(min-width:720px){.mw-parser-output .ambox{margin:0 10%}}@media print{body.ns-0 .mw-parser-output .ambox{display:none!important}}</style><table class="box-Unreferenced_section plainlinks metadata ambox ambox-content ambox-Unreferenced" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><a href="/wiki/File:Question_book-new.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/50px-Question_book-new.svg.png" decoding="async" width="50" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/75px-Question_book-new.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/100px-Question_book-new.svg.png 2x" data-file-width="512" data-file-height="399" /></a></span></div></td><td class="mbox-text"><div class="mbox-text-span">This section <b>does not <a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources">cite</a> any <a href="/wiki/Wikipedia:Verifiability" title="Wikipedia:Verifiability">sources</a></b>.<span class="hide-when-compact"> Please help <a href="/wiki/Special:EditPage/Paleopolyploidy" title="Special:EditPage/Paleopolyploidy">improve this section</a> by <a href="/wiki/Help:Referencing_for_beginners" title="Help:Referencing for beginners">adding citations to reliable sources</a>. Unsourced material may be challenged and <a href="/wiki/Wikipedia:Verifiability#Burden_of_evidence" title="Wikipedia:Verifiability">removed</a>.</span> <span class="date-container"><i>(<span class="date">January 2021</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <p>Duplicated genes can be identified through <a href="/wiki/Sequence_homology" title="Sequence homology">sequence homology</a> on the DNA or protein level. Paleopolyploidy can be identified as massive gene duplication at one time using a <a href="/wiki/Molecular_clock" title="Molecular clock">molecular clock</a>. To distinguish between whole-genome duplication and a collection of (more common) single <a href="/wiki/Gene_duplication" title="Gene duplication">gene duplication</a> events, the following rules are often applied: </p> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:NumbervsKs.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e3/NumbervsKs.jpg/280px-NumbervsKs.jpg" decoding="async" width="280" height="187" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e3/NumbervsKs.jpg/420px-NumbervsKs.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/e/e3/NumbervsKs.jpg 2x" data-file-width="496" data-file-height="332" /></a><figcaption>Detection of paleopolyploidy using Ks.</figcaption></figure> <ul><li>Duplicated genes are located in large duplicated blocks. Single gene duplication is a random process and tends to make duplicated genes scattered throughout the genome.</li> <li>Duplicated blocks are non-overlapping because they were created simultaneously. <b>Segmental duplication</b> within the genome can fulfill the first rule; but multiple independent segmental duplications could overlap each other.<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></li></ul> <p>In theory, the two duplicated genes should have the same "age"; that is, the divergence of the sequence should be equal between the two genes duplicated by paleopolyploidy (<a href="/wiki/Homeolog" class="mw-redirect" title="Homeolog">homeologs</a>). <a href="/wiki/Synonymous_substitution" title="Synonymous substitution">Synonymous substitution</a> rate, <b>Ks</b>, is often used as a molecular clock to determine the time of gene duplication.<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> Thus, paleopolyploidy is identified as a "peak" on the duplicate number vs. Ks graph (shown on the right). </p><p>However, using Ks plots to identify and document ancient polyploid events can be problematic, as the method fails to identify genome duplications that were followed by massive gene elimination and genome refinement. Other mixed model approaches that combined Ks plots with other methods are being developed to better understand paleopolyploidy.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p><p>Duplication events that occurred a long time ago in the history of various evolutionary lineages can be difficult to detect because of subsequent diploidization (such that a polyploid starts to behave cytogenetically as a diploid over time) as mutations and gene translations gradually make one copy of each chromosome unlike its counterpart. This usually results in a low confidence for identifying ancient paleopolyploidy.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Evolutionary_importance">Evolutionary importance</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paleopolyploidy&action=edit&section=3" title="Edit section: Evolutionary importance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Paleopolyploidization events lead to massive cellular changes, including doubling of the genetic material, changes in gene expression and increased cell size. Gene loss during diploidization is not completely random, but heavily selected. Genes from large gene families are duplicated. On the other hand, individual genes are not duplicated.<sup class="noprint Inline-Template" style="margin-left:0.1em; white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Please_clarify" title="Wikipedia:Please clarify"><span title="The text near this tag may need clarification or removal of jargon. (February 2018)">clarification needed</span></a></i>]</sup> Overall, paleopolyploidy can have both short-term and long-term evolutionary effects on an organism's fitness in the natural environment.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2021)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading3"><h3 id="Enhanced_phenotypic_evolution">Enhanced phenotypic evolution</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paleopolyploidy&action=edit&section=4" title="Edit section: Enhanced phenotypic evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Whole genome duplication may increase the rates and efficiency by which organisms acquire new biological traits. However, one test of this hypothesis, which compared evolutionary rates in innovation in early teleost fishes (with duplicate genomes) to early holostean fishes (without duplicated genomes) found little difference between the two.<sup id="cite_ref-:0_8-1" class="reference"><a href="#cite_note-:0-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Genome_diversity">Genome diversity</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paleopolyploidy&action=edit&section=5" title="Edit section: Genome diversity"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Genome doubling provided the organism with redundant alleles that can evolve freely with little selection pressure. The duplicated genes can undergo <a href="/wiki/Neofunctionalization" title="Neofunctionalization">neofunctionalization</a>, <a href="/wiki/Subfunctionalization" title="Subfunctionalization">subfunctionalization</a>, or <a href="/wiki/Gene_redundancy" title="Gene redundancy">nonfunctionalization</a> which could help the organism adapt to the new environment or survive different stress conditions.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Hybrid_vigor">Hybrid vigor</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paleopolyploidy&action=edit&section=6" title="Edit section: Hybrid vigor"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Polyploids often have larger cells and even larger organs. Many important crops, including wheat, maize and <a href="/wiki/Cotton" title="Cotton">cotton</a>, are paleopolyploids which were selected for domestication by ancient peoples.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Speciation">Speciation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paleopolyploidy&action=edit&section=7" title="Edit section: Speciation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>It has been suggested that many polyploidization events created new species, via a gain of adaptive traits, or by sexual incompatibility with their diploid counterparts. An example would be the recent <a href="/wiki/Speciation" title="Speciation">speciation</a> of allopolyploid <i><a href="/wiki/Spartina" title="Spartina">Spartina</a></i> — <i><a href="/wiki/Spartina_anglica" class="mw-redirect" title="Spartina anglica">S. anglica</a></i>; the polyploid plant is so successful that it is listed as an <a href="/wiki/Invasive_species" title="Invasive species">invasive species</a> in many regions.<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> </p> <div class="mw-heading mw-heading2"><h2 id="Allopolyploidy_and_autopolyploidy">Allopolyploidy and autopolyploidy</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paleopolyploidy&action=edit&section=8" title="Edit section: Allopolyploidy and autopolyploidy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There are two major divisions of <a href="/wiki/Polyploidy" title="Polyploidy">polyploidy</a>, allopolyploidy and autopolyploidy. Allopolyploids arise as a result of the hybridization of two related species, while autopolyploids arise from the duplication of a species' genome as a result of hybridization of two conspecific parents,<sup id="cite_ref-Soltis,_PS,_Soltis_DE_2000_7051–7057_25-0" class="reference"><a href="#cite_note-Soltis,_PS,_Soltis_DE_2000_7051–7057-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> or somatic doubling in reproductive tissue of a parent. Allopolyploid species are believed to be much more prevalent in nature,<sup id="cite_ref-Soltis,_PS,_Soltis_DE_2000_7051–7057_25-1" class="reference"><a href="#cite_note-Soltis,_PS,_Soltis_DE_2000_7051–7057-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> possibly because allopolyploids inherit different genomes, resulting in increased <a href="/wiki/Heterozygosity" class="mw-redirect" title="Heterozygosity">heterozygosity</a>, and therefore higher fitness. These different genomes result in an increased likelihood of large genomic reorganizations,<sup id="cite_ref-Soltis,_PS,_Soltis_DE_2000_7051–7057_25-2" class="reference"><a href="#cite_note-Soltis,_PS,_Soltis_DE_2000_7051–7057-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-auto_26-0" class="reference"><a href="#cite_note-auto-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> which can be either deleterious, or advantageous. Autopolyploidy, however, is generally considered to be a neutral process,<sup id="cite_ref-auto_26-1" class="reference"><a href="#cite_note-auto-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> though it has been hypothesized that autopolyploidy may serve as a useful mechanism for inducing speciation, and therefore assisting in the ability of an organism to quickly colonize in new habitats without undergoing the time-intensive and costly period of genomic reorganization experienced by allopolyploid species. One common source of autopolyploidy in plants stems from "<a href="/wiki/Perfect_flower" class="mw-redirect" title="Perfect flower">perfect flowers</a>", which are capable of <a href="/wiki/Self-pollination" title="Self-pollination">self-pollination</a>, or "selfing". This, along with errors in <a href="/wiki/Meiosis" title="Meiosis">meiosis</a> that lead to <a href="/wiki/Aneuploidy" title="Aneuploidy">aneuploidy</a>, can create an environment where autopolyploidy is very likely. This fact can be exploited in a laboratory setting by using <a href="/wiki/Colchicine" title="Colchicine">colchicine</a> to inhibit <a href="/wiki/Chromosome" title="Chromosome">chromosome</a> segregation during meiosis, creating synthetic autopolyploid plants.<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> </p><p>Following polyploidy events, there are several possible fates for duplicated <a href="/wiki/Gene" title="Gene">genes</a>; both copies may be retained as functional genes, change in gene function may occur in one or both copies, <a href="/wiki/Gene_silencing" title="Gene silencing">gene silencing</a> may mask one or both copies, or complete gene loss may occur.<sup id="cite_ref-Soltis,_PS,_Soltis_DE_2000_7051–7057_25-3" class="reference"><a href="#cite_note-Soltis,_PS,_Soltis_DE_2000_7051–7057-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><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> Polyploidy events will result in higher levels of heterozygosity, and, over time, can lead to an increase in the total number of functional genes in the genome. As time passes after a genome duplication event, many genes will change function as a result of either change in duplicate gene function for both allo- and autopolyploid species, or there will be changes in gene expression caused by genomic rearrangements induced by genome duplication in allopolyploids. When both copies of a gene are retained, and thus the number of copies doubled, there is a chance that there will be a proportional increase in expression of that gene, resulting in twice as much <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA</a> transcript being produced. There is also the possibility that transcription of a duplicated gene will be down-regulated, resulting in less than two-fold increase in transcription of that gene, or that the duplication event will yield more than a two-fold increase in transcription.<sup id="cite_ref-Coate_JE_&_Doyle_JJ_2010_534–546_29-0" class="reference"><a href="#cite_note-Coate_JE_&_Doyle_JJ_2010_534–546-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> In one species, <i>Glycine dolichocarpa</i> (a close relative of the <a href="/wiki/Soybean" title="Soybean">soybean</a>, <i>Glycine max</i>), it has been observed that following a genome duplication roughly 500,000 years ago, there has been a 1.4 fold increase in transcription, indicating that there has been a proportional decrease in transcription relative to gene copy number following the duplication event.<sup id="cite_ref-Coate_JE_&_Doyle_JJ_2010_534–546_29-1" class="reference"><a href="#cite_note-Coate_JE_&_Doyle_JJ_2010_534–546-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Vertebrates_as_paleopolyploid">Vertebrates as paleopolyploid</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paleopolyploidy&action=edit&section=9" title="Edit section: Vertebrates as paleopolyploid"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The hypothesis of vertebrate paleopolyploidy originated as early as the 1970s, proposed by the biologist <a href="/wiki/Susumu_Ohno" title="Susumu Ohno">Susumu Ohno</a>. He reasoned that the vertebrate genome could not achieve its complexity without large scale whole-genome duplications. The "two rounds of genome duplication" hypothesis (<a href="/wiki/2R_hypothesis" title="2R hypothesis">2R hypothesis</a>) came about, and gained in popularity, especially among developmental biologists.<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><p>Some researchers have questioned the 2R hypothesis because it predicts that vertebrate genomes should have a 4:1 gene ratio compared with invertebrate genomes, and this is not supported by findings from the 48 vertebrate genome projects available in mid-2011. For example, the human genome consists of ~20,500 protein coding genes according to counts from the Ensembl genome browser<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> while an average invertebrate genome size is about 15,000 genes. The <a href="/wiki/Branchiostoma_lanceolatum" title="Branchiostoma lanceolatum">amphioxus</a> genome sequence provided support for the hypothesis of two rounds of whole genome duplication, followed by loss of duplicate copies of most genes.<sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> Additional arguments against 2R were based on the lack of the (AB)(CD) tree topology amongst four members of a gene family in vertebrates. However, if the two genome duplications occurred close together, we would not expect to find this topology.<sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> A recent study generated the <a href="/wiki/Petromyzon_marinus" class="mw-redirect" title="Petromyzon marinus">sea lamprey</a> genetic map, which yielded strong support for the hypothesis that a single whole-genome duplication occurred in the basal vertebrate lineage, preceded and followed by several evolutionarily independent segmental duplications that occurred over chordate evolution.<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> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paleopolyploidy&action=edit&section=10" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Gene_duplication" title="Gene duplication">Gene duplication</a></li> <li><a href="/wiki/Genomics" title="Genomics">Genomics</a></li> <li><a href="/wiki/Karyotype" title="Karyotype">Karyotype</a></li> <li><a href="/wiki/Ploidy" title="Ploidy">Ploidy</a></li> <li><a href="/wiki/Polyploidy" title="Polyploidy">Polyploidy</a></li> <li><a href="/wiki/Speciation" title="Speciation">Speciation</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paleopolyploidy&action=edit&section=11" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style 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Series B, Biological Sciences</i>. <b>357</b> (1420): <span class="nowrap">531–</span>44. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1098%2Frstb.2001.1035">10.1098/rstb.2001.1035</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/PMC1692965">1692965</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/12028790">12028790</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Philosophical+Transactions+of+the+Royal+Society+of+London.+Series+B%2C+Biological+Sciences&rft.atitle=Were+vertebrates+octoploid%3F&rft.volume=357&rft.issue=1420&rft.pages=%3Cspan+class%3D%22nowrap%22%3E531-%3C%2Fspan%3E44&rft.date=2002-04&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1692965%23id-name%3DPMC&rft_id=info%3Apmid%2F12028790&rft_id=info%3Adoi%2F10.1098%2Frstb.2001.1035&rft.aulast=Furlong&rft.aufirst=RF&rft.au=Holland%2C+PW&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1692965&rfr_id=info%3Asid%2Fen.wikipedia.org%3APaleopolyploidy" class="Z3988"></span></span> </li> <li id="cite_note-34"><span class="mw-cite-backlink"><b><a href="#cite_ref-34">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSmithKeinath2015" class="citation journal cs1">Smith JJ, Keinath MC (August 2015). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4509993">"The sea lamprey meiotic map improves resolution of ancient vertebrate genome duplications"</a>. <i>Genome Research</i>. <b>25</b> (8): <span class="nowrap">1081–</span>90. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1101%2Fgr.184135.114">10.1101/gr.184135.114</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/PMC4509993">4509993</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/26048246">26048246</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Genome+Research&rft.atitle=The+sea+lamprey+meiotic+map+improves+resolution+of+ancient+vertebrate+genome+duplications&rft.volume=25&rft.issue=8&rft.pages=%3Cspan+class%3D%22nowrap%22%3E1081-%3C%2Fspan%3E90&rft.date=2015-08&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4509993%23id-name%3DPMC&rft_id=info%3Apmid%2F26048246&rft_id=info%3Adoi%2F10.1101%2Fgr.184135.114&rft.aulast=Smith&rft.aufirst=JJ&rft.au=Keinath%2C+MC&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4509993&rfr_id=info%3Asid%2Fen.wikipedia.org%3APaleopolyploidy" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Paleopolyploidy&action=edit&section=12" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></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 refbegin-columns references-column-width" style="column-width: 32em"> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAdamsWendel2005" class="citation journal cs1">Adams KL, Wendel JF (April 2005). 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href="/wiki/Ploidy" title="Ploidy">Ploidy</a></li> <li><a href="/wiki/Genome" title="Genome">Genetic material</a>/<a href="/wiki/Genome" title="Genome">Genome</a></li> <li><a href="/wiki/Chromatin" title="Chromatin">Chromatin</a> <ul><li><a href="/wiki/Euchromatin" title="Euchromatin">Euchromatin</a></li> <li><a href="/wiki/Heterochromatin" title="Heterochromatin">Heterochromatin</a></li></ul></li> <li><a href="/wiki/Chromosome" title="Chromosome">Chromosome</a></li> <li><a href="/wiki/Chromatid" title="Chromatid">Chromatid</a></li> <li><a href="/wiki/Nucleosome" title="Nucleosome">Nucleosome</a></li> <li><a href="/wiki/Nuclear_organization" title="Nuclear organization">Nuclear organization</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Types</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/Autosome" title="Autosome">Autosome</a>/<a href="/wiki/Sex-determination_system" title="Sex-determination system">Sex chromosome</a> (or <a href="/wiki/Sex_chromosome" title="Sex chromosome">allosome</a> or heterosome)</li> <li><a href="/wiki/Microchromosome" title="Microchromosome">Macrochromosome/Microchromosome</a></li> <li><a href="/wiki/Circular_bacterial_chromosome" class="mw-redirect" title="Circular bacterial chromosome">Circular chromosome</a>/<a href="/wiki/Linear_chromosome" title="Linear chromosome">Linear chromosome</a></li> <li><a href="/wiki/Aneuploidy" title="Aneuploidy">Extra chromosome</a> (or accessory chromosome)</li> <li><a href="/wiki/Small_supernumerary_marker_chromosome" title="Small supernumerary marker chromosome">Supernumerary chromosome</a></li> <li><a href="/wiki/B_chromosome" title="B chromosome">A chromosome/B chromosome</a></li> <li><a href="/wiki/Lampbrush_chromosome" title="Lampbrush chromosome">Lampbrush chromosome</a></li> <li><a href="/wiki/Polytene_chromosome" title="Polytene chromosome">Polytene chromosome</a></li> <li><a href="/wiki/Dinokaryon" title="Dinokaryon">Dinoflagellate chromosomes</a></li> <li><a href="/wiki/Homologous_chromosome" title="Homologous chromosome">Homologous chromosome</a></li> <li><a href="/wiki/Isochromosome" title="Isochromosome">Isochromosome</a></li> <li><a href="/wiki/Satellite_chromosome" title="Satellite chromosome">Satellite chromosome</a></li> <li><i>Centromere position</i> <ul><li><a href="/wiki/Centromere#Metacentric" title="Centromere">Metacentric</a></li> <li><a href="/wiki/Submetacentric" class="mw-redirect" title="Submetacentric">Submetacentric</a></li> <li><a href="/wiki/Telocentric" class="mw-redirect" title="Telocentric">Telocentric</a></li> <li><a href="/wiki/Acrocentric" class="mw-redirect" title="Acrocentric">Acrocentric</a></li> <li><a href="/wiki/Holocentric_chromosome" title="Holocentric chromosome">Holocentric</a></li></ul></li> <li><i>Centromere number</i> <ul><li><a href="/wiki/Acentric_fragment" title="Acentric fragment">Acentric</a></li> <li><a href="/wiki/Monocentric" class="mw-redirect" title="Monocentric">Monocentric</a></li> <li><a href="/wiki/Dicentric_chromosome" title="Dicentric chromosome">Dicentric</a></li> <li><a href="/wiki/Polycentric_chromosome" title="Polycentric chromosome">Polycentric</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Processes<br />and evolution</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/Mitosis" title="Mitosis">Mitosis</a></li> <li><a href="/wiki/Meiosis" title="Meiosis">Meiosis</a></li> <li><a href="/wiki/Chromosome_abnormality#Structural_abnormalities" title="Chromosome abnormality">Structural alterations</a> <ul><li><a href="/wiki/Chromosomal_inversion" title="Chromosomal inversion">Chromosomal inversion</a></li> <li><a href="/wiki/Chromosomal_translocation" title="Chromosomal translocation">Chromosomal translocation</a></li></ul></li> <li><a href="/wiki/Chromosome_abnormality#Numerical_disorders" title="Chromosome abnormality">Numerical alterations</a> <ul><li><a href="/wiki/Aneuploidy" title="Aneuploidy">Aneuploidy</a></li> <li><a href="/wiki/Ploidy" title="Ploidy">Euploidy</a></li> <li><a href="/wiki/Polyploidy" title="Polyploidy">Polyploidy</a></li> <li><a class="mw-selflink selflink">Paleopolyploidy</a></li></ul></li> <li><a href="/wiki/Polyploidization" class="mw-redirect" title="Polyploidization">Polyploidization</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Structures</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/Telomere" title="Telomere">Telomere</a>: <a href="/wiki/Telomere-binding_protein" title="Telomere-binding protein">Telomere-binding protein</a> (<a href="/wiki/TINF2" title="TINF2">TINF2</a>)</li> <li><a href="/wiki/Protamine" title="Protamine">Protamine</a></li></ul> </div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Histone" title="Histone">Histone</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/Histone_H1" title="Histone H1">H1</a></li> <li><a href="/wiki/Histone_H2A" title="Histone H2A">H2A</a></li> <li><a href="/wiki/Histone_H2B" title="Histone H2B">H2B</a></li> <li><a href="/wiki/Histone_H3" title="Histone H3">H3</a></li> <li><a href="/wiki/Histone_H4" title="Histone H4">H4</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Centromere" title="Centromere">Centromere</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/CENPA" title="CENPA">A</a></li> <li><a href="/wiki/Centromere_protein_B" title="Centromere protein B">B</a></li> <li><a href="/wiki/CENPC1" title="CENPC1">C1</a></li> <li><a href="/w/index.php?title=CENPC2&action=edit&redlink=1" class="new" title="CENPC2 (page does not exist)">C2</a></li> <li><a href="/wiki/Centromere_protein_E" title="Centromere protein E">E</a></li> <li><a href="/wiki/CENPF" title="CENPF">F</a></li> <li><a href="/wiki/CENPH" title="CENPH">H</a></li> <li><a href="/wiki/CENPI" title="CENPI">I</a></li> <li><a href="/wiki/CENPJ" title="CENPJ">J</a></li> <li><a href="/wiki/CENPK" title="CENPK">K</a></li> <li><a href="/wiki/CENPM" title="CENPM">M</a></li> <li><a href="/wiki/CENPN" title="CENPN">N</a></li> <li><a href="/wiki/CENPO" title="CENPO">O</a></li> <li><a href="/w/index.php?title=CENPP&action=edit&redlink=1" class="new" title="CENPP (page does not exist)">P</a></li> <li><a href="/w/index.php?title=CENPQ&action=edit&redlink=1" class="new" title="CENPQ (page does not exist)">Q</a></li> <li><a href="/wiki/CENPT" title="CENPT">T</a></li></ul> </div></td></tr></tbody></table><div> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">See also</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/Extrachromosomal_DNA" title="Extrachromosomal DNA">Extrachromosomal DNA</a> <ul><li><a href="/wiki/Plasmid" title="Plasmid">Plasmid</a></li></ul></li> <li><a href="/wiki/List_of_organisms_by_chromosome_count" title="List of organisms by chromosome count">List of organisms by chromosome count</a></li> <li><a href="/wiki/List_of_sequenced_eukaryotic_genomes" title="List of sequenced eukaryotic genomes">List of sequenced genomes</a></li> <li><a href="/wiki/International_System_for_Human_Cytogenetic_Nomenclature" class="mw-redirect" title="International System for Human Cytogenetic Nomenclature">International System for Human Cytogenetic Nomenclature</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="Speciation841" 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:Speciation" title="Template:Speciation"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Speciation" title="Template talk:Speciation"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Speciation" title="Special:EditPage/Template:Speciation"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Speciation841" style="font-size:114%;margin:0 4em"><a href="/wiki/Speciation" title="Speciation">Speciation</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/w/index.php?title=Introduction_to_speciation&action=edit&redlink=1" class="new" title="Introduction to speciation (page does not exist)">Introduction</a></li> <li><a href="/wiki/History_of_speciation" title="History of speciation">History</a></li> <li><a href="/wiki/Laboratory_experiments_of_speciation" title="Laboratory experiments of speciation">Laboratory experiments</a></li> <li><a href="/wiki/Glossary_of_speciation" class="mw-redirect" title="Glossary of speciation">Glossary</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Basic concepts</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/Species" title="Species">Species</a> (<a href="/wiki/Species_complex" title="Species complex">Species complex</a>)</li> <li><a href="/wiki/Reproductive_isolation" title="Reproductive isolation">Reproductive isolation</a></li> <li><a href="/wiki/Anagenesis" title="Anagenesis">Anagenesis</a></li> <li><a href="/wiki/Cladogenesis" title="Cladogenesis">Cladogenesis</a></li> <li><a href="/wiki/Cospeciation" title="Cospeciation">Cospeciation</a></li> <li><a href="/wiki/Evidence_of_evolution" class="mw-redirect" title="Evidence of evolution">Evidence of evolution</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Geographic modes</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/Allopatric_speciation" title="Allopatric speciation">Allopatric</a> (<a href="/wiki/Peripatric_speciation" title="Peripatric speciation">Peripatric</a> · <a href="/wiki/Peripatric_speciation#Quantum_and_budding_speciation" title="Peripatric speciation">Quantum</a> · <a href="/wiki/Peripatric_speciation#Centrifugal_speciation" title="Peripatric speciation">Centrifugal</a> · <a href="/wiki/Founder_effect" title="Founder effect">Founder effect</a>)</li> <li><a href="/wiki/Parapatric_speciation" title="Parapatric speciation">Parapatric</a> (<a href="/wiki/Cline_(biology)" title="Cline (biology)">Clines</a> · <a href="/wiki/Ring_species" title="Ring species">Ring species</a>)</li> <li><a href="/wiki/Sympatric_speciation" title="Sympatric speciation">Sympatric</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Reproductive_isolation" title="Reproductive isolation">Isolating factors</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/Adaptation" title="Adaptation">Adaptation</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Sexual_selection" title="Sexual selection">Sexual selection</a></li> <li><a href="/wiki/Ecological_speciation" title="Ecological speciation">Ecological speciation</a> (<a href="/wiki/Parallel_speciation" title="Parallel speciation">Parallel speciation</a> · <a href="/wiki/Allochronic_speciation" title="Allochronic speciation">Allochrony</a>)</li> <li><a href="/wiki/Nonecological_speciation" title="Nonecological speciation">Nonecological speciation</a></li> <li><a href="/wiki/Assortative_mating" title="Assortative mating">Assortative mating</a></li> <li><a href="/wiki/Haldane%27s_rule" title="Haldane's rule">Haldane's rule</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Hybrid_(biology)" title="Hybrid (biology)">Hybrid concepts</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/Hybrid_speciation" title="Hybrid speciation">Hybrid speciation</a> (<a href="/wiki/Polyploidy" title="Polyploidy">Polyploidy</a> · <a href="/wiki/Klepton" title="Klepton">Klepton</a> · <a href="/w/index.php?title=Recombinational_speciation&action=edit&redlink=1" class="new" title="Recombinational speciation (page does not exist)">Recombination</a>)</li> <li><a href="/wiki/Reinforcement_(speciation)" title="Reinforcement (speciation)">Reinforcement</a> (<a href="/wiki/Evidence_for_speciation_by_reinforcement" title="Evidence for speciation by reinforcement">evidence</a>)</li> <li><a href="/wiki/Secondary_contact" title="Secondary contact">Secondary contact</a></li> <li><a href="/wiki/Character_displacement" title="Character displacement">Character displacement</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Speciation in taxa</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="/w/index.php?title=Speciation_in_birds&action=edit&redlink=1" class="new" title="Speciation in birds (page does not exist)">Birds</a></li> <li><a href="/w/index.php?title=Speciation_in_fish&action=edit&redlink=1" class="new" title="Speciation in fish (page does not exist)">Fish</a></li> <li><a href="/w/index.php?title=Speciation_in_insects&action=edit&redlink=1" class="new" title="Speciation in insects (page does not exist)">Insects</a></li> <li><a href="/w/index.php?title=Speciation_in_plants&action=edit&redlink=1" class="new" title="Speciation in plants (page does not exist)">Plants</a></li> <li><a href="/w/index.php?title=Speciation_in_the_fossil_record&action=edit&redlink=1" class="new" title="Speciation in the fossil record (page does not exist)">Fossils</a> (<a class="mw-selflink selflink">Paleopolyploidy</a> · <a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a> · <a href="/wiki/Macroevolution" title="Macroevolution">Macroevolution</a> · <a href="/wiki/Chronospecies" title="Chronospecies">Chronospecies</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> <b><a href="/wiki/Category:Speciation" title="Category:Speciation">Category</a></b></li> <li><span class="noviewer" typeof="mw:File"><span title="Commons page"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/12px-Commons-logo.svg.png" decoding="async" width="12" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/18px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/24px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></span></span> <b><a href="https://commons.wikimedia.org/wiki/Category:Speciation" class="extiw" title="commons:Category:Speciation">Commons</a></b></li> <li><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Symbol_portal_class.svg" class="mw-file-description" title="Portal"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/16px-Symbol_portal_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/23px-Symbol_portal_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/31px-Symbol_portal_class.svg.png 2x" data-file-width="180" data-file-height="185" /></a></span> <b><a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Evolutionary biology Portal</a></b></li> <li><span class="noviewer" typeof="mw:File"><span title="WikiProject"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/People_icon.svg/16px-People_icon.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/People_icon.svg/24px-People_icon.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/37/People_icon.svg/32px-People_icon.svg.png 2x" data-file-width="100" data-file-height="100" /></span></span> <b><a href="/wiki/Wikipedia:WikiProject_Evolutionary_biology" title="Wikipedia:WikiProject Evolutionary biology">WikiProject</a></b></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="Genomics334" 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:Genomics" title="Template:Genomics"><abbr title="View this 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href="/wiki/Functional_genomics" title="Functional genomics">Functional genomics</a></li> <li><a href="/wiki/Genome_project" title="Genome project">Genome project</a> <ul><li><a href="/wiki/Human_Genome_Project" title="Human Genome Project">Human Genome Project</a></li></ul></li> <li><a href="/wiki/Metagenomics" title="Metagenomics">Metagenomics</a> <ul><li><a href="/wiki/Human_Microbiome_Project" title="Human Microbiome Project">Human Microbiome Project</a></li></ul></li> <li><a href="/wiki/Oncogenomics" title="Oncogenomics">Oncogenomics</a></li> <li><a href="/wiki/Pangenomics" class="mw-redirect" title="Pangenomics">Pangenomics</a></li> <li><a href="/wiki/Personal_genomics" title="Personal genomics">Personal genomics</a></li> <li><a href="/wiki/Population_genomics" title="Population genomics">Population genomics</a></li> <li><a href="/wiki/Sociogenomics" title="Sociogenomics"><i>Socio</i>genomics</a></li> <li><a href="/wiki/Structural_genomics" title="Structural genomics">Structural genomics</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Bioinformatics" title="Bioinformatics">Bioinformatics</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/Biochip" title="Biochip">Biochip</a></li> <li><a href="/wiki/Cheminformatics" title="Cheminformatics">Cheminformatics</a></li> <li><a href="/wiki/Chemogenomics" title="Chemogenomics">Chemogenomics</a></li> <li><a href="/wiki/Connectomics" title="Connectomics">Connectomics</a> <ul><li><a href="/wiki/Human_Connectome_Project" title="Human Connectome Project">Human Connectome Project</a></li></ul></li> <li><a href="/wiki/Epigenomics" title="Epigenomics">Epigenomics</a> <ul><li><a href="/wiki/Human_Epigenome_Project" title="Human Epigenome Project">Human Epigenome Project</a></li></ul></li> <li><a href="/wiki/Glycomics" title="Glycomics">Glycomics</a></li> <li><a href="/wiki/Immunomics" title="Immunomics">Immunomics</a></li> <li><a href="/wiki/Lipidomics" title="Lipidomics">Lipidomics</a></li> <li><a href="/wiki/Metabolomics" title="Metabolomics">Metabolomics</a></li> <li><a href="/wiki/Microbiome" title="Microbiome">Microbiomics</a></li> <li><a href="/wiki/Nutrigenomics" class="mw-redirect" title="Nutrigenomics">Nutrigenomics</a></li> <li><a class="mw-selflink selflink">Paleopolyploidy</a></li> <li><a href="/wiki/Pharmacogenetics" class="mw-redirect" title="Pharmacogenetics">Pharmacogenetics</a></li> <li><a href="/wiki/Pharmacogenomics" title="Pharmacogenomics">Pharmacogenomics</a></li> <li><a href="/wiki/Systems_biology" title="Systems biology">Systems biology</a></li> <li><a href="/wiki/Toxicogenomics" title="Toxicogenomics">Toxicogenomics</a></li> <li><a href="/wiki/Transcriptomics" class="mw-redirect" title="Transcriptomics">Transcriptomics</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Structural_biology" title="Structural biology">Structural biology</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/Proteomics" title="Proteomics">Proteomics</a> <ul><li><a href="/wiki/Human_proteome_project" class="mw-redirect" title="Human proteome project">Human proteome project</a></li></ul></li> <li><a href="/wiki/Call-map_proteomics" class="mw-redirect" title="Call-map proteomics">Call-map proteomics</a></li> <li><a href="/wiki/Structure-based_drug_design" class="mw-redirect" title="Structure-based drug design">Structure-based drug design</a></li> <li><a href="/wiki/Expression_proteomics" class="mw-redirect" title="Expression proteomics">Expression proteomics</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Research tools</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/2-D_electrophoresis" class="mw-redirect" title="2-D electrophoresis">2-D electrophoresis</a></li> <li><a href="/wiki/Mass_spectrometer" class="mw-redirect" title="Mass spectrometer">Mass spectrometer</a></li> <li><a href="/wiki/Electrospray_ionization" title="Electrospray ionization">Electrospray ionization</a></li> <li><a href="/wiki/Matrix-assisted_laser_desorption_ionization" class="mw-redirect" title="Matrix-assisted laser desorption ionization">Matrix-assisted laser desorption ionization</a></li> <li><a href="/wiki/Matrix-assisted_laser_desorption_ionization-time_of_flight_mass_spectrometer" class="mw-redirect" title="Matrix-assisted laser desorption ionization-time of flight mass spectrometer">Matrix-assisted laser desorption ionization-time of flight mass spectrometer</a></li> <li><a href="/wiki/Microfluidic-based_tools" class="mw-redirect" title="Microfluidic-based tools">Microfluidic-based tools</a></li> <li><a href="/wiki/Isotope_affinity_tags" class="mw-redirect" title="Isotope affinity tags">Isotope affinity tags</a></li> <li><a href="/wiki/Chromosome_conformation_capture" title="Chromosome conformation capture">Chromosome conformation capture</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Organizations</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/DNA_Data_Bank_of_Japan" title="DNA Data Bank of Japan">DNA Data Bank of Japan</a> (JP)</li> <li><a href="/wiki/European_Molecular_Biology_Laboratory" title="European Molecular Biology Laboratory">European Molecular Biology Laboratory</a> (EU)</li> <li><a href="/wiki/National_Institutes_of_Health" title="National Institutes of Health">National Institutes of Health</a> (USA)</li> <li><a href="/wiki/Wellcome_Sanger_Institute" title="Wellcome Sanger Institute">Wellcome Sanger Institute</a> (UK)</li></ul> 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