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Prophage - Wikipedia
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Available in 18 languages" > <label id="p-lang-btn-label" for="p-lang-btn-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--action-progressive mw-portlet-lang-heading-18" aria-hidden="true" ><span class="vector-icon mw-ui-icon-language-progressive mw-ui-icon-wikimedia-language-progressive"></span> <span class="vector-dropdown-label-text">18 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%B7%D9%84%D9%8A%D8%B9%D8%A9_%D8%A7%D9%84%D8%B9%D8%A7%D8%AB%D9%8A%D8%A9" title="طليعة العاثية – Arabic" lang="ar" hreflang="ar" data-title="طليعة العاثية" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Prophage" title="Prophage – German" lang="de" hreflang="de" data-title="Prophage" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Profago" title="Profago – Spanish" lang="es" hreflang="es" data-title="Profago" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Profago" title="Profago – Basque" lang="eu" hreflang="eu" data-title="Profago" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Prophage" title="Prophage – French" lang="fr" hreflang="fr" data-title="Prophage" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%ED%94%84%EB%A1%9C%ED%8C%8C%EC%A7%80" title="프로파지 – Korean" lang="ko" hreflang="ko" data-title="프로파지" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Profago" title="Profago – Italian" lang="it" hreflang="it" data-title="Profago" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/%E1%83%9E%E1%83%A0%E1%83%9D%E1%83%A4%E1%83%90%E1%83%92%E1%83%98" title="პროფაგი – Georgian" lang="ka" hreflang="ka" data-title="პროფაგი" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target"><span>ქართული</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%9F%D1%80%D0%BE%D1%84%D0%B0%D0%B3" title="Профаг – Kazakh" lang="kk" hreflang="kk" data-title="Профаг" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-ky mw-list-item"><a href="https://ky.wikipedia.org/wiki/%D0%9F%D1%80%D0%BE%D1%84%D0%B0%D0%B3" title="Профаг – Kyrgyz" lang="ky" hreflang="ky" data-title="Профаг" data-language-autonym="Кыргызча" data-language-local-name="Kyrgyz" class="interlanguage-link-target"><span>Кыргызча</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E3%83%97%E3%83%AD%E3%83%95%E3%82%A1%E3%83%BC%E3%82%B8" title="プロファージ – Japanese" lang="ja" hreflang="ja" data-title="プロファージ" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-uz mw-list-item"><a href="https://uz.wikipedia.org/wiki/Profag" title="Profag – Uzbek" lang="uz" hreflang="uz" data-title="Profag" data-language-autonym="Oʻzbekcha / ўзбекча" data-language-local-name="Uzbek" class="interlanguage-link-target"><span>Oʻzbekcha / ўзбекча</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Profag" title="Profag – Polish" lang="pl" hreflang="pl" data-title="Profag" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Profago" title="Profago – Portuguese" lang="pt" hreflang="pt" data-title="Profago" 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<div class="vector-body-before-content"> <div class="mw-indicators"> </div> <div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Bacteriophage genome that is integrated into a bacterial cell</div> <p class="mw-empty-elt"> </p> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Prophage.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b3/Prophage.svg/225px-Prophage.svg.png" decoding="async" width="225" height="250" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b3/Prophage.svg/338px-Prophage.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b3/Prophage.svg/450px-Prophage.svg.png 2x" data-file-width="540" data-file-height="600" /></a><figcaption>Formation of a prophage</figcaption></figure><p>A <b>prophage</b> is a <a href="/wiki/Bacteriophage" title="Bacteriophage">bacteriophage</a> (often shortened to "phage") <a href="/wiki/Genome" title="Genome">genome</a> that is integrated into the <a href="/wiki/Circular_bacterial_chromosome" class="mw-redirect" title="Circular bacterial chromosome">circular bacterial chromosome</a> or exists as an extrachromosomal <a href="/wiki/Plasmid" title="Plasmid">plasmid</a> within the bacterial <a href="/wiki/Cell_(biology)" title="Cell (biology)">cell</a>.<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> Integration of prophages into the bacterial host is the characteristic step of the <a href="/wiki/Lysogenic_cycle" title="Lysogenic cycle">lysogenic cycle</a> of <a href="/wiki/Temperateness_(virology)" title="Temperateness (virology)">temperate</a> phages. Prophages remain <a href="/wiki/Viral_latency" class="mw-redirect" title="Viral latency">latent</a> in the genome through multiple cell divisions until activation by an external factor, such as UV light, leading to production of new phage particles that will <a href="/wiki/Lysis" title="Lysis">lyse</a> the cell and spread. As ubiquitous <a href="/wiki/Mobile_genetic_elements" title="Mobile genetic elements">mobile genetic elements</a>, prophages play important roles in bacterial genetics and evolution, such as in the acquisition of <a href="/wiki/Virulence" title="Virulence">virulence</a> factors. </p><meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Prophage_induction">Prophage induction</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Prophage&action=edit&section=1" title="Edit section: Prophage induction"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Upon detection of host cell damage by UV light or certain chemicals, the prophage is excised from the bacterial chromosome in a process called prophage induction. After induction, viral replication begins via the <a href="/wiki/Lytic_cycle" title="Lytic cycle">lytic cycle</a>. In the lytic cycle, the virus commandeers the cell's reproductive machinery. The cell may fill with new viruses until it lyses or bursts, or it may release the new viruses one at a time in an exocytotic process. The period from infection to lysis is termed the latent period. A virus following a lytic cycle is called a <a href="/wiki/Virulent" class="mw-redirect" title="Virulent">virulent</a> virus. Prophages are important agents of <a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">horizontal gene transfer</a>, and are considered part of the <a href="/wiki/Mobilome" title="Mobilome">mobilome</a>. Genes are transferred via <a href="/wiki/Transduction_(genetics)" title="Transduction (genetics)">transduction</a> as the prophage genome is imperfectly excised from the host chromosome and integrated into a new host (specialized transduction) or as fragments of host DNA are packaged into the phage particles and introduced into a new host (generalized transduction).<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> All families of bacterial viruses that have circular (single-stranded or double-stranded) DNA genomes or replicate their genomes through <a href="/wiki/Rolling_circle_replication" title="Rolling circle replication">rolling circle replication</a> (e.g., <a href="/wiki/Caudovirales" class="mw-redirect" title="Caudovirales">Caudovirales</a>) have temperate members.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>During infections by the bacterial pathogen <i><a href="/wiki/Clostridioides_difficile" title="Clostridioides difficile">Clostridioides difficile</a></i>, spontaneous prophage release from the bacterial <a href="/wiki/Chromosome" title="Chromosome">chromosome</a> is common.<sup id="cite_ref-Schuler2024_4-0" class="reference"><a href="#cite_note-Schuler2024-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> The presence of <a href="/wiki/Deoxycholic_acid" title="Deoxycholic acid">deoxycholic acid</a> in the intestinal environment can promote induction of <i>C. difficle</i> <a href="/wiki/Biofilm" title="Biofilm">biofilm</a> formation as well as the induction of prophage release.<sup id="cite_ref-Schuler2024_4-1" class="reference"><a href="#cite_note-Schuler2024-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Zygotic_induction">Zygotic induction</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Prophage&action=edit&section=2" title="Edit section: Zygotic induction"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Zygotic_induction" title="Zygotic induction">Zygotic induction</a></div> <p>Zygotic induction occurs when a bacterial cell carrying the DNA of a bacterial virus transfers its own DNA along with the viral DNA (prophage) into the new host cell. This has the effect of causing the host cell to break apart.<sup id="cite_ref-Griffiths_2002_5-0" class="reference"><a href="#cite_note-Griffiths_2002-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> The DNA of the bacterial cell is silenced before entry into the cell by a <a href="/wiki/Repressor_protein" class="mw-redirect" title="Repressor protein">repressor protein</a> which is encoded for by the prophage. Upon the transfer of the bacterial cell's DNA into the host cell, the repressor protein is no longer encoded for, and the bacterial cell's original DNA is then turned on in the host cell. This mechanism eventually will lead to the release of the virus as the host cell splits open and the viral DNA is able to spread.<sup id="cite_ref-Griffiths_2002_5-1" class="reference"><a href="#cite_note-Griffiths_2002-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> This new discovery provided key insights into <a href="/wiki/Bacterial_conjugation" title="Bacterial conjugation">bacterial conjugation</a> and contributed to the early repression model of gene regulation, which provided an explanation as to how the <a href="/wiki/Lac_operon" title="Lac operon">lac operon</a> and λ <a href="/wiki/Bacteriophage" title="Bacteriophage">bacteriophage</a> genes are negatively regulated.<sup id="cite_ref-Blanco_1973_6-0" class="reference"><a href="#cite_note-Blanco_1973-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Prophage_reactivation">Prophage reactivation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Prophage&action=edit&section=3" title="Edit section: Prophage reactivation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Lambda_phage" title="Lambda phage">Bacteriophage λ</a> is able to undergo a type of <a href="/wiki/Homologous_recombination" title="Homologous recombination">recombinational</a> repair called prophage reactivation.<sup id="cite_ref-Blanco_1973_6-1" class="reference"><a href="#cite_note-Blanco_1973-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid6261109_7-0" class="reference"><a href="#cite_note-pmid6261109-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> Prophage reactivation can occur by recombination between a UV-damaged infecting phage λ chromosome and a homologous phage <a href="/wiki/Genome" title="Genome">genome</a> integrated into the bacterial DNA and existing in a prophage state. Prophage reactivation in the case of phage λ appears to be an accurate recombinational repair process that is mediated by the <a href="/wiki/RecA" title="RecA">recA</a>+ and red+ gene products.<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. (October 2022)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading2"><h2 id="Cost/benefit_to_the_host"><span id="Cost.2Fbenefit_to_the_host"></span>Cost/benefit to the host</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Prophage&action=edit&section=4" title="Edit section: Cost/benefit to the host"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Lysis of host cells during prophage induction can cause the collapse of a microbial population.<sup id="cite_ref-Haaber_2016_8-0" class="reference"><a href="#cite_note-Haaber_2016-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Hu_2021_9-0" class="reference"><a href="#cite_note-Hu_2021-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> On the other hand, induction, transduction and superinfection exclusion mechanisms confer many beneficial functions to the host. Induction of prophages allows hosts to compete in the microbial ecology by infecting and lysing susceptible bacteria.<sup id="cite_ref-Wendling_2021_10-0" class="reference"><a href="#cite_note-Wendling_2021-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Phages also enable the host to pick up and integrate antibiotic resistance genes from nearby cells.<sup id="cite_ref-Hu_2021_9-1" class="reference"><a href="#cite_note-Hu_2021-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Wendling_2021_10-1" class="reference"><a href="#cite_note-Wendling_2021-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Haaber_2016_8-1" class="reference"><a href="#cite_note-Haaber_2016-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Fortier_2013_11-0" class="reference"><a href="#cite_note-Fortier_2013-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Additionally, phages can enable the host to acquire virulence and pathogenicity genes.<sup id="cite_ref-Hu_2021_9-2" class="reference"><a href="#cite_note-Hu_2021-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Fortier_2013_11-1" class="reference"><a href="#cite_note-Fortier_2013-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Modulation of biofilm formation is also affected by infection by lysogenic phages.<sup id="cite_ref-Fortier_2013_11-2" class="reference"><a href="#cite_note-Fortier_2013-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Superinfection exclusion, or protection against infection by multiple phages, can be conferred by prophage integration.<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> Additionally, phage-mediated recombination mechanisms may remodel the host chromosome and provide new ways for cells to regulate metabolism and gene expression, such as those involved in sporulation and competence.<sup id="cite_ref-Fortier_2013_11-3" class="reference"><a href="#cite_note-Fortier_2013-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><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> <div class="mw-heading mw-heading2"><h2 id="Applications">Applications</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Prophage&action=edit&section=5" title="Edit section: Applications"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Prophages can tell researchers a lot about the relationship between a bacterium and a host.<sup id="cite_ref-Canchaya-et-al-2003_14-0" class="reference"><a href="#cite_note-Canchaya-et-al-2003-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> With data from more nonpathogenic bacteria, researchers will be able to gather evidence as to whether or not prophages contribute to the survival value of the host. Prophage genomics has the potential to lead to ecological adaptations of the relationships between bacteria.<sup id="cite_ref-Canchaya-et-al-2003_14-1" class="reference"><a href="#cite_note-Canchaya-et-al-2003-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> Another important area of interest is the control of prophage gene expression with many of the lysogenic conversion genes (<a href="/wiki/Gene_conversion" title="Gene conversion">gene conversion</a>) being tightly regulated.<sup id="cite_ref-Feiner-et-al-2015_15-0" class="reference"><a href="#cite_note-Feiner-et-al-2015-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> This process is capable of converting non-pathogenic bacteria into pathogenic bacteria that can now produce harmful toxins<sup id="cite_ref-Feiner-et-al-2015_15-1" class="reference"><a href="#cite_note-Feiner-et-al-2015-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> such as in <a href="/wiki/Staph_infections" class="mw-redirect" title="Staph infections">staph infections</a>. Since the specific mechanisms of prophage are not yet detailed, this research could provide the community with this tool for future research.<sup id="cite_ref-Canchaya-et-al-2003_14-2" class="reference"><a href="#cite_note-Canchaya-et-al-2003-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Economic_impact">Economic impact</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Prophage&action=edit&section=6" title="Edit section: Economic impact"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Exotoxins" class="mw-redirect" title="Exotoxins">Exotoxins</a> encoded by prophages cause pathogenic outcomes in <a href="/wiki/Agricultural_microbiology" title="Agricultural microbiology">agriculture</a> and <a href="/wiki/Aquaculture" title="Aquaculture">aquaculture</a>.<sup id="cite_ref-Cobian-Guemes-et-al-2016_16-0" class="reference"><a href="#cite_note-Cobian-Guemes-et-al-2016-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p> <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=Prophage&action=edit&section=7" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFSaussereauDebarbieux2012" class="citation book cs1">Saussereau E, Debarbieux L (2012). 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title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annual+Review+of+Virology&rft.atitle=Viruses+as+Winners+in+the+Game+of+Life&rft.volume=3&rft.issue=1&rft.pages=%3Cspan+class%3D%22nowrap%22%3E197-%3C%2Fspan%3E214&rft.date=2016-09&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A36517589%23id-name%3DS2CID&rft_id=info%3Apmid%2F27741409&rft_id=info%3Adoi%2F10.1146%2Fannurev-virology-100114-054952&rft.aulast=Cobi%C3%A1n+G%C3%BCemes&rft.aufirst=AG&rft.au=Youle%2C+M&rft.au=Cant%C3%BA%2C+VA&rft.au=Felts%2C+B&rft.au=Nulton%2C+J&rft.au=Rohwer%2C+F&rfr_id=info%3Asid%2Fen.wikipedia.org%3AProphage" class="Z3988"></span></span> </li> </ol></div></div> <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=Prophage&action=edit&section=8" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output .side-box-text{flex:1;min-width:0}}@media(min-width:720px){.mw-parser-output .side-box{width:238px}.mw-parser-output .side-box-right{clear:right;float:right;margin-left:1em}.mw-parser-output .side-box-left{margin-right:1em}}</style><style data-mw-deduplicate="TemplateStyles:r1237033735">@media print{body.ns-0 .mw-parser-output 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style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bacteria" title="Bacteria">Bacteria</a></li> <li><a href="/wiki/Archaea" title="Archaea">Archaea</a></li> <li><a href="/wiki/Eukaryote" title="Eukaryote">Eukaryota</a> <ul><li><a href="/wiki/Animal" title="Animal">Animalia</a></li> <li><a href="/wiki/Fungus" title="Fungus">Fungi</a></li> <li><a href="/wiki/Plant" title="Plant">Plantae</a></li> <li><a href="/wiki/Protist" title="Protist">Protista</a></li></ul></li> <li><i><a href="/wiki/Incertae_sedis" title="Incertae sedis">Incertae sedis</a></i> <ul><li><i><a href="/wiki/Parakaryon" title="Parakaryon">Parakaryon</a></i></li> <li><a href="/wiki/Biological_dark_matter" title="Biological dark matter">Biological dark matter</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%;background:#CEDAF2;"><a href="/wiki/Virus" title="Virus">Virus</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/DNA_virus#Group_I:_dsDNA_viruses" title="DNA virus">dsDNA virus</a> <ul><li><a href="/wiki/Giant_virus" title="Giant virus">Giant virus</a></li></ul></li> <li><a href="/wiki/DNA_virus#Group_II:_ssDNA_viruses" title="DNA virus">ssDNA virus</a></li> <li><a href="/wiki/Double-stranded_RNA_viruses" title="Double-stranded RNA viruses">dsRNA virus</a></li> <li><a href="/wiki/RNA_virus#Group_IV—positive-sense_ssRNA_viruses" title="RNA virus">(+)ssRNA virus</a></li> <li><a href="/wiki/RNA_virus#Group_V—negative-sense_ssRNA_viruses" title="RNA virus">(−)ssRNA virus</a></li> <li><a href="/wiki/Retrovirus" title="Retrovirus">ssRNA-RT virus</a></li> <li><a href="/wiki/DsDNA-RT_virus" class="mw-redirect" title="DsDNA-RT virus">dsDNA-RT virus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%;background:#CEDAF2;"><a href="/wiki/Subviral_agents" class="mw-redirect" title="Subviral agents">Subviral<br />agents</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;;background:#CEDAF2;"><a href="/wiki/Viroid" title="Viroid">Viroid</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a></i></li> <li><i><a href="/wiki/Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a></i></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;;background:#CEDAF2;"><a href="/wiki/Helper_virus" title="Helper virus">Helper-virus<br />dependent</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;;background:#CEDAF2;"><a href="/wiki/Satellite_(biology)" title="Satellite (biology)">Satellite</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>ssRNA satellite virus</li> <li>dsDNA satellite virus (<a href="/wiki/Virophage" title="Virophage">Virophage</a>)</li> <li>ssDNA satellite virus</li> <li>ssDNA satellite</li> <li>dsRNA satellite</li> <li>ssRNA satellite (<a href="/wiki/Virusoid" title="Virusoid">Virusoid</a>)</li> <li>Satellite-like nucleic acids <ul><li>RNA</li> <li>DNA</li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;;background:#CEDAF2;">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Defective_interfering_particle" title="Defective interfering particle">Defective interfering particle</a> <ul><li>RNA</li> <li>DNA</li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;"><a href="/wiki/Prion" title="Prion">Prion</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Prion" title="Prion">Mammalian prion</a></li> <li><a href="/wiki/Fungal_prion" title="Fungal prion">Fungal prion</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Nucleic_acid" title="Nucleic acid">Nucleic acid</a><br />self-replication</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;"><a href="/wiki/Mobile_genetic_elements" title="Mobile genetic elements">Mobile genetic<br />elements</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mobilome" title="Mobilome">Mobilome</a> <ul><li><a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">Horizontal gene transfer</a></li> <li><a href="/wiki/Genomic_island" title="Genomic island">Genomic island</a></li></ul></li> <li><a href="/wiki/Transposable_element" title="Transposable element">Transposable element</a> <ul><li><a href="/wiki/Retrotransposon" title="Retrotransposon">Class I or retrotransposon</a></li> <li><a href="/wiki/DNA_transposon" title="DNA transposon">Class II or DNA transposon</a></li></ul></li> <li><a href="/wiki/Plasmid" title="Plasmid">Plasmid</a> <ul><li><a href="/wiki/Fertility_factor_(bacteria)" class="mw-redirect" title="Fertility factor (bacteria)">Fertility</a></li> <li><a href="/wiki/R-factor" class="mw-redirect" title="R-factor">Resistance</a></li> <li><a href="/wiki/Colicin" title="Colicin">Col</a></li> <li>Degradative</li> <li><a href="/wiki/Virulence_factor" title="Virulence factor">Virulence</a>/<a href="/wiki/Ti_plasmid" title="Ti plasmid">Ti</a></li> <li>Cryptic</li></ul></li> <li><a href="/wiki/Cosmid" title="Cosmid">Cosmid</a> <ul><li><a href="/wiki/Fosmid" title="Fosmid">Fosmid</a></li></ul></li> <li><a href="/wiki/Phagemid" title="Phagemid">Phagemid</a></li> <li><a href="/wiki/Group_I_catalytic_intron" title="Group I catalytic intron">Group I intron</a></li> <li><a href="/wiki/Group_II_intron" title="Group II intron">Group II intron</a></li> <li><a href="/wiki/Retrozyme" title="Retrozyme">Retrozyme</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;">Other aspects</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a> <ul><li><a href="/wiki/RNA-dependent_RNA_polymerase" title="RNA-dependent RNA polymerase">RNA replication</a></li></ul></li> <li><a href="/wiki/Chromosome" title="Chromosome">Chromosome</a> <ul><li><a href="/wiki/Linear_chromosome" title="Linear chromosome">Linear</a></li> <li><a href="/wiki/Circular_chromosome" title="Circular chromosome">Circular</a></li> <li><a href="/wiki/Extrachromosomal_DNA" title="Extrachromosomal DNA">Extrachromosomal DNA</a></li> <li><a href="/wiki/Secondary_chromosome" title="Secondary chromosome">Secondary chromosome</a></li></ul></li> <li><a href="/wiki/Genome" title="Genome">Genome</a> <ul><li><a href="/wiki/Gene" title="Gene">Gene</a></li> <li><a href="/wiki/Gene_duplication" title="Gene duplication">Gene duplication</a></li> <li><a href="/wiki/Non-coding_DNA" title="Non-coding DNA">Non-coding DNA</a></li></ul></li> <li><a href="/wiki/Origin_of_replication" title="Origin of replication">Origin of replication</a> <ul><li><a href="/wiki/Replicon_(genetics)" title="Replicon (genetics)">Replicon</a></li></ul></li> <li><a href="/wiki/Endogenous_viral_element" title="Endogenous viral element">Endogenous viral element</a> <ul><li><a href="/wiki/Provirus" title="Provirus">Provirus</a></li> <li><a class="mw-selflink selflink">Prophage</a></li> <li><a href="/wiki/Endogenous_retrovirus" title="Endogenous retrovirus">Endogenous retrovirus</a></li> <li><a href="/wiki/Transpoviron" title="Transpoviron">Transpoviron</a></li></ul></li> <li><a href="/wiki/Repeated_sequence_(DNA)" title="Repeated sequence (DNA)">Repeated sequences in DNA</a> <ul><li><a href="/wiki/Tandem_repeat" title="Tandem repeat">Tandem repeat</a></li> <li><a href="/wiki/Interspersed_repeat" title="Interspersed repeat">Interspersed repeat</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Endosymbiont" title="Endosymbiont">Endosymbiosis</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mitochondrion" title="Mitochondrion">Mitochondrion</a> <ul><li><a href="/wiki/Mitosome" title="Mitosome">Mitosome</a></li> <li><a href="/wiki/Hydrogenosome" title="Hydrogenosome">Hydrogenosome</a></li></ul></li> <li><a href="/wiki/Plastid" title="Plastid">Plastid</a> <ul><li><a href="/wiki/Chloroplast" title="Chloroplast">Chloroplast</a></li> <li><a href="/wiki/Chromoplast" title="Chromoplast">Chromoplast</a></li> <li><a href="/wiki/Gerontoplast" title="Gerontoplast">Gerontoplast</a></li> <li><a href="/wiki/Leucoplast" title="Leucoplast">Leucoplast</a></li> <li><a href="/wiki/Apicoplast" title="Apicoplast">Apicoplast</a></li></ul></li> <li><a href="/wiki/Kappa_organism" title="Kappa organism">Kappa organism</a></li> <li>Organs <ul><li><a href="/wiki/Bacteriome" title="Bacteriome">Bacteriome</a></li> <li><a href="/wiki/Trophosome" title="Trophosome">Trophosome</a></li></ul></li> <li><a href="/wiki/Nitroplast" title="Nitroplast">Nitroplast</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Abiogenesis" title="Abiogenesis">Abiogenesis</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Last_universal_common_ancestor" title="Last universal common ancestor">Last universal common ancestor</a></li> <li><a href="/wiki/Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li> <li>?<a href="/wiki/RNA_world" title="RNA world">RNA life</a> <ul><li><a href="/wiki/Ribozyme" title="Ribozyme">Ribozyme</a></li></ul></li> <li>†<a href="/wiki/Protocell" title="Protocell">Protocell</a></li> <li><a href="/wiki/Coacervate" title="Coacervate">Coacervate</a></li> <li><a href="/wiki/Proteinoid" title="Proteinoid">Proteinoid</a></li> <li><a href="/wiki/Sulphobes" title="Sulphobes">Sulphobe</a></li> <li>Research <ul><li><a href="/wiki/Model_lipid_bilayer" title="Model lipid bilayer">Model lipid bilayer</a></li> <li><a href="/wiki/Jeewanu" title="Jeewanu">Jeewanu</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%">See also</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Organism" title="Organism">Organism</a></li> <li><a href="/wiki/Cell_(biology)" title="Cell (biology)">Cell</a> <ul><li><a href="/wiki/Cell_division" title="Cell division">Cell division</a></li> <li><a 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