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class="vector-toc-link" href="#Historical_discovery"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Historical discovery</span> </div> </a> <ul id="toc-Historical_discovery-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Replication,_partitioning_and_maintenance" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Replication,_partitioning_and_maintenance"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Replication, partitioning and maintenance</span> </div> </a> <button aria-controls="toc-Replication,_partitioning_and_maintenance-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Replication, partitioning and maintenance subsection</span> </button> <ul id="toc-Replication,_partitioning_and_maintenance-sublist" class="vector-toc-list"> <li id="toc-Replication_of_the_Ti_plasmid" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Replication_of_the_Ti_plasmid"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Replication of the Ti plasmid</span> </div> </a> <ul id="toc-Replication_of_the_Ti_plasmid-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Partitioning_of_the_Ti_plasmid" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Partitioning_of_the_Ti_plasmid"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Partitioning of the Ti plasmid</span> </div> </a> <ul id="toc-Partitioning_of_the_Ti_plasmid-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Maintenance_of_the_Ti_plasmid" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Maintenance_of_the_Ti_plasmid"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.3</span> <span>Maintenance of the Ti plasmid</span> </div> </a> <ul id="toc-Maintenance_of_the_Ti_plasmid-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Features" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Features"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Features</span> </div> </a> <button aria-controls="toc-Features-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Features subsection</span> </button> <ul id="toc-Features-sublist" class="vector-toc-list"> <li id="toc-Virulence_operon" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Virulence_operon"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Virulence operon</span> </div> </a> <ul id="toc-Virulence_operon-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Transfer_DNA_(T-DNA)" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Transfer_DNA_(T-DNA)"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2</span> <span>Transfer DNA (T-DNA)</span> </div> </a> <ul id="toc-Transfer_DNA_(T-DNA)-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Transfer_apparatus" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Transfer_apparatus"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.3</span> <span>Transfer apparatus</span> </div> </a> <ul id="toc-Transfer_apparatus-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Uses_in_bioengineering" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Uses_in_bioengineering"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Uses in bioengineering</span> </div> </a> <ul id="toc-Uses_in_bioengineering-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Similar_plasmids" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Similar_plasmids"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Similar plasmids</span> </div> </a> <button aria-controls="toc-Similar_plasmids-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Similar plasmids subsection</span> </button> <ul id="toc-Similar_plasmids-sublist" class="vector-toc-list"> <li id="toc-Root-inducing_(Ri)_plasmid" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Root-inducing_(Ri)_plasmid"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1</span> <span>Root-inducing (Ri) plasmid</span> </div> </a> <ul id="toc-Root-inducing_(Ri)_plasmid-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Symbiotic_(sym)_plasmids_of_Rhizobia" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Symbiotic_(sym)_plasmids_of_Rhizobia"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.2</span> <span>Symbiotic (sym) plasmids of Rhizobia</span> </div> </a> <ul id="toc-Symbiotic_(sym)_plasmids_of_Rhizobia-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of 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Available in 17 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-17" 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">17 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%AA%D9%8A_%D8%A8%D9%84%D8%A7%D8%B2%D9%85%D9%8A%D8%AF" title="تي بلازميد – Arabic" lang="ar" hreflang="ar" data-title="تي بلازميد" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Plasmidi_Ti" title="Plasmidi Ti – Catalan" lang="ca" hreflang="ca" data-title="Plasmidi Ti" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Ti_plasmid" title="Ti plasmid – Danish" lang="da" hreflang="da" data-title="Ti plasmid" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Ti-Plasmid" title="Ti-Plasmid – German" lang="de" hreflang="de" data-title="Ti-Plasmid" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%A0%CE%BB%CE%B1%CF%83%CE%BC%CE%AF%CE%B4%CE%B9%CE%BF_Ti" title="Πλασμίδιο Ti – Greek" lang="el" hreflang="el" data-title="Πλασμίδιο Ti" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Ti_plasmidoa" title="Ti plasmidoa – Basque" lang="eu" hreflang="eu" data-title="Ti plasmidoa" 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/Plasmide_Ti" title="Plasmide Ti – French" lang="fr" hreflang="fr" data-title="Plasmide Ti" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Pl%C3%A1smido_Ti" title="Plásmido Ti – Galician" lang="gl" hreflang="gl" data-title="Plásmido Ti" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%ED%8B%B0%EC%95%84%EC%9D%B4_%ED%94%8C%EB%9D%BC%EC%8A%A4%EB%AF%B8%EB%93%9C" title="티아이 플라스미드 – Korean" lang="ko" hreflang="ko" data-title="티아이 플라스미드" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Plasmid_Ti" title="Plasmid Ti – Indonesian" lang="id" hreflang="id" data-title="Plasmid Ti" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-jv mw-list-item"><a href="https://jv.wikipedia.org/wiki/Plasmid_Ti" title="Plasmid Ti – Javanese" lang="jv" hreflang="jv" data-title="Plasmid Ti" data-language-autonym="Jawa" data-language-local-name="Javanese" class="interlanguage-link-target"><span>Jawa</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Ti-plasmide" title="Ti-plasmide – Dutch" lang="nl" hreflang="nl" data-title="Ti-plasmide" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/Ti%E3%83%97%E3%83%A9%E3%82%B9%E3%83%9F%E3%83%89" title="Tiプラスミド – Japanese" lang="ja" hreflang="ja" data-title="Tiプラスミド" 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-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Plazmid_Ti" title="Plazmid Ti – Polish" lang="pl" hreflang="pl" data-title="Plazmid Ti" 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/Plasm%C3%ADdeo_Ti" title="Plasmídeo Ti – Portuguese" lang="pt" hreflang="pt" data-title="Plasmídeo Ti" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ru badge-Q17559452 badge-recommendedarticle mw-list-item" title="recommended article"><a href="https://ru.wikipedia.org/wiki/Ti-%D0%BF%D0%BB%D0%B0%D0%B7%D0%BC%D0%B8%D0%B4%D0%B0" title="Ti-плазмида – Russian" lang="ru" hreflang="ru" data-title="Ti-плазмида" data-language-autonym="Русский" data-language-local-name="Russian" 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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">Circular plasmid used in creation of transgenic plants</div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Ti_plasmid.svg" class="mw-file-description"><img alt="A diagram showing the structure of the Ti plasmid, with various important regions labeled" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d1/Ti_plasmid.svg/350px-Ti_plasmid.svg.png" decoding="async" width="350" height="265" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d1/Ti_plasmid.svg/525px-Ti_plasmid.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d1/Ti_plasmid.svg/700px-Ti_plasmid.svg.png 2x" data-file-width="1035" data-file-height="783" /></a><figcaption>The structure of the Ti plasmid</figcaption></figure> <p>A <b>tumour inducing (Ti)</b> <a href="/wiki/Plasmid" title="Plasmid">plasmid</a> is a plasmid found in pathogenic species of <i><a href="/wiki/Agrobacterium" title="Agrobacterium">Agrobacterium</a></i>, including <a href="/wiki/Agrobacterium_tumefaciens" title="Agrobacterium tumefaciens"><i>A. tumefaciens</i></a>, <a href="/wiki/Rhizobium_rhizogenes" title="Rhizobium rhizogenes"><i>A. rhizogenes</i></a>, <i>A. rubi</i> and <a href="/wiki/Allorhizobium_vitis" title="Allorhizobium vitis"><i>A. vitis</i></a>. </p><p>Evolutionarily, the Ti plasmid is part of a family of plasmids carried by many species of <a href="/wiki/Alphaproteobacteria" title="Alphaproteobacteria">Alphaproteobacteria</a>. Members of this plasmid family are defined by the presence of a conserved DNA region known as the <i>repABC</i> gene cassette, which mediates the <a href="/wiki/DNA_replication" title="DNA replication">replication</a> of the plasmid, the partitioning of the plasmid into daughter cells during <a href="/wiki/Cell_division" title="Cell division">cell division</a> as well as the maintenance of the plasmid at low copy numbers in a cell.<sup id="cite_ref-Gordon2014_1-0" class="reference"><a href="#cite_note-Gordon2014-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The Ti plasmids themselves are sorted into different categories based on the type of molecule, or <a href="/wiki/Opine" title="Opine">opine</a>, they allow the bacteria to break down as an energy source.<sup id="cite_ref-Hooykaas1994_2-0" class="reference"><a href="#cite_note-Hooykaas1994-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p><p>The presence of this Ti plasmid is essential for the bacteria to cause crown gall disease in plants.<sup id="cite_ref-Gordon2014_1-1" class="reference"><a href="#cite_note-Gordon2014-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> This is facilitated via certain crucial regions in the Ti plasmid, including the <i>vir</i> region, which encodes for virulence genes, and the <a href="/wiki/Transfer_DNA" title="Transfer DNA">transfer DNA</a> (T-DNA) region, which is a section of the Ti plasmid that is <a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">transferred</a> via <a href="/wiki/Bacterial_conjugation" title="Bacterial conjugation">conjugation</a> into host plant cells after an injury site is sensed by the bacteria. These regions have features that allow the delivery of T-DNA into host plant cells, and can modify the host plant cell to cause the synthesis of molecules like <a href="/wiki/Plant_hormone" title="Plant hormone">plant hormones</a> (e.g. <a href="/wiki/Auxin" title="Auxin">auxins</a>, <a href="/wiki/Cytokinin" title="Cytokinin">cytokinins</a>) and opines and the formation of crown gall tumours.<sup id="cite_ref-Gordon2014_1-2" class="reference"><a href="#cite_note-Gordon2014-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>Because the T-DNA region of the Ti plasmid can be transferred from bacteria to plant cells, it represented an exciting avenue for the transfer of DNA between <a href="/wiki/Kingdom_(biology)" title="Kingdom (biology)">kingdoms</a> and spurred large amounts of research on the Ti plasmid and its possible uses in bioengineering. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Nomenclature_and_classification">Nomenclature and classification</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ti_plasmid&action=edit&section=1" title="Edit section: Nomenclature and classification"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The Ti plasmid is a member of the <a href="/w/index.php?title=RepABC&action=edit&redlink=1" class="new" title="RepABC (page does not exist)">RepABC</a> plasmid family found in Alphaproteobacteria.<sup id="cite_ref-Pinto2012_3-0" class="reference"><a href="#cite_note-Pinto2012-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> These plasmids are often relatively large in size, ranging from 100kbp to 2Mbp. They are also often termed <a href="/wiki/Replicon_(genetics)" title="Replicon (genetics)">replicons</a>, as their replication begins at a single site. Members of this family have a characteristic <i>repABC</i> gene cassette.<sup id="cite_ref-Cevallos2008_4-0" class="reference"><a href="#cite_note-Cevallos2008-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Another notable member of this family is the root inducing (Ri) plasmid carried by <i>A. rhizogenes</i>, which causes another plant disease known as hairy root disease.<sup id="cite_ref-Gordon2014_1-3" class="reference"><a href="#cite_note-Gordon2014-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>A key feature of Ti plasmids is their ability to drive the production of opines, which are derivatives of various <a href="/wiki/Amino_acid" title="Amino acid">amino acids</a> or <a href="/wiki/Sugar_phosphates" title="Sugar phosphates">sugar phosphates</a>, in host plant cells. These opines can then be used as a nutrient for the infecting bacteria, which <a href="/wiki/Catabolism" title="Catabolism">catabolizes</a> the respective opines using genes encoded in the Ti plasmid. </p><p>Accordingly, Ti plasmids have been classified based on the type of opine they catabolize, namely: <a href="/wiki/Nopaline" title="Nopaline">nopaline-</a>, <a href="/wiki/Octopine" title="Octopine">octopine-</a> or mannityl-types, which are amino acid derivatives, or agrocinopine-type, which are sugar phosphate derivatives.<sup id="cite_ref-Gordon2014_1-4" class="reference"><a href="#cite_note-Gordon2014-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Historical_discovery">Historical discovery</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ti_plasmid&action=edit&section=2" title="Edit section: Historical discovery"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The identification of <i>A. tumefaciens</i> as the cause of gall tumours in plants paved the way for insights into the molecular basis of crown gall disease.<sup id="cite_ref-Kado_5-0" class="reference"><a href="#cite_note-Kado-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>The first indication of a genetic effect on host plant cells came in 1942-1943, where plant cells of secondary tumours were found to lack any bacterial cells within. However, these tumour cells did possess the ability to produce opines metabolized by the infecting bacterial strain.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Crucially, the production of the respective opines occurred regardless of the plant species and occasionally only within crown gall tissues, indicating that the bacteria had transferred some genetic material to the host plant cells in order to allow opine synthesis.<sup id="cite_ref-Kado_5-1" class="reference"><a href="#cite_note-Kado-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>However, how and to what extend did DNA transfer occur remained an open question. Adding <i>A. tumefaciens</i> DNA alone did not cause tumors in plants,<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> while very little <i>A. tumefaciens</i> DNA was found to be integrated into the host plant cell genome.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> The addition of <a href="/wiki/Deoxyribonuclease" title="Deoxyribonuclease">deoxyribonucleases</a> (DNases) to degrade DNA also failed to prevent the formation and growth of the plant tumors.<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> These suggested that little, if any, of the <i>A. tumefaciens</i> DNA is transferred to the host plant cell to cause disease and, if DNA is indeed transferred from the bacteria to the plant, it must occur in a protected manner. </p><p>Subsequently, <a href="/wiki/Oncogenic" class="mw-redirect" title="Oncogenic">oncogenic</a> bacterial strains were found to be able to convert non-pathogenic bacteria into pathogens via the process of conjugation, where the genes responsible for virulence were transferred to the non-pathogenic cells.<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> The role of a plasmid in this pathogenic ability was further supported when large plasmids were found only in pathogenic bacteria but not avirulent bacteria.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Eventually, the detection of parts of bacterial plasmids in host plant cells was established, confirming that this was the genetic material responsible for the genetic effect of infection.<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> </p><p>With the identification of the Ti plasmid, many studies were carried out to determine the characteristics of the Ti plasmid and how the genetic material is transferred from the <i>Agrobacterium</i> to the plant host. Some notable early milestones in the studies of Ti plasmids include the mapping of a Ti plasmid in 1978 and the studying of sequence similarity between different Ti plasmids in 1981.<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><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> </p><p>Between 1980–2000, the characterization of the T-DNA region and the 'vir' region was also pursued. Studies into the T-DNA region determined their process of transfer and identified genes allowing the synthesis of plant hormones and opines.<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> Separately, early work aimed to determine the functions of the genes encoded in the 'vir' region - these were broadly categorized into those that allowed bacterial-host interactions and those that enabled T-DNA delivery.<sup id="cite_ref-Hooykaas1994_2-1" class="reference"><a href="#cite_note-Hooykaas1994-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Replication,_partitioning_and_maintenance"><span id="Replication.2C_partitioning_and_maintenance"></span>Replication, partitioning and maintenance</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ti_plasmid&action=edit&section=3" title="Edit section: Replication, partitioning and maintenance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-center" typeof="mw:File/Thumb"><a href="/wiki/File:RepABC_gene_cassette_of_Ti_plasmids.png" class="mw-file-description"><img alt="A schematic of the repABC gene cassette, along with the activity of their gene products" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d8/RepABC_gene_cassette_of_Ti_plasmids.png/330px-RepABC_gene_cassette_of_Ti_plasmids.png" decoding="async" width="330" height="233" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d8/RepABC_gene_cassette_of_Ti_plasmids.png/495px-RepABC_gene_cassette_of_Ti_plasmids.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d8/RepABC_gene_cassette_of_Ti_plasmids.png/660px-RepABC_gene_cassette_of_Ti_plasmids.png 2x" data-file-width="1123" data-file-height="794" /></a><figcaption>The <i>repABC gene</i> cassette of Ti plasmids in Agrobacteria, with a schematic of their gene product and activities</figcaption></figure> <p>The replication, partitioning and maintenance of the Ti plasmid depends on the <i>repABC</i> gene cassette, which is mainly made up of three genes: <i>repA</i>, <i>repB</i> and <i>repC</i>. <i>repA</i> and <i>repB</i> each encode for proteins involved in plasmid partitioning, while <i>repC</i> encodes a replication initiator.<sup id="cite_ref-Gordon2014_1-5" class="reference"><a href="#cite_note-Gordon2014-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> These genes are expressed from 4 different promoters located upstream of <i>repA</i>. <i>repE</i> encodes for a small <a href="/wiki/Antisense_RNA" title="Antisense RNA">antisense RNA</a> and is located between <i>repB</i> and <i>repC</i>.<sup id="cite_ref-Cevallos2008_4-1" class="reference"><a href="#cite_note-Cevallos2008-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Additionally, there is a partitioning site (<i>parS</i>) and an <a href="/wiki/Origin_of_replication" title="Origin of replication">origin of replication</a> (<i>oriV</i>) present within the <i>repABC</i> cassette.<sup id="cite_ref-Gordon2014_1-6" class="reference"><a href="#cite_note-Gordon2014-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Replication_of_the_Ti_plasmid">Replication of the Ti plasmid</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ti_plasmid&action=edit&section=4" title="Edit section: Replication of the Ti plasmid"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The replication of the Ti plasmid is driven by the RepC initiator protein (<span class="plainlinks"><a rel="nofollow" class="external text" href="https://www.uniprot.org/uniprot/P05684">P05684</a></span>), which possesses two <a href="/wiki/Protein_domain" title="Protein domain">protein domains</a>: an <a href="/wiki/N-terminal" class="mw-redirect" title="N-terminal">N-terminal</a> domain (NTD) that binds to DNA and a <a href="/wiki/C-terminal" class="mw-redirect" title="C-terminal">C-terminal</a> domain (CTD). Mutational analyses have shown that without a functional RepC protein, the Ti plasmid is unable to replicate.<sup id="cite_ref-Cevallos2008_4-2" class="reference"><a href="#cite_note-Cevallos2008-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Meanwhile, the <i>oriV</i> sequence is around 150 nucleotides in length and is found within the <i>repC</i> gene.<sup id="cite_ref-Pinto2012_3-1" class="reference"><a href="#cite_note-Pinto2012-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Laboratory experiments have shown that the RepC protein binds to this region, suggesting its role as the origin of replication.<sup id="cite_ref-Pinto2011_16-0" class="reference"><a href="#cite_note-Pinto2011-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> Therefore, while the complete process behind the replication of the Ti plasmid has not been fully described, the initial step of replication would likely depend on the expression of RepC and its binding to <i>oriV</i>. Of note, the RepC protein only acts in <i>cis</i>, where it only drives the replication of the plasmid it is encoded in and not any other plasmid also present in the bacterial cell.<sup id="cite_ref-Pinto2011_16-1" class="reference"><a href="#cite_note-Pinto2011-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Partitioning_of_the_Ti_plasmid">Partitioning of the Ti plasmid</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ti_plasmid&action=edit&section=5" title="Edit section: Partitioning of the Ti plasmid"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <table class="wikitable"> <caption style="text-align:left;">Components involved in the RepA/RepB partitioning system of Ti plasmids<sup id="cite_ref-Gordon2014_1-7" class="reference"><a href="#cite_note-Gordon2014-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </caption> <tbody><tr> <th>Component </th> <th>Function </th></tr> <tr> <td>RepA (ParA), <span class="plainlinks"><a rel="nofollow" class="external text" href="https://www.uniprot.org/uniprot/P05682">P05682</a></span> </td> <td>A weak <a href="/wiki/ATPase" title="ATPase">ATPase</a> that negatively <a href="/wiki/Autoregulation" title="Autoregulation">autoregulates</a> the expression of the <i>repABC</i> cassette and can form filaments to aid in the partitioning of the plasmid during cell division </td></tr> <tr> <td>RepB (ParB), <span class="plainlinks"><a rel="nofollow" class="external text" href="https://www.uniprot.org/uniprot/P05683">P05683</a></span> </td> <td>A DNA binding protein that serves as an adaptor between RepA and the <i>parS</i> site </td></tr> <tr> <td><i>parS</i> </td> <td>The palindromic binding site for the ParB protein; consensus <samp class="dna-sequence" style="word-break: break-all;">GTTNNCNGCNGNNAAC</samp> </td></tr></tbody></table> <p>The <a href="/wiki/Plasmid_partition_system" title="Plasmid partition system">partitioning system</a> of the Ti plasmid is similar to the ParA/ParB system used in other plasmids and bacterial chromosomes and is thought to act in the same way.<sup id="cite_ref-Bignell2001_17-0" class="reference"><a href="#cite_note-Bignell2001-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Mutations in either of the proteins RepA or RepB have resulted in a decrease in plasmid stability, indicating their role and importance in plasmid partitioning.<sup id="cite_ref-Cevallos2008_4-3" class="reference"><a href="#cite_note-Cevallos2008-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> The ability of RepA to form filaments allows it to create a physical bridge along which DNA can be pulled to opposite poles of a dividing cell. Meanwhile, the RepB protein can bind specifically to the <i>parS</i> sequence, forming a complex with DNA that can be recognized by RepA.<sup id="cite_ref-Gordon2014_1-8" class="reference"><a href="#cite_note-Gordon2014-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Cevallos2008_4-4" class="reference"><a href="#cite_note-Cevallos2008-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> This system is particularly important for the proper partitioning of the Ti plasmid, as the plasmid is only present in few copy numbers in the bacterial cell. </p> <div class="mw-heading mw-heading3"><h3 id="Maintenance_of_the_Ti_plasmid">Maintenance of the Ti plasmid</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ti_plasmid&action=edit&section=6" title="Edit section: Maintenance of the Ti plasmid"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The Ti plasmid is maintained at low copy numbers within a bacterial cell. This is partly achieved by influencing the expression of the replication initiator RepC.<sup id="cite_ref-Gordon2014_1-9" class="reference"><a href="#cite_note-Gordon2014-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> When bound to <a href="/wiki/Adenosine_diphosphate" title="Adenosine diphosphate">ADP</a>, RepA is activated to work with RepB, acting as a negative <a href="/wiki/Regulation_of_gene_expression" title="Regulation of gene expression">regulator</a> of the <i>repABC</i> cassette.<sup id="cite_ref-Pinto2012_3-2" class="reference"><a href="#cite_note-Pinto2012-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> The levels of RepC is therefore kept low within a cell, preventing too many rounds of replication from occurring during each cell division cycle. Furthermore, there is a small RNA known as RepE encoded between <i>repB</i> and <i>repC</i> that lowers the expression of <i>repC</i>.<sup id="cite_ref-Chai2005_18-0" class="reference"><a href="#cite_note-Chai2005-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> RepE is <a href="/wiki/Complementarity_(molecular_biology)" title="Complementarity (molecular biology)">complementary</a> to RepC and will bind with the <i>repC</i> <a href="/wiki/Messenger_RNA" title="Messenger RNA">mRNA</a> to form a double-stranded molecule. This can then block the <a href="/wiki/Translation_(biology)" title="Translation (biology)">translational</a> production of the RepC protein.<sup id="cite_ref-Chai2005_18-1" class="reference"><a href="#cite_note-Chai2005-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p><p>Separately, the expression of the <i>repABC</i> cassette and hence the copy number of the Ti plasmid is also influenced via a <a href="/wiki/Quorum_sensing" title="Quorum sensing">quorum sensing</a> system in <i>Agrobacterium</i>.<sup id="cite_ref-Cevallos2008_4-5" class="reference"><a href="#cite_note-Cevallos2008-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Quorum sensing systems respond to bacterial population densities by sensing a molecule, known as an autoinducer, that is produced by the bacterial cells at low levels and would build up to a threshold level when there is a high density of bacteria present.<sup id="cite_ref-Chai2005_18-2" class="reference"><a href="#cite_note-Chai2005-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> In this case, the autoinducer is the N-3-oxooctanoyl-L-homoserine lactone (3-O-C<sub>8</sub>-AHL) molecule, which is sensed by a regulator known as TraR.<sup id="cite_ref-Cevallos2008_4-6" class="reference"><a href="#cite_note-Cevallos2008-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> When activated, TraR will bind to regions known as <i>tra</i> boxes in the <i>repABC</i> gene cassette's promoter regions to drive expression. Therefore, a high level of population density increases the number of plasmids present within each bacterial cell, likely to support pathogenesis in the plant host.<sup id="cite_ref-Cevallos2008_4-7" class="reference"><a href="#cite_note-Cevallos2008-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Features">Features</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ti_plasmid&action=edit&section=7" title="Edit section: Features"><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-Missing_information plainlinks metadata ambox ambox-content" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><a href="/wiki/File:Wiki_letter_w.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/en/thumb/6/6c/Wiki_letter_w.svg/44px-Wiki_letter_w.svg.png" decoding="async" width="44" height="44" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/6/6c/Wiki_letter_w.svg/66px-Wiki_letter_w.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/6/6c/Wiki_letter_w.svg/88px-Wiki_letter_w.svg.png 2x" data-file-width="44" data-file-height="44" /></a></span></div></td><td class="mbox-text"><div class="mbox-text-span">This section <b>is missing information</b> about UniProt links for easier exploration.<span class="hide-when-compact"> Please expand the section to include this information. Further details may exist on the <a href="/wiki/Talk:Ti_plasmid" title="Talk:Ti plasmid">talk page</a>.</span> <span class="date-container"><i>(<span class="date">December 2020</span>)</i></span></div></td></tr></tbody></table> <div class="mw-heading mw-heading3"><h3 id="Virulence_operon">Virulence operon</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ti_plasmid&action=edit&section=8" title="Edit section: Virulence operon"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-center" typeof="mw:File/Thumb"><a href="/wiki/File:Vir_region_of_Ti_plasmids.png" class="mw-file-description"><img alt="A diagram showing the composition of the vir region of Ti plasmids" src="//upload.wikimedia.org/wikipedia/commons/thumb/4/49/Vir_region_of_Ti_plasmids.png/440px-Vir_region_of_Ti_plasmids.png" decoding="async" width="440" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/49/Vir_region_of_Ti_plasmids.png/660px-Vir_region_of_Ti_plasmids.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/49/Vir_region_of_Ti_plasmids.png/880px-Vir_region_of_Ti_plasmids.png 2x" data-file-width="1117" data-file-height="373" /></a><figcaption>The composition of the <i>vir</i> region of octopine-type Ti plasmids</figcaption></figure> <p>The expression of the <i>vir</i> region is usually repressed under normal conditions, and only becomes activated when the bacteria senses plant-derived signals from wound sites. This activation is necessary for the production of Vir proteins and the transfer of DNA and proteins into host plant cells.<sup id="cite_ref-Gordon2014_1-10" class="reference"><a href="#cite_note-Gordon2014-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>VirA and VirG form a <a href="/wiki/Two-component_regulatory_system" title="Two-component regulatory system">two-component regulatory system</a> within <i>Agrobacterium</i>.<sup id="cite_ref-Winans1991_19-0" class="reference"><a href="#cite_note-Winans1991-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> This is a type of sensing and signalling system found commonly in bacteria; in this case, they act to sense plant-derived signals to drive the expression of the <i>vir</i> region. During the sensing, VirA, a histidine sensor kinase, will become <a href="/wiki/Phosphorylation" title="Phosphorylation">phosphorylated</a> before passing on this phosphate group to the response regulator VirG.<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> The activated response regulator VirG can then bind to a region of DNA known as the <i>vir</i> box, located upstream of each <i>vir</i> promoter, to activate the expression of the <i>vir</i> region.<sup id="cite_ref-Gordon2014_1-11" class="reference"><a href="#cite_note-Gordon2014-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Winans1991_19-1" class="reference"><a href="#cite_note-Winans1991-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> One possible downstream functions of the sensing mediated by VirA and VirG is the directional movement, or <a href="/wiki/Chemotaxis" title="Chemotaxis">chemotaxis</a>, of the bacteria towards plant-derived signals; this allows the <i>Agrobacterium</i> to move towards the wound site in plants.<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> Furthermore, with the induction of the <i>vir</i> region, the transfer of T-DNA can be mediated by the Vir proteins.<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><p>The <i>virB</i> <a href="/wiki/Operon" title="Operon">operon</a> is the largest operon in the <i>vir</i> region, encoding for 11 VirB proteins involved in the transfer process of T-DNA and bacterial proteins into host plant cells (see transfer apparatus below).<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><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><p>The <i>virC</i> operon encodes for two proteins: VirC1 and VirC2. These proteins influence the pathogenesis of the <i>Agrobacterium</i> towards different plant hosts, and mutations can reduce but not remove the virulence of the bacteria.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> Both the <i>virC</i> and <i>virD</i> operons can be repressed by a chromosomally encoded protein known as Ros.<sup id="cite_ref-Cooley1991_26-0" class="reference"><a href="#cite_note-Cooley1991-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-DSouza1993_27-0" class="reference"><a href="#cite_note-DSouza1993-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> Ros binds to a region of DNA that overlaps with the binding site of the VirG regulator, and therefore competes with VirG to control their expression levels.<sup id="cite_ref-Cooley1991_26-1" class="reference"><a href="#cite_note-Cooley1991-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-DSouza1993_27-1" class="reference"><a href="#cite_note-DSouza1993-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> Functionally, VirC1 and VirC2 promote the assembly of a <a href="/wiki/Relaxosome" title="Relaxosome">relaxosome</a> complex during the conjugative transfer of T-DNA from the bacteria to the host plant cell.<sup id="cite_ref-Atmakuri2007_28-0" class="reference"><a href="#cite_note-Atmakuri2007-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> This is an energy-dependent process mediated via their NTPase activity, and occurs as they bind to a region of DNA known as <i>overdrive</i>.<sup id="cite_ref-Atmakuri2007_28-1" class="reference"><a href="#cite_note-Atmakuri2007-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> As a result, they act to increase the amount of T-DNA strands produced. Following the production of the DNA strand to be transferred (transfer strand, T-strand), the VirC proteins can also help to direct the transfer strand to the transfer apparatus.<sup id="cite_ref-Atmakuri2007_28-2" class="reference"><a href="#cite_note-Atmakuri2007-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p><p>The <i>virD</i> operon encodes for 4 proteins: VirD1-D4.<sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> VirD1 and VirD2 are involved in the processing of T-DNA during conjugation to produce the T-strand; this is the single-stranded DNA molecule that is transported to the host plant cell (see transfer apparatus below).<sup id="cite_ref-Ghai1989_30-0" class="reference"><a href="#cite_note-Ghai1989-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> During the processing, VirD1 will act as a <a href="/wiki/Topoisomerase" title="Topoisomerase">topoisomerase</a> to unwind the DNA strands.<sup id="cite_ref-Ghai1989_30-1" class="reference"><a href="#cite_note-Ghai1989-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> VirD2, a <a href="/wiki/Relaxase" title="Relaxase">relaxase</a>, will then nick one of the DNA strands and remain bound to the DNA as it is transferred to the recipient cell.<sup id="cite_ref-Zechner2012_31-0" class="reference"><a href="#cite_note-Zechner2012-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup><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> Within the recipient cell, VirD2 will also work together with VirE2 to direct the transferred DNA to the recipient cell's nucleus. There are suggestions that VirD2 may be phosphorylated and dephosphorylated by different proteins, affecting its ability to deliver DNA.<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> Conversely, little is known about VirD3, and mutational analyses have not provided any support for its role in the virulence of <i>Agrobacterium</i>.<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> Finally, VirD4 is a crucial part of the conjugation process, serving as a coupling factor that recognizes and transfers the T-strand to the transport channel.<sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p><p>The <i>virE</i> operon encodes for 2 proteins: VirE1 and VirE2.<sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> VirE2 is an effector protein translocated together with the T-strand into host plant cells. There, it binds to the T-strand to direct its delivery to the <a href="/wiki/Cell_nucleus" title="Cell nucleus">nucleus</a> of the host plant cell.<sup id="cite_ref-Galvin2012_37-0" class="reference"><a href="#cite_note-Galvin2012-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> Part of this activity involves the presence of <a href="/wiki/Nuclear_localization_sequence" title="Nuclear localization sequence">nuclear localization sequences</a> within the protein, which marks the protein and the associated DNA for entry into the nucleus. It also protects the T-strand from <a href="/wiki/Nuclease" title="Nuclease">nuclease</a> attack.<sup id="cite_ref-Schrammeijer_39-0" class="reference"><a href="#cite_note-Schrammeijer-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> There is some speculation regarding the role of VirE2 as a protein channel, allowing DNA to move through the plant <a href="/wiki/Cytoplasmic_membrane" class="mw-redirect" title="Cytoplasmic membrane">cytoplasmic membrane</a>.<sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> On the other hand, VirE1 may be involved in promoting the transfer of the VirE2 protein into the host plant cell.<sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> It binds to the ssDNA-binding domain of VirE2, therefore preventing the VirE2 protein from prematurely binding to the T-strand within the bacterial cell.<sup id="cite_ref-42" class="reference"><a href="#cite_note-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> </p><p><i>virF</i> is a host specificity factor found in some but not all types of Ti plasmids; for example, octopine-type Ti plasmids possess <i>virF</i> but nopaline-types do not.<sup id="cite_ref-Jarchow1991_43-0" class="reference"><a href="#cite_note-Jarchow1991-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Melchers1990_44-0" class="reference"><a href="#cite_note-Melchers1990-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> The ability of <i>A. tumefaciens</i> to induce crown gall tumours in certain plant species but not others has been attributed to the presence or absence of this <i>virF</i> gene.<sup id="cite_ref-Jarchow1991_43-1" class="reference"><a href="#cite_note-Jarchow1991-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Melchers1990_44-1" class="reference"><a href="#cite_note-Melchers1990-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> </p><p>The <i>virH</i> operon encodes for 2 proteins: VirH1 and VirH2.<sup id="cite_ref-Kalogeraki1999_45-0" class="reference"><a href="#cite_note-Kalogeraki1999-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> A <a href="/wiki/Bioinformatics" title="Bioinformatics">bioinformatics</a> study of the amino acid sequences of the VirH protein showed similarities between them and a superfamily of proteins known as <a href="/wiki/Cytochrome_P450" title="Cytochrome P450">cytochrome P450</a> enzymes.<sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> VirH2 was then discovered to metabolize certain phenolic compounds detected by VirA.<sup id="cite_ref-Kalogeraki1999_45-1" class="reference"><a href="#cite_note-Kalogeraki1999-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Transfer_DNA_(T-DNA)"><span id="Transfer_DNA_.28T-DNA.29"></span>Transfer DNA (T-DNA)</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ti_plasmid&action=edit&section=9" title="Edit section: Transfer DNA (T-DNA)"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The T-DNA of <i>Agrobacterium</i> is approximately 15-20 kbp in length and will become integrated into the host plant genome upon its transfer via a process known as <a href="/wiki/Genetic_recombination" title="Genetic recombination">recombination</a>. This process utilizes preexisting gaps in the host plant cell's genome to allow the T-DNA to pair with short sequences in the genome, priming the process of <a href="/wiki/Ligation_(molecular_biology)" title="Ligation (molecular biology)">DNA ligation</a>, where the T-DNA is permanently joint to the plant genome.<sup id="cite_ref-Galvin2012_37-1" class="reference"><a href="#cite_note-Galvin2012-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> The T-DNA region is flanked at both ends by 24bp sequences. </p><p>Within the host plant cell's genome, the T-DNA of <i>Agrobacterium</i> is expressed to produce two main groups of proteins.<sup id="cite_ref-Gordon2014_1-12" class="reference"><a href="#cite_note-Gordon2014-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> One group is responsible for the production of plant growth hormones. As these hormones are produced, there will be an increase in the rate of cell division and therefore the formation of crown gall tumors.<sup id="cite_ref-Zhu2000_47-0" class="reference"><a href="#cite_note-Zhu2000-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> The second group of proteins are responsible for driving the synthesis of opines in the host plant cells. The specific opines produced depends on the type of the Ti plasmid but not on the plant host. These opines cannot be utilized by the plant host, and will instead be exported out of the plant cell where it can be taken up by the <i>Agrobacterium</i> cells. The bacteria possess genes in other regions of the Ti plasmid that allows the catabolism of opines.<sup id="cite_ref-Gordon2014_1-13" class="reference"><a href="#cite_note-Gordon2014-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Transfer_apparatus">Transfer apparatus</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ti_plasmid&action=edit&section=10" title="Edit section: Transfer apparatus"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Transfer apparatuses encoded within the Ti plasmid have to achieve two objectives: allow the conjugative transfer of the Ti plasmid between bacteria and allow the delivery of the T-DNA and certain effector proteins into host plant cells. These are achieved by the Tra/Trb system and the VirB/VirD4 system respectively, which are members of the <a href="/wiki/Bacterial_secretion_system" title="Bacterial secretion system">type IV secretion system</a> (T4SS).<sup id="cite_ref-Zhu2000_47-1" class="reference"><a href="#cite_note-Zhu2000-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> </p><p>For the Ti plasmid and T-DNA to be transferred via conjugation, they must first be processed by different proteins, such as the relaxase enzyme (TraA/VirD2) and the DNA transfer and replication (Dtr) proteins. Together, these proteins will recognize and bind to a region known as the <a href="/wiki/Origin_of_transfer" title="Origin of transfer">origin of transfer</a> (<i>oriT</i>) in the Ti plasmid to form the relaxosome complex. For the T-DNA, a nick will be created at the T-DNA's border sequence, and the nicked T-strand will be transported to the cell membrane, where the rest of the transfer machinery is present.<sup id="cite_ref-Zechner2012_31-1" class="reference"><a href="#cite_note-Zechner2012-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> </p><p>Within the VirB/VirD4 system, the VirD2 relaxase is aided by the accessory factors VirD1, VirC1 and VirC2 while it processes the DNA substrate.<sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> Furthermore, the VirD2 relaxase and the VirC proteins will contribute to the delivery of the DNA strand to the VirD4 receptor at the cell membrane.<sup id="cite_ref-Atmakuri2007_28-3" class="reference"><a href="#cite_note-Atmakuri2007-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> This receptor is an essential component of T4SSs, and is thought to energize and mediate the transfer of the DNA into the translocation channel between two cells.<sup id="cite_ref-49" class="reference"><a href="#cite_note-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> The table below summarizes the proteins encoded in the <i>virB</i> operon that makes up the translocation channel of the VirB/VirD4 system.<sup id="cite_ref-Gordon2014_1-14" class="reference"><a href="#cite_note-Gordon2014-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <table class="wikitable"> <tbody><tr> <th>Protein(s) </th> <th>Function </th></tr> <tr> <td>VirB4, VirB11 </td> <td>ATPases that provide the energy for DNA transfer<sup id="cite_ref-christie2005_50-0" class="reference"><a href="#cite_note-christie2005-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-51" class="reference"><a href="#cite_note-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>VirB3, VirB6, VirB8 </td> <td>Subunits of a putative inner membrane <a href="/wiki/Translocase" title="Translocase">translocase</a><sup id="cite_ref-christie2005_50-1" class="reference"><a href="#cite_note-christie2005-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-52" class="reference"><a href="#cite_note-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-53" class="reference"><a href="#cite_note-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>VirB7, VirB9, VirB10 </td> <td>Forms a core complex that stabilizes the channel subunits<sup id="cite_ref-christie2005_50-2" class="reference"><a href="#cite_note-christie2005-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-54" class="reference"><a href="#cite_note-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>VirB2 </td> <td>The major <a href="/wiki/Pilin" title="Pilin">pilin</a> subunit of the conjugative <a href="/wiki/Pilus" title="Pilus">pilus</a><sup id="cite_ref-christie2005_50-3" class="reference"><a href="#cite_note-christie2005-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>VirB1, VirB5 </td> <td>Minor components of the conjugative pilus<sup id="cite_ref-55" class="reference"><a href="#cite_note-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-56" class="reference"><a href="#cite_note-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> </td></tr></tbody></table> <div class="mw-heading mw-heading2"><h2 id="Uses_in_bioengineering">Uses in bioengineering</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ti_plasmid&action=edit&section=11" title="Edit section: Uses in bioengineering"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Agrobacterium_tumefaciens" title="Agrobacterium tumefaciens">Agrobacterium tumefaciens</a></div> <p>The ability of <i>Agrobacterium</i> to deliver DNA into plant cells opened new doors for plant <a href="/wiki/Genome_engineering" class="mw-redirect" title="Genome engineering">genome engineering</a>, allowing the production of <a href="/wiki/Genetically_modified_plant" title="Genetically modified plant">genetically modified plants</a> (transgenic plants).<sup id="cite_ref-57" class="reference"><a href="#cite_note-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> Proteins involved in mediating the transfer of T-DNA will first recognize the border sequences of the T-DNA region. Therefore, it is possible for scientists to use T-DNA border sequences to flank any desired sequence of interest - such a product can then be inserted into a plasmid and introduced into <i>Agrobacterium</i> cells.<sup id="cite_ref-58" class="reference"><a href="#cite_note-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> There, the border sequences will be recognized by the transfer apparatus of <i>A. tumefaciens</i> and delivered in a standard manner into the target plant cell.<sup id="cite_ref-Gordon2014_1-15" class="reference"><a href="#cite_note-Gordon2014-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Moreover, by leaving behind only the border sequences of the T-DNA, the resulting product will edit the plant genome without causing any tumours in plants.<sup id="cite_ref-59" class="reference"><a href="#cite_note-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> This method has been used to modify several crop plants, including rice,<sup id="cite_ref-60" class="reference"><a href="#cite_note-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> barley<sup id="cite_ref-61" class="reference"><a href="#cite_note-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> and wheat.<sup id="cite_ref-62" class="reference"><a href="#cite_note-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> Further work have since extended the targets of <i>A. tumefaciens</i> to include fungi and human cell lines.<sup id="cite_ref-63" class="reference"><a href="#cite_note-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-64" class="reference"><a href="#cite_note-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Similar_plasmids">Similar plasmids</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ti_plasmid&action=edit&section=12" title="Edit section: Similar plasmids"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Root-inducing_(Ri)_plasmid"><span id="Root-inducing_.28Ri.29_plasmid"></span>Root-inducing (Ri) plasmid</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ti_plasmid&action=edit&section=13" title="Edit section: Root-inducing (Ri) plasmid"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Symbiotic_(sym)_plasmids_of_Rhizobia"><span id="Symbiotic_.28sym.29_plasmids_of_Rhizobia"></span>Symbiotic (sym) plasmids of Rhizobia</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ti_plasmid&action=edit&section=14" title="Edit section: Symbiotic (sym) plasmids of Rhizobia"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></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=Ti_plasmid&action=edit&section=15" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Mary-Dell_Chilton" title="Mary-Dell Chilton">Mary-Dell Chilton</a></li> <li><a href="/wiki/Jeff_Schell" class="mw-redirect" title="Jeff Schell">Jeff Schell</a></li> <li><a href="/wiki/Marc_Van_Montagu" title="Marc Van Montagu">Marc Van Montagu</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=Ti_plasmid&action=edit&section=16" 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-Gordon2014-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Gordon2014_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Gordon2014_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Gordon2014_1-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Gordon2014_1-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Gordon2014_1-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-Gordon2014_1-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-Gordon2014_1-6"><sup><i><b>g</b></i></sup></a> <a href="#cite_ref-Gordon2014_1-7"><sup><i><b>h</b></i></sup></a> <a href="#cite_ref-Gordon2014_1-8"><sup><i><b>i</b></i></sup></a> <a href="#cite_ref-Gordon2014_1-9"><sup><i><b>j</b></i></sup></a> <a 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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="CITEREFGordonChristie2014" class="citation journal cs1">Gordon JE, Christie PJ (December 2014). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4292801">"The Agrobacterium Ti Plasmids"</a>. <i>Microbiology Spectrum</i>. <b>2</b> (6). <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1128%2Fmicrobiolspec.PLAS-0010-2013">10.1128/microbiolspec.PLAS-0010-2013</a></span>. <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/PMC4292801">4292801</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/25593788">25593788</a>.</cite><span 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(November 1997). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC158560">"Genetic Transformation of Wheat Mediated by Agrobacterium tumefaciens"</a>. <i>Plant Physiology</i>. <b>115</b> (3): 971–980. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1104%2Fpp.115.3.971">10.1104/pp.115.3.971</a></span>. <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/PMC158560">158560</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/12223854">12223854</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Plant+Physiology&rft.atitle=Genetic+Transformation+of+Wheat+Mediated+by+Agrobacterium+tumefaciens&rft.volume=115&rft.issue=3&rft.pages=971-980&rft.date=1997-11&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC158560%23id-name%3DPMC&rft_id=info%3Apmid%2F12223854&rft_id=info%3Adoi%2F10.1104%2Fpp.115.3.971&rft.aulast=Cheng&rft.aufirst=M&rft.au=Fry%2C+JE&rft.au=Pang%2C+S&rft.au=Zhou%2C+H&rft.au=Hironaka%2C+CM&rft.au=Duncan%2C+DR&rft.au=Conner%2C+TW&rft.au=Wan%2C+Y&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC158560&rfr_id=info%3Asid%2Fen.wikipedia.org%3ATi+plasmid" class="Z3988"></span></span> </li> <li id="cite_note-63"><span class="mw-cite-backlink"><b><a href="#cite_ref-63">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFTzfiraCitovsky2007" class="citation book cs1">Tzfira T, <a href="/wiki/Vitaly_Citovsky" title="Vitaly Citovsky">Citovsky V</a> (2007). <i><span></span></i>Agrobacterium<i>: from biology to biotechnology</i>. Springer Science & Business Media.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Agrobacterium%3A+from+biology+to+biotechnology&rft.pub=Springer+Science+%26+Business+Media&rft.date=2007&rft.aulast=Tzfira&rft.aufirst=T&rft.au=Citovsky%2C+V&rfr_id=info%3Asid%2Fen.wikipedia.org%3ATi+plasmid" class="Z3988"></span></span> </li> <li id="cite_note-64"><span class="mw-cite-backlink"><b><a href="#cite_ref-64">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKunikTzfiraKapulnikGafni2001" class="citation journal cs1">Kunik T, Tzfira T, Kapulnik Y, Gafni Y, Dingwall C, Citovsky V (February 2001). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC29349">"Genetic transformation of HeLa cells by Agrobacterium"</a>. <i>Proceedings of the National Academy of Sciences of the United States of America</i>. <b>98</b> (4): 1871–6. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2001PNAS...98.1871K">2001PNAS...98.1871K</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.98.4.1871">10.1073/pnas.98.4.1871</a></span>. <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/PMC29349">29349</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/11172043">11172043</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+National+Academy+of+Sciences+of+the+United+States+of+America&rft.atitle=Genetic+transformation+of+HeLa+cells+by+Agrobacterium&rft.volume=98&rft.issue=4&rft.pages=1871-6&rft.date=2001-02&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC29349%23id-name%3DPMC&rft_id=info%3Apmid%2F11172043&rft_id=info%3Adoi%2F10.1073%2Fpnas.98.4.1871&rft_id=info%3Abibcode%2F2001PNAS...98.1871K&rft.aulast=Kunik&rft.aufirst=T&rft.au=Tzfira%2C+T&rft.au=Kapulnik%2C+Y&rft.au=Gafni%2C+Y&rft.au=Dingwall%2C+C&rft.au=Citovsky%2C+V&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC29349&rfr_id=info%3Asid%2Fen.wikipedia.org%3ATi+plasmid" class="Z3988"></span></span> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a 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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 class="mw-selflink selflink">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 href="/wiki/Prophage" title="Prophage">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 href="/wiki/Artificial_cell" title="Artificial cell">Artificial cell</a></li></ul></li> <li><a href="/wiki/Non-cellular_life" title="Non-cellular life">Non-cellular life</a></li> <li><a href="/wiki/Synthetic_virology" title="Synthetic virology">Synthetic virus</a> <ul><li><a href="/wiki/Viral_vector" title="Viral vector">Viral vector</a></li> <li><a href="/wiki/Helper_dependent_virus" title="Helper dependent virus">Helper dependent virus</a></li></ul></li> <li>?<a href="/wiki/Nanobacterium" title="Nanobacterium">Nanobacterium</a></li> <li>?<a href="/wiki/Nanobe" title="Nanobe">Nanobe</a></li> <li><a href="/wiki/Cancer_cell" title="Cancer cell">Cancer cell</a> <ul><li><a href="/wiki/HeLa" title="HeLa">HeLa</a></li> <li><a href="/wiki/Clonally_transmissible_cancer" title="Clonally transmissible cancer">Clonally transmissible cancer</a></li></ul></li> <li><a href="/wiki/Virome" title="Virome">Virome</a></li></ul> </div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1" alt="" width="1" 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