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class="vector-toc-link" href="#Transmission"> <div class="vector-toc-text"> 3 Transmission </div> </a> <button aria-controls="toc-Transmission-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Transmission subsection </button> <ul id="toc-Transmission-sublist" class="vector-toc-list"> <li id="toc-Horizontal" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Horizontal"> <div class="vector-toc-text"> 3.1 Horizontal </div> </a> <ul id="toc-Horizontal-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Vertical" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Vertical"> <div class="vector-toc-text"> 3.2 Vertical </div> </a> <ul id="toc-Vertical-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Hosts" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Hosts"> <div class="vector-toc-text"> 4 Hosts </div> </a> <button aria-controls="toc-Hosts-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Hosts subsection </button> <ul id="toc-Hosts-sublist" class="vector-toc-list"> <li id="toc-Invertebrates" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Invertebrates"> <div class="vector-toc-text"> 4.1 Invertebrates </div> </a> <ul id="toc-Invertebrates-sublist" class="vector-toc-list"> <li id="toc-Insects" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Insects"> <div class="vector-toc-text"> 4.1.1 Insects </div> </a> <ul id="toc-Insects-sublist" class="vector-toc-list"> <li id="toc-Primary" class="vector-toc-list-item vector-toc-level-4"> <a class="vector-toc-link" href="#Primary"> <div class="vector-toc-text"> 4.1.1.1 Primary </div> </a> <ul id="toc-Primary-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Secondary" 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Available in 16 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-16" aria-hidden="true" > 16 languages </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/%D9%85%D8%B9%D8%A7%D9%8A%D8%B4_%D8%AC%D9%88%D8%A7%D9%86%D9%8A" title="معايش جواني – Arabic" lang="ar" hreflang="ar" data-title="معايش جواني" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target">العربية</a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%85%E0%A6%A8%E0%A7%8D%E0%A6%A4%E0%A6%83%E0%A6%AE%E0%A6%BF%E0%A6%A5%E0%A7%8B%E0%A6%9C%E0%A7%80%E0%A6%AC%E0%A7%80" title="অন্তঃমিথোজীবী – Bangla" lang="bn" hreflang="bn" data-title="অন্তঃমিথোজীবী" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target">বাংলা</a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Endosimbionte" title="Endosimbionte – Spanish" lang="es" hreflang="es" data-title="Endosimbionte" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target">Español</a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Endosimbioto" title="Endosimbioto – Esperanto" lang="eo" hreflang="eo" data-title="Endosimbioto" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target">Esperanto</a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%AF%D8%B1%D9%88%D9%86%E2%80%8C%D9%87%D9%85%D8%B2%DB%8C%D8%B3%D8%AA" title="درون‌همزیست – Persian" lang="fa" hreflang="fa" data-title="درون‌همزیست" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target">فارسی</a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Endosymbiote" title="Endosymbiote – French" lang="fr" hreflang="fr" data-title="Endosymbiote" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target">Français</a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%85%E0%A4%82%E0%A4%A4%E0%A4%B0%E0%A5%8D%E0%A4%B8%E0%A4%B9%E0%A4%9C%E0%A5%80%E0%A4%B5%E0%A5%80" title="अंतर्सहजीवी – Hindi" lang="hi" hreflang="hi" data-title="अंतर्सहजीवी" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target">हिन्दी</a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Endosimbion" title="Endosimbion – Indonesian" lang="id" hreflang="id" data-title="Endosimbion" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target">Bahasa Indonesia</a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%90%D7%A0%D7%93%D7%95%D7%A1%D7%99%D7%9E%D7%91%D7%99%D7%95%D7%A0%D7%98" title="אנדוסימביונט – Hebrew" lang="he" hreflang="he" data-title="אנדוסימביונט" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target">עברית</a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E5%86%85%E7%94%9F%E7%94%9F%E7%89%A9" title="内生生物 – Japanese" lang="ja" hreflang="ja" data-title="内生生物" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target">日本語</a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Endosymbiont" title="Endosymbiont – Polish" lang="pl" hreflang="pl" data-title="Endosymbiont" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target">Polski</a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Endossimbionte" title="Endossimbionte – Portuguese" lang="pt" hreflang="pt" data-title="Endossimbionte" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target">Português</a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%AD%D0%BD%D0%B4%D0%BE%D1%81%D0%B8%D0%BC%D0%B1%D0%B8%D0%BE%D0%BD%D1%82" title="Эндосимбионт – Russian" lang="ru" hreflang="ru" data-title="Эндосимбионт" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target">Русский</a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Endosimbiyont" title="Endosimbiyont – Turkish" lang="tr" hreflang="tr" data-title="Endosimbiyont" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target">Türkçe</a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%95%D0%BD%D0%B4%D0%BE%D1%81%D0%B8%D0%BC%D0%B1%D1%96%D0%BE%D0%BD%D1%82%D0%B8" title="Ендосимбіонти – Ukrainian" lang="uk" hreflang="uk" data-title="Ендосимбіонти" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target">Українська</a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/N%E1%BB%99i_c%E1%BB%99ng_sinh" title="Nội cộng sinh – Vietnamese" lang="vi" hreflang="vi" data-title="Nội cộng sinh" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target">Tiếng Việt</a></li> </ul> <div 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</div> </div> <div id="bodyContent" class="vector-body" aria-labelledby="firstHeading" data-mw-ve-target-container> <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">Organism that lives within the body or cells of another organism</div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Endosymbiosis.PNG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c5/Endosymbiosis.PNG/200px-Endosymbiosis.PNG" decoding="async" width="200" height="223" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c5/Endosymbiosis.PNG/300px-Endosymbiosis.PNG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c5/Endosymbiosis.PNG/400px-Endosymbiosis.PNG 2x" data-file-width="690" data-file-height="768" /></a><figcaption>A representation of the <a href="/wiki/Endosymbiotic_theory" class="mw-redirect" title="Endosymbiotic theory">endosymbiotic theory</a></figcaption></figure> An endosymbiont or endobiont<a href="#cite_note-KingDom-1">[1]</a> is an <a href="/wiki/Organism" title="Organism">organism</a> that lives within the body or cells of another organism. Typically the two organisms are in a <a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">mutualistic</a> relationship. Examples are <a href="/wiki/Nitrogen-fixing" class="mw-redirect" title="Nitrogen-fixing">nitrogen-fixing</a> <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> (called <a href="/wiki/Rhizobia" title="Rhizobia">rhizobia</a>), which live in the <a href="/wiki/Root_nodule" title="Root nodule">root nodules</a> of <a href="/wiki/Legume" title="Legume">legumes</a>, single-cell <a href="/wiki/Algae" title="Algae">algae</a> inside <a href="/wiki/Coral_reef" title="Coral reef">reef-building</a> <a href="/wiki/Coral" title="Coral">corals</a>, and bacterial endosymbionts that provide essential nutrients to <a href="/wiki/Insect" title="Insect">insects</a>.<a href="#cite_note-pmid29393944-2">[2]</a><a href="#cite_note-pmid15178799-3">[3]</a> Endosymbiosis played key roles in the development of <a href="/wiki/Eukaryotes" class="mw-redirect" title="Eukaryotes">eukaryotes</a> and plants. Roughly 2.2 billion years ago an <a href="/wiki/Archaeon" class="mw-redirect" title="Archaeon">archaeon</a> absorbed a <a href="/wiki/Bacterium" class="mw-redirect" title="Bacterium">bacterium</a> through <a href="/wiki/Phagocytosis" title="Phagocytosis">phagocytosis</a>, that eventually became the <a href="/wiki/Mitochondria" class="mw-redirect" title="Mitochondria">mitochondria</a> that provide energy to almost all living <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotic</a> cells. Approximately 1 billion years ago, some of those cells absorbed <a href="/wiki/Cyanobacteria" title="Cyanobacteria">cyanobacteria</a> that eventually became <a href="/wiki/Chloroplasts" class="mw-redirect" title="Chloroplasts">chloroplasts</a>, <a href="/wiki/Organelles" class="mw-redirect" title="Organelles">organelles</a> that produce energy from sunlight.<a href="#cite_note-4">[4]</a> Approximately 100 million years ago, a lineage of amoeba in the genus <a href="/wiki/Paulinella" title="Paulinella">Paulinella</a> independently engulfed a cyanobacteria that evolved to be functionally synonymous with traditional chloroplasts, called chromatophores.<a href="#cite_note-5">[5]</a> Some 100 million years ago, <a href="/wiki/UCYN-A" class="mw-redirect" title="UCYN-A">UCYN-A</a>, a nitrogen-fixing bacterium, became an endosymbiont of the marine alga <a href="/wiki/Braarudosphaera_bigelowii" title="Braarudosphaera bigelowii">Braarudosphaera bigelowii</a>, eventually evolving into a <a href="/wiki/Nitroplast" title="Nitroplast">nitroplast</a>, which fixes nitrogen.<a href="#cite_note-nature.com-6">[6]</a> Similarly, <a href="/wiki/Diatom" title="Diatom">diatoms</a> in the family Rhopalodiaceae have cyanobacterial endosymbionts, called spheroid bodies or diazoplasts, which have been proposed to be in the early stages of organelle evolution.<a href="#cite_note-7">[7]</a><a href="#cite_note-8">[8]</a> Symbionts are either obligate (require their host to survive) or facultative (can survive independently).<a href="#cite_note-Bright-2010-9">[9]</a> The most common examples of obligate endosymbiosis are <a href="/wiki/Mitochondria" class="mw-redirect" title="Mitochondria">mitochondria</a> and <a href="/wiki/Chloroplast" title="Chloroplast">chloroplasts</a>; however, they do not reproduce via <a href="/wiki/Mitosis" title="Mitosis">mitosis</a> in tandem with their host cells. Instead, they replicate via <a href="/wiki/Fission_(biology)" title="Fission (biology)">binary fission</a>, a replication process uncoupled from the host cells in which they reside.<a href="#cite_note-10">[10]</a><a href="#cite_note-11">[11]</a> Some human parasites, e.g. <a href="/wiki/Wuchereria_bancrofti" title="Wuchereria bancrofti">Wuchereria bancrofti</a> and <a href="/wiki/Mansonella_perstans" title="Mansonella perstans">Mansonella perstans</a>, thrive in their intermediate insect hosts because of an obligate endosymbiosis with <a href="/wiki/Wolbachia" title="Wolbachia">Wolbachia</a> spp.<a href="#cite_note-Slatko-2010-12">[12]</a> They can both be eliminated by treatments that target their bacterial host.<a href="#cite_note-13">[13]</a> <style data-mw-deduplicate="TemplateStyles:r886046785">.mw-parser-output .toclimit-2 .toclevel-1 ul,.mw-parser-output .toclimit-3 .toclevel-2 ul,.mw-parser-output .toclimit-4 .toclevel-3 ul,.mw-parser-output .toclimit-5 .toclevel-4 ul,.mw-parser-output .toclimit-6 .toclevel-5 ul,.mw-parser-output .toclimit-7 .toclevel-6 ul{display:none}</style><div class="toclimit-3"><meta property="mw:PageProp/toc" /></div> <div class="mw-heading mw-heading2"><h2 id="Etymology">Etymology</h2>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=1" title="Edit section: Etymology">edit</a>]</div> Endosymbiosis comes from the <a href="/wiki/Greek_language" title="Greek language">Greek</a>: ἔνδον endon "within", σύν syn "together" and βίωσις biosis "living". <div class="mw-heading mw-heading2"><h2 id="Symbiogenesis">Symbiogenesis</h2>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=2" title="Edit section: Symbiogenesis">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Endosymbiotic_theory.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/cf/Endosymbiotic_theory.svg/220px-Endosymbiotic_theory.svg.png" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/cf/Endosymbiotic_theory.svg/330px-Endosymbiotic_theory.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/cf/Endosymbiotic_theory.svg/440px-Endosymbiotic_theory.svg.png 2x" data-file-width="512" data-file-height="384" /></a><figcaption>An overview of the endosymbiosis theory of eukaryote origin (symbiogenesis).</figcaption></figure><a href="/wiki/Symbiogenesis" title="Symbiogenesis">Symbiogenesis</a> theory holds that eukaryotes evolved via absorbing <a href="/wiki/Prokaryotes" class="mw-redirect" title="Prokaryotes">prokaryotes</a>. Typically, one organism envelopes a bacterium and the two evolve a mutualistic relationship. The absorbed bacteria (the endosymbiont) eventually lives exclusively within the host cells. This fits the concept of observed organelle development.<a href="#cite_note-14">[14]</a><a href="#cite_note-McCutcheon-15">[15]</a><a href="#cite_note-Callier-16">[16]</a><a href="#cite_note-17">[17]</a><a href="#cite_note-Gabaldón-18">[18]</a> Typically the endosymbiont's genome shrinks, discarding genes whose roles are displaced by the host.<a href="#cite_note-pmid12415315-19">[19]</a> For example, the Hodgkinia genome of <a href="/wiki/Magicicada" class="mw-redirect" title="Magicicada">Magicicada</a> <a href="/wiki/Cicadas" class="mw-redirect" title="Cicadas">cicadas</a> is much different from the prior freestanding bacteria. The cicada life cycle involves years of stasis underground. The symbiont produces many generations during this phase, experiencing little <a href="/wiki/Natural_selection" title="Natural selection">selection pressure</a>, allowing their genomes to diversify. Selection is episodic (when the cicadas reproduce). The original Hodgkinia genome split into three much simpler endosymbionts, each encoding only a few genes—an instance of <a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">punctuated equilibrium</a> producing distinct lineages. The host requires all three symbionts.<a href="#cite_note-pmid29129532-20">[20]</a> <div class="mw-heading mw-heading2"><h2 id="Transmission">Transmission</h2>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=3" title="Edit section: Transmission">edit</a>]</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 articles: <a href="/wiki/Horizontal_transmission" title="Horizontal transmission">Horizontal transmission</a> and <a href="/wiki/Vertical_transmission" title="Vertical transmission">Vertical transmission</a></div> Symbiont transmission is the process where the host acquires its symbiont. Since symbionts are not produced by host cells, they must find their own way to reproduce and populate daughter cells as host cells divide. Horizontal, vertical, and mixed-mode (hybrid of horizonal and vertical) transmission are the three paths for symbiont transfer. <div class="mw-heading mw-heading3"><h3 id="Horizontal">Horizontal</h3>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=4" title="Edit section: Horizontal">edit</a>]</div> Horizontal symbiont transfer (<a href="/wiki/Horizontal_transmission" title="Horizontal transmission">horizontal transmission</a>) is a process where a host acquires a facultative symbiont from the environment or another host.<a href="#cite_note-Bright-2010-9">[9]</a> The Rhizobia-Legume symbiosis (bacteria-plant endosymbiosis) is a prime example of this modality.<a href="#cite_note-Gage-2004-21">[21]</a> The Rhizobia-legume symbiotic relationship is important for processes such as the formation of root nodules. It starts with flavonoids released by the legume host, which causes the rhizobia species (endosymbiont) to activate its Nod genes.<a href="#cite_note-Gage-2004-21">[21]</a> These Nod genes generate <a href="/wiki/Lipopolysaccharide" title="Lipopolysaccharide">lipooligosaccharide</a> signals that the legume detects, leading to root nodule formation.<a href="#cite_note-pmid109930772-22">[22]</a> This process bleeds into other processes such as nitrogen fixation in plants.<a href="#cite_note-Gage-2004-21">[21]</a> The evolutionary advantage of such an interaction allows genetic exchange between both organisms involved to increase the propensity for novel functions as seen in the plant-bacterium interaction (<a href="/wiki/Holobiont" title="Holobiont">holobiont</a> formation).<a href="#cite_note-23">[23]</a> <div class="mw-heading mw-heading3"><h3 id="Vertical">Vertical</h3>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=5" title="Edit section: Vertical">edit</a>]</div> Vertical transmission takes place when the symbiont moves directly from parent to offspring.<a href="#cite_note-McCutcheon2-24">[24]</a><a href="#cite_note-Callier2-25">[25]</a> In horizontal transmission each generation acquires symbionts from the environment. An example is nitrogen-fixing bacteria in certain plant roots, such as <a href="/wiki/Acyrthosiphon_pisum" title="Acyrthosiphon pisum">pea aphid</a> symbionts. A third type is mixed-mode transmission, where symbionts move horizontally for some generations, after which they are acquired vertically.<a href="#cite_note-Wierz2-26">[26]</a><a href="#cite_note-Ebert2-27">[27]</a><a href="#cite_note-pmid201573403-28">[28]</a> <a href="/wiki/Wigglesworthia" class="mw-redirect" title="Wigglesworthia">Wigglesworthia</a>, a tsetse fly symbiont,<a href="#cite_note-pmid201573403-28">[28]</a> is vertically transmitted (via mother's milk).<a href="#cite_note-pmid201573403-28">[28]</a> In <a href="/wiki/Vertical_transmission_(symbiont)" class="mw-redirect" title="Vertical transmission (symbiont)">vertical transmission</a>, the symbionts do not need to survive independently, often leading them to have a reduced genome. For instance, <a href="/wiki/Acyrthosiphon_pisum" title="Acyrthosiphon pisum">pea aphid</a> symbionts have lost genes for essential molecules and rely on the host to supply them. In return, the symbionts synthesize essential <a href="/wiki/Amino_acids" class="mw-redirect" title="Amino acids">amino acids</a> for the aphid host.<a href="#cite_note-pmid109930772-22">[22]</a> When a symbiont reaches this stage, it begins to resemble a cellular <a href="/wiki/Organelle" title="Organelle">organelle</a>, similar to <a href="/wiki/Mitochondria" class="mw-redirect" title="Mitochondria">mitochondria</a> or <a href="/wiki/Chloroplasts" class="mw-redirect" title="Chloroplasts">chloroplasts</a>. Such dependent hosts and symbionts form a <a href="/wiki/Holobiont" title="Holobiont">holobiont</a>. In the event of a bottleneck, a decrease in symbiont diversity could compromise host-symbiont interactions, as deleterious mutations accumulate.<a href="#cite_note-29">[29]</a> <div class="mw-heading mw-heading2"><h2 id="Hosts">Hosts</h2>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=6" title="Edit section: Hosts">edit</a>]</div> <div class="mw-heading mw-heading3"><h3 id="Invertebrates">Invertebrates</h3>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=7" title="Edit section: Invertebrates">edit</a>]</div> The best-studied examples of endosymbiosis are in <a href="/wiki/Invertebrate" title="Invertebrate">invertebrates</a>. These symbioses affect organisms with global impact, including <a href="/wiki/Symbiodinium" title="Symbiodinium">Symbiodinium</a> (corals), or <a href="/wiki/Wolbachia" title="Wolbachia">Wolbachia</a> (insects). Many insect agricultural pests and human disease vectors have intimate relationships with primary endosymbionts.<a href="#cite_note-30">[30]</a> <div class="mw-heading mw-heading4"><h4 id="Insects">Insects</h4>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=8" title="Edit section: Insects">edit</a>]</div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Cospeciation_(5_processes)_-_with_key.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/51/Cospeciation_%285_processes%29_-_with_key.png/220px-Cospeciation_%285_processes%29_-_with_key.png" decoding="async" width="220" height="352" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/51/Cospeciation_%285_processes%29_-_with_key.png/330px-Cospeciation_%285_processes%29_-_with_key.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/51/Cospeciation_%285_processes%29_-_with_key.png/440px-Cospeciation_%285_processes%29_-_with_key.png 2x" data-file-width="2400" data-file-height="3837" /></a><figcaption>Diagram of cospeciation, where parasites or endosymbionts speciate or branch alongside their hosts. This process is more common in hosts with primary endosymbionts.</figcaption></figure> Scientists classify insect endosymbionts as Primary or Secondary. Primary endosymbionts (P-endosymbionts) have been associated with their <a href="/wiki/Insect" title="Insect">insect</a> hosts for millions of years (from ten to several hundred million years). They form obligate associations and display <a href="/wiki/Cospeciation" title="Cospeciation">cospeciation</a> with their insect hosts. Secondary endosymbionts more recently associated with their hosts, may be horizontally transferred, live in the <a href="/wiki/Hemolymph" title="Hemolymph">hemolymph</a> of the insects (not specialized bacteriocytes, see below), and are not obligate.<a href="#cite_note-31">[31]</a> <div class="mw-heading mw-heading5"><h5 id="Primary">Primary</h5>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=9" title="Edit section: Primary">edit</a>]</div> Among primary endosymbionts of insects, the best-studied are the pea <a href="/wiki/Aphid" title="Aphid">aphid</a> (<a href="/wiki/Acyrthosiphon_pisum" title="Acyrthosiphon pisum">Acyrthosiphon pisum</a>) and its endosymbiont <a href="/wiki/Buchnera_(proteobacteria)" class="mw-redirect" title="Buchnera (proteobacteria)">Buchnera</a> sp. APS,<a href="#cite_note-32">[32]</a><a href="#cite_note-pmid109930772-22">[22]</a> the <a href="/wiki/Tsetse_fly" title="Tsetse fly">tsetse fly</a> Glossina morsitans morsitans and its endosymbiont <a href="/wiki/Wigglesworthia_glossinidia_brevipalpis" class="mw-redirect" title="Wigglesworthia glossinidia brevipalpis">Wigglesworthia glossinidia brevipalpis</a> and the endosymbiotic <a href="/wiki/Protists" class="mw-redirect" title="Protists">protists</a> in lower <a href="/wiki/Termite" title="Termite">termites</a>. As with endosymbiosis in other insects, the symbiosis is obligate. Nutritionally-enhanced diets allow symbiont-free specimens to survive, but they are unhealthy, and at best survive only a few generations.<a href="#cite_note-33">[33]</a> In some insect groups, these endosymbionts live in specialized insect cells called <a href="/wiki/Bacteriocyte" title="Bacteriocyte">bacteriocytes</a> (also called mycetocytes), and are maternally-transmitted, i.e. the mother transmits her endosymbionts to her offspring. In some cases, the bacteria are transmitted in the <a href="/wiki/Egg_(biology)" class="mw-redirect" title="Egg (biology)">egg</a>, as in Buchnera; in others like Wigglesworthia, they are transmitted via milk to the embryo. In termites, the endosymbionts reside within the hindguts and are transmitted through <a href="/wiki/Trophallaxis" title="Trophallaxis">trophallaxis</a> among colony members.<a href="#cite_note-34">[34]</a> Primary endosymbionts are thought to help the host either by providing essential nutrients or by metabolizing insect waste products into safer forms. For example, the putative primary role of Buchnera is to synthesize <a href="/wiki/Essential_amino_acid" title="Essential amino acid">essential amino acids</a> that the aphid cannot acquire from its diet of plant sap. The primary role of Wigglesworthia is to synthesize <a href="/wiki/Vitamin" title="Vitamin">vitamins</a> that the tsetse fly does not get from the <a href="/wiki/Blood" title="Blood">blood</a> that it eats. In lower termites, the endosymbiotic protists play a major role in the digestion of lignocellulosic materials that constitute a bulk of the termites' diet. Bacteria benefit from the reduced exposure to <a href="/wiki/Predator" class="mw-redirect" title="Predator">predators</a> and competition from other bacterial species, the ample supply of nutrients and relative environmental stability inside the host. Primary endosymbionts of insects have among the smallest of known bacterial genomes and have <a href="/wiki/Genome_reduction" class="mw-redirect" title="Genome reduction">lost many genes</a> commonly found in closely related bacteria. One theory claimed that some of these genes are not needed in the environment of the host insect cell. A complementary theory suggests that the relatively small numbers of bacteria inside each insect decrease the efficiency of natural selection in 'purging' deleterious mutations and small mutations from the population, resulting in a loss of genes over many millions of years. Research in which a parallel <a href="/wiki/Phylogeny" class="mw-redirect" title="Phylogeny">phylogeny</a> of bacteria and insects was inferred supports the assumption hat primary endosymbionts are transferred only vertically.<a href="#cite_note-35">[35]</a><a href="#cite_note-36">[36]</a> Attacking obligate bacterial endosymbionts may present a way to control their hosts, many of which are pests or human disease carriers. For example, aphids are crop pests and the tsetse fly carries the organism <a href="/wiki/Trypanosoma_brucei" title="Trypanosoma brucei">Trypanosoma brucei</a> that causes African <a href="/wiki/African_trypanosomiasis" title="African trypanosomiasis">sleeping sickness</a>.<a href="#cite_note-37">[37]</a> Studying insect endosymbionts can aid understanding the origins of symbioses in general, as a proxy for understanding endosymbiosis in other species. The best-studied ant endosymbionts are <a href="/wiki/Blochmannia" title="Blochmannia">Blochmannia</a> bacteria, which are the primary endosymbiont of <a href="/wiki/Camponotus" class="mw-redirect" title="Camponotus">Camponotus</a> ants. In 2018 a new ant-associated symbiont, Candidatus Westeberhardia Cardiocondylae, was discovered in <a href="/wiki/Cardiocondyla" title="Cardiocondyla">Cardiocondyla</a>. It is reported to be a primary symbiont.<a href="#cite_note-38">[38]</a> <div class="mw-heading mw-heading5"><h5 id="Secondary">Secondary</h5>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=10" title="Edit section: Secondary">edit</a>]</div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:HEMI_Aphididae_Aphidius_attacking_pea_aphid.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b7/HEMI_Aphididae_Aphidius_attacking_pea_aphid.png/220px-HEMI_Aphididae_Aphidius_attacking_pea_aphid.png" decoding="async" width="220" height="188" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b7/HEMI_Aphididae_Aphidius_attacking_pea_aphid.png/330px-HEMI_Aphididae_Aphidius_attacking_pea_aphid.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b7/HEMI_Aphididae_Aphidius_attacking_pea_aphid.png/440px-HEMI_Aphididae_Aphidius_attacking_pea_aphid.png 2x" data-file-width="3542" data-file-height="3020" /></a><figcaption>Pea aphids are commonly infested by parasitic wasps. Their secondary endosymbionts attack the infesting parasitoid wasp larvae promoting the survival of both the aphid host and its endosymbionts.</figcaption></figure> The pea aphid (<a href="/wiki/Acyrthosiphon_pisum" title="Acyrthosiphon pisum">Acyrthosiphon pisum</a>) contains at least three secondary endosymbionts, <a href="/wiki/Hamiltonella_defensa" title="Hamiltonella defensa">Hamiltonella defensa</a>, <a href="/wiki/Regiella_insecticola" title="Regiella insecticola">Regiella insecticola</a>, and <a href="/wiki/Serratia_symbiotica" title="Serratia symbiotica">Serratia symbiotica</a>. Hamiltonella defensa defends its aphid host from parasitoid wasps.<a href="#cite_note-pmid18029301-39">[39]</a> This symbiosis replaces lost elements of the insect's immune response.<a href="#cite_note-pmid20186266-40">[40]</a> One of the best-understood defensive symbionts is the spiral bacteria <a href="/wiki/Spiroplasma_poulsonii" title="Spiroplasma poulsonii">Spiroplasma poulsonii</a>. Spiroplasma sp. can be reproductive manipulators, but also defensive symbionts of <a href="/wiki/Drosophila" title="Drosophila">Drosophila</a> flies. In <a href="/wiki/Drosophila_neotestacea" title="Drosophila neotestacea">Drosophila neotestacea</a>, S. poulsonii has spread across North America owing to its ability to defend its fly host against <a href="/wiki/Nematode" title="Nematode">nematode</a> parasites.<a href="#cite_note-41">[41]</a> This defence is mediated by toxins called "<a href="/wiki/Ribosome" title="Ribosome">ribosome</a>-inactivating <a href="/wiki/Proteins" class="mw-redirect" title="Proteins">proteins</a>" that attack the molecular machinery of invading parasites.<a href="#cite_note-42">[42]</a><a href="#cite_note-43">[43]</a> These toxins represent one of the first understood examples of a defensive symbiosis with a mechanistic understanding for defensive symbiosis between an insect endosymbiont and its host.<a href="#cite_note-Ballinger-2017-44">[44]</a> <a href="/wiki/Sodalis_glossinidius" title="Sodalis glossinidius">Sodalis glossinidius</a> is a secondary endosymbiont of tsetse flies that lives inter- and intracellularly in various host tissues, including the midgut and hemolymph. Phylogenetic studies do not report a correlation between evolution of <a href="/wiki/Sodalis_(genus)" title="Sodalis (genus)">Sodalis</a> and tsetse.<a href="#cite_note-45">[45]</a> Unlike Wigglesworthia, Sodalis has been cultured in vitro.<a href="#cite_note-pmid3662675-46">[46]</a> <a href="/wiki/Cardinium" title="Cardinium">Cardinium</a> and many other insects have secondary endosymbionts.<a href="#cite_note-pmid15189221-47">[47]</a><a href="#cite_note-pmid12415315-19">[19]</a> <div class="mw-heading mw-heading4"><h4 id="Marine">Marine</h4>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=11" title="Edit section: Marine">edit</a>]</div> Extracellular endosymbionts are represented in all four extant classes of <a href="/wiki/Echinodermata" class="mw-redirect" title="Echinodermata">Echinodermata</a> (<a href="/wiki/Crinoidea" class="mw-redirect" title="Crinoidea">Crinoidea</a>, <a href="/wiki/Ophiuroidea" class="mw-redirect" title="Ophiuroidea">Ophiuroidea</a>, <a href="/wiki/Echinoidea" class="mw-redirect" title="Echinoidea">Echinoidea</a>, and <a href="/wiki/Holothuroidea" class="mw-redirect" title="Holothuroidea">Holothuroidea</a>). Little is known of the nature of the association (mode of infection, transmission, metabolic requirements, etc.) but <a href="/wiki/Phylogenetic" class="mw-redirect" title="Phylogenetic">phylogenetic</a> analysis indicates that these symbionts belong to the class <a href="/wiki/Alphaproteobacteria" title="Alphaproteobacteria">Alphaproteobacteria</a>, relating them to <a href="/wiki/Rhizobium" title="Rhizobium">Rhizobium</a> and <a href="/wiki/Thiobacillus" title="Thiobacillus">Thiobacillus</a>. Other studies indicate that these subcuticular bacteria may be both abundant within their hosts and widely distributed among the Echinoderms.<a href="#cite_note-48">[48]</a> Some marine <a href="/wiki/Oligochaeta" title="Oligochaeta">oligochaeta</a> (e.g., <a href="/wiki/Olavius_algarvensis" title="Olavius algarvensis">Olavius algarvensis</a> and <a href="/wiki/Inanidrilus" title="Inanidrilus">Inanidrillus</a> spp.) have obligate extracellular endosymbionts that fill the entire body of their host. These marine worms are nutritionally dependent on their symbiotic <a href="/wiki/Chemoautotroph" class="mw-redirect" title="Chemoautotroph">chemoautotrophic</a> bacteria lacking any digestive or excretory system (no gut, mouth, or <a href="/wiki/Nephridia" class="mw-redirect" title="Nephridia">nephridia</a>).<a href="#cite_note-49">[49]</a> The sea slug <a href="/wiki/Elysia_chlorotica" title="Elysia chlorotica">Elysia chlorotica</a>'s endosymbiont is the <a href="/wiki/Algae" title="Algae">algae</a> <a href="/wiki/Vaucheria_litorea" title="Vaucheria litorea">Vaucheria litorea</a>. The <a href="/wiki/Jellyfish" title="Jellyfish">jellyfish</a> <a href="/wiki/Mastigias" title="Mastigias">Mastigias</a> have a similar relationship with an algae. <a href="/wiki/Elysia_chlorotica" title="Elysia chlorotica">Elysia chlorotica</a> forms this relationship intracellularly with the algae's chloroplasts. These chloroplasts retain their photosynthetic capabilities and structures for several months after entering the slug's cells.<a href="#cite_note-50">[50]</a> <a href="/wiki/Trichoplax" title="Trichoplax">Trichoplax</a> have two bacterial endosymbionts. Ruthmannia lives inside the animal's digestive cells. Grellia lives permanently inside the <a href="/wiki/Endoplasmic_reticulum" title="Endoplasmic reticulum">endoplasmic reticulum</a> (ER), the first known symbiont to do so.<a href="#cite_note-51">[51]</a> <a href="/wiki/Paracatenula" title="Paracatenula">Paracatenula</a> is a <a href="/wiki/Flatworm" title="Flatworm">flatworm</a> which have lived in symbiosis with an endosymbiotic bacteria for 500 million years. The bacteria produce numerous small, droplet-like vesicles that provide the host with needed nutrients.<a href="#cite_note-52">[52]</a> <div class="mw-heading mw-heading5"><h5 id="Dinoflagellates">Dinoflagellates</h5>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=12" title="Edit section: Dinoflagellates">edit</a>]</div> <a href="/wiki/Dinoflagellate" title="Dinoflagellate">Dinoflagellate</a> endosymbionts of the genus <a href="/wiki/Symbiodinium" title="Symbiodinium">Symbiodinium</a>, commonly known as <a href="/wiki/Zooxanthella" class="mw-redirect" title="Zooxanthella">zooxanthellae</a>, are found in <a href="/wiki/Corals" class="mw-redirect" title="Corals">corals</a>, <a href="/wiki/Mollusk" class="mw-redirect" title="Mollusk">mollusks</a> (esp. <a href="/wiki/Giant_clam" title="Giant clam">giant clams</a>, the Tridacna), <a href="/wiki/Sea_sponge" class="mw-redirect" title="Sea sponge">sponges</a>, and the unicellular <a href="/wiki/Foraminifera" title="Foraminifera">foraminifera</a>. These endosymbionts capture sunlight and provide their hosts with energy via <a href="/wiki/Carbonate" title="Carbonate">carbonate</a> deposition.<a href="#cite_note-Baker2003-53">[53]</a> Previously thought to be a single species, molecular <a href="/wiki/Phylogenetic" class="mw-redirect" title="Phylogenetic">phylogenetic</a> evidence reported diversity in Symbiodinium. In some cases, the host requires a specific Symbiodinium <a href="/wiki/Clade" title="Clade">clade</a>. More often, however, the distribution is ecological, with symbionts switching among hosts with ease. When reefs become environmentally stressed, this distribution is related to the observed pattern of <a href="/wiki/Coral_bleaching" title="Coral bleaching">coral bleaching</a> and recovery. Thus, the distribution of Symbiodinium on coral reefs and its role in coral bleaching is an important in coral reef ecology.<a href="#cite_note-Baker2003-53">[53]</a> <div class="mw-heading mw-heading4"><h4 id="Phytoplankton">Phytoplankton</h4>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=13" title="Edit section: Phytoplankton">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Microbial_loop" title="Microbial loop">Microbial loop</a></div> In marine environments,<a href="#cite_note-Villareal-1994-54">[54]</a><a href="#cite_note-Carpenter-1999-55">[55]</a><a href="#cite_note-Foster-2007-56">[56]</a><a href="#cite_note-57">[57]</a> endosymbiont relationships are especially prevalent in <a href="/wiki/Trophic_state_index" title="Trophic state index">oligotrophic</a> or nutrient-poor regions of the ocean like that of the North Atlantic.<a href="#cite_note-Villareal-1994-54">[54]</a><a href="#cite_note-Goebel-2010-58">[58]</a><a href="#cite_note-Carpenter-1999-55">[55]</a><a href="#cite_note-Foster-2007-56">[56]</a> In such waters, cell growth of larger <a href="/wiki/Phytoplankton" title="Phytoplankton">phytoplankton</a> such as <a href="/wiki/Diatom" title="Diatom">diatoms</a> is limited by (insufficient) <a href="/wiki/Nitrate" title="Nitrate">nitrate</a> concentrations.<a href="#cite_note-Foster-2011-59">[59]</a>  Endosymbiotic bacteria fix nitrogen for their hosts and in turn receive organic carbon from photosynthesis.<a href="#cite_note-Goebel-2010-58">[58]</a> These symbioses play an important role in global <a href="/wiki/Carbon_cycle" title="Carbon cycle">carbon cycling</a>.<a href="#cite_note-60">[60]</a><a href="#cite_note-Carpenter-1999-55">[55]</a><a href="#cite_note-Foster-2007-56">[56]</a> One known symbiosis between the diatom <a href="/wiki/Cyanobiont#Diatoms" title="Cyanobiont">Hemialus</a> spp. and the cyanobacterium <a href="/wiki/Richelia_intracellularis" class="mw-redirect" title="Richelia intracellularis">Richelia intracellularis</a> has been reported in North Atlantic, Mediterranean, and Pacific waters.<a href="#cite_note-Villareal-1994-54">[54]</a><a href="#cite_note-Carpenter-1999-55">[55]</a><a href="#cite_note-61">[61]</a> Richelia is found within the <a href="/wiki/Diatom_frustule" class="mw-redirect" title="Diatom frustule">diatom frustule</a> of Hemiaulus spp., and has a reduced genome.<a href="#cite_note-Hilton-2013-62">[62]</a> A 2011 study measured nitrogen fixation by the <a href="/wiki/Cyanobacteria" title="Cyanobacteria">cyanobacterial</a> host Richelia intracellularis well above intracellular requirements, and found the cyanobacterium was likely fixing nitrogen for its host.<a href="#cite_note-Foster-2011-59">[59]</a> Additionally, both host and symbiont cell growth were much greater than free-living Richelia intracellularis or symbiont-free Hemiaulus spp.<a href="#cite_note-Foster-2011-59">[59]</a> The Hemaiulus-Richelia symbiosis is not obligatory, especially in nitrogen-replete areas.<a href="#cite_note-Villareal-1994-54">[54]</a> Richelia intracellularis is also found in Rhizosolenia spp., a diatom found in oligotrophic oceans.<a href="#cite_note-Goebel-2010-58">[58]</a><a href="#cite_note-Foster-2011-59">[59]</a><a href="#cite_note-Foster-2007-56">[56]</a> Compared to the Hemaiulus host, the endosymbiosis with Rhizosolenia is much more consistent, and Richelia intracellularis is generally found in Rhizosolenia.<a href="#cite_note-Villareal-1994-54">[54]</a> There are some asymbiotic (occurs without an endosymbiont) Rhizosolenia, however there appears to be mechanisms limiting growth of these organisms in low nutrient conditions.<a href="#cite_note-Villareal-1989-63">[63]</a> Cell division for both the diatom host and cyanobacterial symbiont can be uncoupled and mechanisms for passing bacterial symbionts to daughter cells during cell division are still relatively unknown.<a href="#cite_note-Villareal-1989-63">[63]</a> Other endosymbiosis with nitrogen fixers in open oceans include <a href="/wiki/Calothrix" title="Calothrix">Calothrix</a> in <a href="/wiki/Chaetoceros" title="Chaetoceros">Chaetoceros</a> spp. and UNCY-A in <a href="/wiki/Prymnesiophyte" class="mw-redirect" title="Prymnesiophyte">prymnesiophyte</a> microalga.<a href="#cite_note-Zehr-2015-64">[64]</a>  The Chaetoceros-Calothrix endosymbiosis is hypothesized to be more recent, as the Calothrix genome is generally intact. While other species like that of the UNCY-A symbiont and Richelia have reduced genomes.<a href="#cite_note-Hilton-2013-62">[62]</a> This reduction in genome size occurs within nitrogen metabolism pathways indicating endosymbiont species are generating nitrogen for their hosts and losing the ability to use this nitrogen independently.<a href="#cite_note-Hilton-2013-62">[62]</a> This endosymbiont reduction in genome size, might be a step that occurred in the evolution of organelles (above).<a href="#cite_note-Zehr-2015-64">[64]</a> <div class="mw-heading mw-heading3"><h3 id="Protists">Protists</h3>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=14" title="Edit section: Protists">edit</a>]</div> <a href="/wiki/Mixotricha_paradoxa" title="Mixotricha paradoxa">Mixotricha paradoxa</a> is a <a href="/wiki/Protozoan" class="mw-redirect" title="Protozoan">protozoan</a> that lacks mitochondria. However, spherical bacteria live inside the cell and serve the function of the mitochondria. Mixotricha has three other species of symbionts that live on the surface of the cell.<a href="#cite_note-65">[65]</a> <a href="/wiki/Paramecium_bursaria" title="Paramecium bursaria">Paramecium bursaria</a>, a species of <a href="/wiki/Ciliate" title="Ciliate">ciliate</a>, has a mutualistic symbiotic relationship with green alga called <a href="/wiki/Zoochlorella" title="Zoochlorella">Zoochlorella</a>. The algae live in its cytoplasm.<a href="#cite_note-66">[66]</a> Platyophrya chlorelligera is a freshwater ciliate that harbors <a href="/wiki/Chlorella" title="Chlorella">Chlorella</a> that perform photosynthesis.<a href="#cite_note-67">[67]</a><a href="#cite_note-68">[68]</a> Strombidium purpureum is a marine ciliate that uses endosymbiotic, purple, non-sulphur bacteria for anoxygenic photosynthesis.<a href="#cite_note-69">[69]</a><a href="#cite_note-70">[70]</a> <a href="/wiki/Paulinella" title="Paulinella">Paulinella chromatophora</a> is a freshwater <a href="/wiki/Amoeboid" class="mw-redirect" title="Amoeboid">amoeboid</a> that has a <a href="/wiki/Cyanobacterium" class="mw-redirect" title="Cyanobacterium">cyanobacterium</a> endosymbiont. Many <a href="/wiki/Foraminifera" title="Foraminifera">foraminifera</a> are hosts to several types of algae, such as <a href="/wiki/Red_algae" title="Red algae">red algae</a>, <a href="/wiki/Diatom" title="Diatom">diatoms</a>, <a href="/wiki/Dinoflagellate" title="Dinoflagellate">dinoflagellates</a> and <a href="/wiki/Chlorophyta" title="Chlorophyta">chlorophyta</a>.<a href="#cite_note-71">[71]</a> These endosymbionts can be transmitted vertically to the next generation via asexual reproduction of the host, but because the endosymbionts are larger than the foraminiferal <a href="/wiki/Gamete" title="Gamete">gametes</a>, they need to acquire algae horizontally following sexual reproduction.<a href="#cite_note-72">[72]</a> Several species of <a href="/wiki/Radiolaria" title="Radiolaria">radiolaria</a> have photosynthetic symbionts. In some species the host digests algae to keep the population at a constant level.<a href="#cite_note-73">[73]</a> <a href="/wiki/Hatena_arenicola" title="Hatena arenicola">Hatena arenicola</a> is a flagellate <a href="/wiki/Protist" title="Protist">protist</a> with a complicated feeding apparatus that feeds on other microbes. When it engulfs a green <a href="/wiki/Nephroselmis" title="Nephroselmis">Nephroselmis</a> alga, the feeding apparatus disappears and it becomes photosynthetic. During <a href="/wiki/Mitosis" title="Mitosis">mitosis</a> the algae is transferred to only one of the daughter cells, while the other cell restarts the cycle. In 1966, biologist Kwang W. Jeon found that a lab strain of <a href="/wiki/Amoeba_proteus" title="Amoeba proteus">Amoeba proteus</a> had been infected by bacteria that lived inside the cytoplasmic <a href="/wiki/Vacuoles" class="mw-redirect" title="Vacuoles">vacuoles</a>.<a href="#cite_note-74">[74]</a> This infection killed almost all of the infected protists. After the equivalent of 40 host generations, the two organisms become mutually interdependent. A genetic exchange between the <a href="/wiki/Prokaryotes" class="mw-redirect" title="Prokaryotes">prokaryotes</a> and protists occurred.<a href="#cite_note-75">[75]</a><a href="#cite_note-76">[76]</a><a href="#cite_note-77">[77]</a> <div class="mw-heading mw-heading3"><h3 id="Vertebrates">Vertebrates</h3>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=15" title="Edit section: Vertebrates">edit</a>]</div> The spotted salamander (<a href="/wiki/Ambystoma_maculatum" class="mw-redirect" title="Ambystoma maculatum">Ambystoma maculatum</a>) lives in a relationship with the algae <a href="/wiki/Oophila_amblystomatis" class="mw-redirect" title="Oophila amblystomatis">Oophila amblystomatis</a>, which grows in its egg cases.<a href="#cite_note-pmid21464324-78">[78]</a> <div class="mw-heading mw-heading3"><h3 id="Plants">Plants</h3>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=16" title="Edit section: Plants">edit</a>]</div> All vascular plants harbor endosymbionts or endophytes in this context. They include <a href="/wiki/Bacteria" title="Bacteria">bacteria</a>, <a href="/wiki/Fungus" title="Fungus">fungi</a>, <a href="/wiki/Virus" title="Virus">viruses</a>, <a href="/wiki/Protozoa" title="Protozoa">protozoa</a> and even <a href="/wiki/Microalgae" title="Microalgae">microalgae</a>. Endophytes aid in processes such as growth and development, nutrient uptake, and defense against biotic and abiotic stresses like <a href="/wiki/Drought" title="Drought">drought</a>, <a href="/wiki/Salinity" title="Salinity">salinity</a>, heat, and herbivores.<a href="#cite_note-Baron_39–55-79">[79]</a> Plant symbionts can be categorized into <a href="/wiki/Epiphyte" title="Epiphyte">epiphytic</a>, <a href="/wiki/Endophyte" title="Endophyte">endophytic</a>, and <a href="/wiki/Mycorrhiza" title="Mycorrhiza">mycorrhizal</a>. These relations can also be categorized as beneficial, <a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">mutualistic</a>, neutral, and <a href="/wiki/Pathogen" title="Pathogen">pathogenic</a>.<a href="#cite_note-Hardoim_293–320-80">[80]</a><a href="#cite_note-81">[81]</a> <a href="/wiki/Microorganism" title="Microorganism">Microorganisms</a> living as endosymbionts in plants can enhance their host's primary productivity either by producing or capturing important resources.<a href="#cite_note-Fungal_plant_endosymbionts_alter_li-82">[82]</a> These endosymbionts can also enhance plant productivity by producing toxic metabolites that aid plant defenses against <a href="/wiki/Herbivore" title="Herbivore">herbivores</a>.<a href="#cite_note-83">[83]</a><a href="#cite_note-84">[84]</a> Plants are dependent on <a href="/wiki/Plastid" title="Plastid">plastid</a> or <a href="/wiki/Chloroplast" title="Chloroplast">chloroplast</a> organelles. The chloroplast is derived from a cyanobacterial primary endosymbiosis that began over one billion years ago. An oxygenic, photosynthetic free-living <a href="/wiki/Cyanobacteria" title="Cyanobacteria">cyanobacterium</a> was engulfed and kept by a heterotrophic <a href="/wiki/Protist" title="Protist">protist</a> and eventually evolved into the present intracellular organelle.<a href="#cite_note-85">[85]</a>   Mycorrhizal endosymbionts appear only in <a href="/wiki/Fungus" title="Fungus">fungi</a>. Typically, plant endosymbiosis studies focus on a single category or species to better understand their individual biological processes and functions.<a href="#cite_note-86">[86]</a> <div class="mw-heading mw-heading4"><h4 id="Fungal_endophytes">Fungal endophytes</h4>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=17" title="Edit section: Fungal endophytes">edit</a>]</div> Fungal endophytes can be found in all plant tissues. Fungi living below the ground amidst plant roots are known as <a href="/wiki/Mycorrhiza" title="Mycorrhiza">mycorrhiza</a>, but are further categorized based on their location inside the root, with prefixes such as ecto, endo, arbuscular, ericoid, etc. Fungal endosymbionts that live in the roots and extend their extraradical <a href="/wiki/Hyphae" class="mw-redirect" title="Hyphae">hyphae</a> into the outer <a href="/wiki/Rhizosphere" title="Rhizosphere">rhizosphere</a> are known as ectendosymbionts.<a href="#cite_note-Salhi-2022-87">[87]</a><a href="#cite_note-88">[88]</a> <div class="mw-heading mw-heading5"><h5 id="Arbuscular_Mycorrhizal_Fungi_(AMF)">Arbuscular Mycorrhizal Fungi (AMF)</h5>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=18" title="Edit section: Arbuscular Mycorrhizal Fungi (AMF)">edit</a>]</div> <a href="/wiki/Arbuscular_mycorrhiza" title="Arbuscular mycorrhiza">Arbuscular mycorrhizal fungi</a> or AMF are the most diverse plant microbial endosymbionts. With exceptions such as the <a href="/wiki/Ericaceae" title="Ericaceae">Ericaceae</a> family, almost all vascular plants harbor <a href="/wiki/Arbuscular_mycorrhiza" title="Arbuscular mycorrhiza">AMF</a> endosymbionts as endo and ecto as well. AMF plant endosymbionts systematically colonize <a href="/wiki/Root" title="Root">plant roots</a> and help the plant host acquire soil <a href="/wiki/Nutrient" title="Nutrient">nutrients</a> such as nitrogen. In return it absorbs plant organic carbon products.<a href="#cite_note-Salhi-2022-87">[87]</a> <a href="/wiki/Plant_root_exudates" title="Plant root exudates">Plant root exudates</a> contain diverse secondary metabolites, especially <a href="/wiki/Flavonoids" class="mw-redirect" title="Flavonoids">flavonoids</a> and <a href="/w/index.php?title=Strigolactones&action=edit&redlink=1" class="new" title="Strigolactones (page does not exist)">strigolactones</a> that act as <a href="/wiki/Plant_communication" title="Plant communication">chemical signals</a> and attracts the AMF.<a href="#cite_note-89">[89]</a> AMF <a href="/wiki/Gigasporaceae" title="Gigasporaceae">Gigaspora</a> margarita lives as a plant endosymbiont and also harbors further endosymbiont intracytoplasmic bacterium-like organisms.<a href="#cite_note-90">[90]</a> AMF generally promote plant health and growth and alleviate <a href="/wiki/Abiotic_stress" title="Abiotic stress">abiotic stresses</a> such as salinity, drought, heat, poor nutrition, and <a href="/wiki/Metal_toxicity" title="Metal toxicity">metal toxicity</a>.<a href="#cite_note-91">[91]</a> Individual AMF species have different effects in different hosts – introducing the AMF of one plant to another plant can reduce the latter's growth.<a href="#cite_note-92">[92]</a> <div class="mw-heading mw-heading5"><h5 id="Endophytic_fungi">Endophytic fungi</h5>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=19" title="Edit section: Endophytic fungi">edit</a>]</div> Endophytic fungi in <a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">mutualistic</a> relations directly benefit and benefit from their host plants. They also can help their hosts succeed in polluted environments such as those contaminated with toxic metals.<a href="#cite_note-93">[93]</a> Fungal <a href="/wiki/Endophyte" title="Endophyte">endophytes</a> are taxonomically diverse and are divided into categories based on mode of transmission, <a href="/wiki/Biodiversity" title="Biodiversity">biodiversity</a>, in planta colonization and host plant type.<a href="#cite_note-Rodriguez-2009-94">[94]</a><a href="#cite_note-95">[95]</a> Clavicipitaceous fungi systematically colonize temperate season grasses. Non-clavicipitaceous fungi colonize higher plants and even roots and divide into subcategories.<a href="#cite_note-96">[96]</a> <a href="/wiki/Aureobasidium" title="Aureobasidium">Aureobasidium</a> and <a href="/wiki/Preussia_(fungus)" title="Preussia (fungus)">preussia</a> species of endophytic fungi isolated from <a href="/wiki/Boswellia_sacra" title="Boswellia sacra">Boswellia sacra</a> produce <a href="/wiki/Indole-3-acetic_acid" title="Indole-3-acetic acid">indole acetic acid</a> <a href="/wiki/Hormone" title="Hormone">hormone</a> to promote plant health and development.<a href="#cite_note-97">[97]</a> <a href="/wiki/Aphid" title="Aphid">Aphids</a> can be found in most plants. Carnivorous <a href="/wiki/Coccinellidae" title="Coccinellidae">ladybirds</a> are aphid predators and are used in <a href="/wiki/Pest_control" title="Pest control">pest control</a>. Plant endophytic fungus <a href="/wiki/Neotyphodium_lolii" class="mw-redirect" title="Neotyphodium lolii">Neotyphodium lolii</a> produces <a href="/wiki/Alkaloid" title="Alkaloid">alkaloid</a> <a href="/wiki/Mycotoxin" title="Mycotoxin">mycotoxins</a> in response to <a href="/wiki/Aphid" title="Aphid">aphid</a> invasions. In response, ladybird predators exhibited reduced <a href="/wiki/Fertility" title="Fertility">fertility</a> and abnormal reproduction, suggesting that the mycotoxins are transmitted along the <a href="/wiki/Food_chain" title="Food chain">food chain</a> and affect the <a href="/wiki/Predation" title="Predation">predators</a>.<a href="#cite_note-Fungal_plant_endosymbionts_alter_li-82">[82]</a> <div class="mw-heading mw-heading4"><h4 id="Endophytic_bacteria">Endophytic bacteria</h4>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=20" title="Edit section: Endophytic bacteria">edit</a>]</div> Endophytic bacteria belong to a diverse group of plant endosymbionts characterized by systematic colonization of plant tissues. The most common genera include <a href="/wiki/Pseudomonas" title="Pseudomonas">Pseudomonas</a>, <a href="/wiki/Bacillus" title="Bacillus">Bacillus</a>, <a href="/wiki/Acinetobacter" title="Acinetobacter">Acinetobacter</a>, <a href="/wiki/Actinobacteria" class="mw-redirect" title="Actinobacteria">Actinobacteria</a>, <a href="/wiki/Sphingomonas" title="Sphingomonas">Sphingomonas</a>. Some endophytic bacteria, such as <a href="/wiki/Bacillus_amyloliquefaciens" title="Bacillus amyloliquefaciens">Bacillus amyloliquefaciens</a>, a seed-born endophytic bacteria, produce plant growth by producing <a href="/wiki/Gibberellins" class="mw-redirect" title="Gibberellins">gibberellins</a>, which are potent plant growth hormones. <a href="/wiki/Bacillus_amyloliquefaciens" title="Bacillus amyloliquefaciens">Bacillus amyloliquefaciens</a> promotes the taller height of <a href="/wiki/Transgenic_rice" class="mw-redirect" title="Transgenic rice">transgenic</a> dwarf rice plants.<a href="#cite_note-98">[98]</a> Some endophytic bacteria genera additionally belong to the <a href="/wiki/Enterobacteriaceae" title="Enterobacteriaceae">Enterobacteriaceae</a> family.<a href="#cite_note-99">[99]</a> Endophytic bacteria typically colonize the leaf tissues from plant roots, but can also enter the plant through the leaves through leaf <a href="/wiki/Stomata" class="mw-redirect" title="Stomata">stomata</a>.<a href="#cite_note-100">[100]</a> Generally, the endophytic bacteria are isolated from the plant tissues by surface <a href="/wiki/Sterilization_(microbiology)" title="Sterilization (microbiology)">sterilization</a> of the plant tissue in a sterile environment.<a href="#cite_note-101">[101]</a> Passenger endophytic bacteria eventually colonize inner tissue of plant by <a href="/wiki/Stochastic" title="Stochastic">stochastic</a> events while True endophytes possess adaptive traits because of which they live strictly in association with plants.<a href="#cite_note-102">[102]</a> The in vitro-cultivated endophytic <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> association with plants is considered a more intimate relationship that helps plants acclimatize to conditions and promotes health and growth. Endophytic bacteria are considered to be plant's essential endosymbionts because virtually all plants harbor them, and these endosymbionts play essential roles in host survival.<a href="#cite_note-103">[103]</a> This endosymbiotic relation is important in terms of <a href="/wiki/Ecology" title="Ecology">ecology</a>, <a href="/wiki/Evolution" title="Evolution">evolution</a> and diversity. Endophytic bacteria such as <a href="/wiki/Sphingomonas" title="Sphingomonas">Sphingomonas</a> sp. and <a href="/wiki/Serratia" title="Serratia">Serratia</a> sp. that are isolated from arid land plants regulate endogenous <a href="/wiki/Hormone" title="Hormone">hormone</a> content and promote growth.<a href="#cite_note-104">[104]</a> <div class="mw-heading mw-heading4"><h4 id="Archaea_endosymbionts">Archaea endosymbionts</h4>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=21" title="Edit section: Archaea endosymbionts">edit</a>]</div> <a href="/wiki/Archaea" title="Archaea">Archaea</a> are members of most <a href="/wiki/Microbiome" title="Microbiome">microbiomes</a>. While archaea are abundant in extreme environments, they are less abundant and diverse in association with eukaryotic hosts. Nevertheless, archaea are a substantial constituent of plant-associated ecosystems in the aboveground and belowground phytobiome, and play a role in host plant's health, growth and survival amid biotic and abiotic stresses. However, few studies have investigated the role of archaea in plant health and its symbiotic relationships.<a href="#cite_note-Jung-2020-105">[105]</a> Most plant endosymbiosis studies focus on fungal or bacteria using <a href="/wiki/Metagenomic" class="mw-redirect" title="Metagenomic">metagenomic</a> approaches.<a href="#cite_note-106">[106]</a> The characterization of archaea includes crop plants such as <a href="/wiki/Rice" title="Rice">rice</a><a href="#cite_note-107">[107]</a> and <a href="/wiki/Maize" title="Maize">maize</a>, but also aquatic plants.<a href="#cite_note-Jung-2020-105">[105]</a> The abundance of archaea varies by tissue type; for example archaea are more abundant in the <a href="/wiki/Rhizosphere" title="Rhizosphere">rhizosphere</a> than the <a href="/wiki/Phyllosphere" title="Phyllosphere">phyllosphere</a> and <a href="/wiki/Endosphere" title="Endosphere">endosphere</a>.<a href="#cite_note-108">[108]</a> This archaeal abundance is associated with plant species type, environment and the plant's developmental stage.<a href="#cite_note-109">[109]</a> In a study on plant <a href="/wiki/Genotype" title="Genotype">genotype</a>-specific archaeal and bacterial endophytes, 35% of archaeal sequences were detected in overall sequences (achieved using <a href="/wiki/Amplicon_sequencing" class="mw-redirect" title="Amplicon sequencing">amplicon sequencing</a> and verified by <a href="/wiki/Real-time_polymerase_chain_reaction" title="Real-time polymerase chain reaction">real time-PCR</a>). The archaeal sequences belong to the phyla <a href="/wiki/Thaumarchaeota" class="mw-redirect" title="Thaumarchaeota">Thaumarchaeota</a>, <a href="/wiki/Crenarchaeota" class="mw-redirect" title="Crenarchaeota">Crenarchaeota</a>, and <a href="/wiki/Euryarchaeota" title="Euryarchaeota">Euryarchaeota</a>.<a href="#cite_note-110">[110]</a> <div class="mw-heading mw-heading3"><h3 id="Bacteria">Bacteria</h3>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=22" title="Edit section: Bacteria">edit</a>]</div> Some <a href="/wiki/Betaproteobacteria" title="Betaproteobacteria">Betaproteobacteria</a> have <a href="/wiki/Gammaproteobacteria" title="Gammaproteobacteria">Gammaproteobacteria</a> endosymbionts.<a href="#cite_note-111">[111]</a> <div class="mw-heading mw-heading3"><h3 id="Fungi">Fungi</h3>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=23" title="Edit section: Fungi">edit</a>]</div> Fungi host endohyphal bacteria;<a href="#cite_note-Shaffer-2022-112">[112]</a> the effects of the bacteria are not well studied. Many such fungi in turn live within plants.<a href="#cite_note-Shaffer-2022-112">[112]</a> These fungi are otherwise known as fungal <a href="/wiki/Endophyte" title="Endophyte">endophytes</a>. It is hypothesized that the fungi offers a safe haven for the <a href="/wiki/Bacteria" title="Bacteria">bacteria</a>, and the diverse bacteria that they attract create a micro-ecosystem.<a href="#cite_note-113">[113]</a> These interactions may impact the way that fungi interact with the environment by modulating their <a href="/wiki/Phenotypes" class="mw-redirect" title="Phenotypes">phenotypes</a>.<a href="#cite_note-Shaffer-2022-112">[112]</a> The bacteria do this by altering the fungi's <a href="/wiki/Gene_expression" title="Gene expression">gene expression</a>.<a href="#cite_note-Shaffer-2022-112">[112]</a> For example, <a href="/wiki/Luteibacter" title="Luteibacter">Luteibacter</a> sp. has been shown to naturally infect the <a href="/wiki/Ascomycota" title="Ascomycota">ascomycetous</a> <a href="/wiki/Endophyte" title="Endophyte">endophyte</a> <a href="/wiki/Pestalotiopsis" title="Pestalotiopsis">Pestalotiopsis</a> sp. isolated from <a href="/wiki/Platycladus" title="Platycladus">Platycladus orientalis</a>.<a href="#cite_note-Shaffer-2022-112">[112]</a> The Luteibacter sp. influences the <a href="/wiki/Auxin" title="Auxin">auxin</a> and enzyme production within its host, which, in turn, may influence the effect the fungus has on its plant host.<a href="#cite_note-Shaffer-2022-112">[112]</a> Another interesting example of a bacterium living in symbiosis with a fungus is the fungus <a href="/wiki/Mortierella" title="Mortierella">Mortierella</a>. This soil-dwelling fungus lives in close association with a toxin-producing bacteria, Mycoavidus, which helps the fungus defend against <a href="/wiki/Nematode" title="Nematode">nematodes</a>.<a href="#cite_note-114">[114]</a> In 2024, researchers injected individual cells of the bacterium Mycetohabitans rhizoxinica into cells of the fungus <a href="/wiki/Rhizopus_microsporus" title="Rhizopus microsporus">Rhizopus microsporus</a> and were able to propagate the pair of cells for ten rounds using <a href="/wiki/Cell_sorting#Fluorescence-activated" title="Cell sorting">fluorescence-activated cell sorting</a> to select fungal cells containing the bacterium. They found that the fungus's DNA changed during the rounds of propagation.<a href="#cite_note-115">[115]</a> This was claimed to be the first time that endosymbiosis was artifically induced in a laboratory.<a href="#cite_note-116">[116]</a> <div class="mw-heading mw-heading2"><h2 id="Virus_endosymbionts">Virus endosymbionts</h2>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=24" title="Edit section: Virus endosymbionts">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Endogenous_retrovirus" title="Endogenous retrovirus">Endogenous retrovirus</a></div> The <a href="/wiki/Human_genome_project" class="mw-redirect" title="Human genome project">human genome project</a> found several thousand <a href="/wiki/Endogenous_retrovirus" title="Endogenous retrovirus">endogenous retroviruses</a>, <a href="/wiki/Endogenous_viral_element" title="Endogenous viral element">endogenous viral elements</a> in the <a href="/wiki/Genome" title="Genome">genome</a> that closely resemble and can be derived from <a href="/wiki/Retrovirus" title="Retrovirus">retroviruses</a>, organized into 24 families.<a href="#cite_note-117">[117]</a><a href="#cite_note-118">[118]</a> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=25" title="Edit section: See also">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col" style="column-width: 35em;"> <ul><li><a href="/wiki/Anagenesis" title="Anagenesis">Anagenesis</a> – Gradual evolutionary change in a species without splitting</li> <li><a href="/wiki/Cyanobiont" title="Cyanobiont">Cyanobiont</a> – Symbiotic bacterium</li> <li><a href="/wiki/Ectosymbiosis" title="Ectosymbiosis">Ectosymbiosis</a> – Symbiosis in which the symbiont lives on the body surface of the host</li> <li><a href="/wiki/Endophyte" title="Endophyte">Endophyte</a> – Endosymbiotic bacterium or fungus</li> <li><a href="/wiki/Epibiont" title="Epibiont">Epibiont</a> – Organism that lives on surface of another living organism</li> <li><a href="/wiki/Fungal-bacterial_endosymbiosis" title="Fungal-bacterial endosymbiosis">Fungal-bacterial endosymbiosis</a></li> <li><a href="/wiki/List_of_symbiotic_organisms" class="mw-redirect" title="List of symbiotic organisms">List of symbiotic organisms</a> – Close, long-term biological interaction between distinct organisms (usually species)Pages displaying short descriptions of redirect targets</li> <li><a href="/wiki/List_of_symbiotic_relationships" class="mw-redirect" title="List of symbiotic relationships">List of symbiotic relationships</a> – Close, long-term biological interaction between distinct organisms (usually species)Pages displaying short descriptions of redirect targets</li> <li><a href="/wiki/Multigenomic_organism" class="mw-redirect" title="Multigenomic organism">Multigenomic organism</a> – Close, long-term biological interaction between distinct organisms (usually species)Pages displaying short descriptions of redirect targets</li> <li><a href="/wiki/Nitroplast" title="Nitroplast">Nitroplast</a> – Nitrogen-fixing organelle</li> <li><a href="/wiki/Protocell" title="Protocell">Protocell</a> – Lipid globule proposed as a precursor of living cells</li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2>[<a href="/w/index.php?title=Endosymbiont&action=edit&section=26" title="Edit section: References">edit</a>]</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 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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=Long-term+reinfection+of+the+human+genome+by+endogenous+retroviruses&rft.volume=101&rft.issue=14&rft.pages=%3Cspan+class%3D%22nowrap%22%3E4894-%3C%2Fspan%3E4899&rft.date=2004-04&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC387345%23id-name%3DPMC&rft_id=info%3Apmid%2F15044706&rft_id=info%3Adoi%2F10.1073%2Fpnas.0307800101&rft_id=info%3Abibcode%2F2004PNAS..101.4894B&rft.aulast=Belshaw&rft.aufirst=R&rft.au=Pereira%2C+V&rft.au=Katzourakis%2C+A&rft.au=Talbot%2C+G&rft.au=Paces%2C+J&rft.au=Burt%2C+A&rft.au=Tristem%2C+M&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC387345&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEndosymbiont" class="Z3988"> </li> </ol></div></div> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output 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class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Life" title="Life">Cellular life</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/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><a href="/wiki/Incertae_sedis" title="Incertae sedis">Incertae sedis</a> <ul><li><a href="/wiki/Parakaryon" title="Parakaryon">Parakaryon</a></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 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><a href="/wiki/Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a></li> <li><a href="/wiki/Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a></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 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> 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 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 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 class="mw-selflink selflink">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 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