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Available in 11 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-11" 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">11 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ceb mw-list-item"><a href="https://ceb.wikipedia.org/wiki/Euprymna_scolopes" title="Euprymna scolopes – Cebuano" lang="ceb" hreflang="ceb" data-title="Euprymna scolopes" data-language-autonym="Cebuano" data-language-local-name="Cebuano" class="interlanguage-link-target"><span>Cebuano</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Euprymna_scolopes" title="Euprymna scolopes – German" lang="de" hreflang="de" data-title="Euprymna scolopes" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Euprymna_scolopes" title="Euprymna scolopes – Spanish" lang="es" hreflang="es" data-title="Euprymna scolopes" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Euprymna_scolopes" title="Euprymna scolopes – Basque" lang="eu" hreflang="eu" data-title="Euprymna scolopes" 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/Euprymna_scolopes" title="Euprymna scolopes – French" lang="fr" hreflang="fr" data-title="Euprymna scolopes" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Euprymna_scolopes" title="Euprymna scolopes – Italian" lang="it" hreflang="it" data-title="Euprymna scolopes" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Euprymna_scolopes" title="Euprymna scolopes – Dutch" lang="nl" hreflang="nl" data-title="Euprymna scolopes" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link 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class="infobox biota" style="text-align: left; width: 200px; font-size: 100%"> <tbody><tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(235,235,210)"><i>Euprymna scolopes</i> </th></tr> <tr> <td colspan="2" style="text-align: center"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Euprymna_scolopes,_South_shore_of_Oahu,_Hawaii.tiff" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5c/Euprymna_scolopes%2C_South_shore_of_Oahu%2C_Hawaii.tiff/lossy-page1-220px-Euprymna_scolopes%2C_South_shore_of_Oahu%2C_Hawaii.tiff.jpg" decoding="async" width="220" height="169" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5c/Euprymna_scolopes%2C_South_shore_of_Oahu%2C_Hawaii.tiff/lossy-page1-330px-Euprymna_scolopes%2C_South_shore_of_Oahu%2C_Hawaii.tiff.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5c/Euprymna_scolopes%2C_South_shore_of_Oahu%2C_Hawaii.tiff/lossy-page1-440px-Euprymna_scolopes%2C_South_shore_of_Oahu%2C_Hawaii.tiff.jpg 2x" data-file-width="278" data-file-height="213" /></a></span> </td></tr> <tr> <td colspan="2" style="text-align: center; font-size: 88%">The Hawaiian bobtail squid, <i>Euprymna scolopes</i>, swimming in the water column off the south shore of Oahu </td></tr> <tr style="color:inherit; text-align: center; background-color: rgb(235,235,210)"> <th colspan="2"><div style="text-align: center"><a href="/wiki/Conservation_status" title="Conservation status">Conservation status</a></div> </th></tr> <tr> <td colspan="2"><div style="text-align: center"><span class="mw-default-size skin-invert-image" typeof="mw:File/Frameless"><span><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/4/4f/Status_iucn3.1_blank.svg/220px-Status_iucn3.1_blank.svg.png" decoding="async" width="220" height="59" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/4f/Status_iucn3.1_blank.svg/330px-Status_iucn3.1_blank.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/4f/Status_iucn3.1_blank.svg/440px-Status_iucn3.1_blank.svg.png 2x" data-file-width="512" data-file-height="137" /></span></span><br /><a href="/wiki/Data_Deficient" class="mw-redirect" title="Data Deficient">Data Deficient</a> <small> (<a href="/wiki/IUCN_Red_List" title="IUCN Red List">IUCN 3.1</a>)<sup id="cite_ref-IUCN_1-0" class="reference"><a href="#cite_note-IUCN-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup></small></div> </td></tr> <tr> <th colspan="2" style="color:inherit; min-width:15em; text-align: center; background-color: rgb(235,235,210)"><a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">Scientific classification</a> <span class="plainlinks taxobox-edit-taxonomy skin-invert" style="font-size:smaller; float:right; padding-right:0.4em; margin-left:-3em;"><span typeof="mw:File"><a href="/wiki/Template:Taxonomy/Euprymna" title="Edit this classification"><img alt="Edit this classification" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/15px-OOjs_UI_icon_edit-ltr.svg.png" decoding="async" width="15" height="15" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/23px-OOjs_UI_icon_edit-ltr.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/30px-OOjs_UI_icon_edit-ltr.svg.png 2x" data-file-width="20" data-file-height="20" /></a></span></span> </th></tr> <tr> <td>Domain: </td> <td><a href="/wiki/Eukaryote" title="Eukaryote">Eukaryota</a> </td></tr> <tr> <td>Kingdom: </td> <td><a href="/wiki/Animal" title="Animal">Animalia</a> </td></tr> <tr> <td>Phylum: </td> <td><a href="/wiki/Mollusca" title="Mollusca">Mollusca</a> </td></tr> <tr> <td>Class: </td> <td><a href="/wiki/Cephalopod" title="Cephalopod">Cephalopoda</a> </td></tr> <tr> <td>Order: </td> <td><a href="/wiki/Bobtail_squid" title="Bobtail squid">Sepiolida</a> </td></tr> <tr> <td>Family: </td> <td><a href="/wiki/Sepiolidae" title="Sepiolidae">Sepiolidae</a> </td></tr> <tr> <td>Subfamily: </td> <td><a href="/wiki/Sepiolinae" title="Sepiolinae">Sepiolinae</a> </td></tr> <tr> <td>Genus: </td> <td><a href="/wiki/Euprymna" title="Euprymna"><i>Euprymna</i></a> </td></tr> <tr> <td>Species: </td> <td><div style="display:inline" class="species"><i><b>E. scolopes</b></i></div> </td></tr> <tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(235,235,210)"><a href="/wiki/Binomial_nomenclature" title="Binomial nomenclature">Binomial name</a> </th></tr> <tr> <td colspan="2" style="text-align: center"><b><span class="binomial"><span style="font-weight:normal;"></span><i>Euprymna scolopes</i></span></b><br /><div style="font-size: 85%;"><a href="/wiki/S._Stillman_Berry" title="S. Stillman Berry">Berry</a>, 1913<sup id="cite_ref-WoRMS_2-0" class="reference"><a href="#cite_note-WoRMS-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup></div> </td></tr> </tbody></table><style data-mw-deduplicate="TemplateStyles:r1238732961">@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox.biota tr{background:transparent!important}html.skin-theme-clientpref-night .mw-parser-output .infobox.biota img{background:transparent}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox.biota tr{background:transparent!important}html.skin-theme-clientpref-os .mw-parser-output .infobox.biota img{background:white}}.mw-parser-output .infobox.biota .taxobox-edit-taxonomy img{background:transparent!important}body.skin-vector .mw-parser-output table.biota.infobox{margin-top:0.5em}body.skin--responsive .mw-parser-output table.biota.infobox tr.taxonrow td{padding:2px 10px}</style> <p><i><b>Euprymna scolopes</b></i>, also known as the <b>Hawaiian bobtail squid</b>, is a <a href="/wiki/Species" title="Species">species</a> of <a href="/wiki/Bobtail_squid" title="Bobtail squid">bobtail squid</a> in the <a href="/wiki/Family_(biology)" title="Family (biology)">family</a> <a href="/wiki/Sepiolidae" title="Sepiolidae">Sepiolidae</a> native to the central <a href="/wiki/Pacific_Ocean" title="Pacific Ocean">Pacific Ocean</a>, where it occurs in shallow coastal waters off the <a href="/wiki/Hawaiian_Islands" title="Hawaiian Islands">Hawaiian Islands</a> and <a href="/wiki/Midway_Atoll" title="Midway Atoll">Midway Island</a>.<sup id="cite_ref-Reid_3-0" class="reference"><a href="#cite_note-Reid-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> The type specimen was collected off the Hawaiian Islands and is located at the <a href="/wiki/National_Museum_of_Natural_History" title="National Museum of Natural History">National Museum of Natural History</a> in <a href="/wiki/Washington,_D.C." title="Washington, D.C.">Washington, D.C.</a><sup id="cite_ref-Recent_5-0" class="reference"><a href="#cite_note-Recent-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p><i>Euprymna scolopes</i> grows to 30 mm (1.2 in) in <a href="/wiki/Mantle_(mollusc)" title="Mantle (mollusc)">mantle</a> length.<sup id="cite_ref-Reid_3-1" class="reference"><a href="#cite_note-Reid-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Hatchlings weigh 0.005 g (0.00018 oz) and mature in 80 days. Adults weigh up to 2.67 g (0.094 oz).<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> </p><p>In the wild, <i>E. scolopes</i> feeds on species of <a href="/wiki/Shrimp" title="Shrimp">shrimp</a>, including <i><a href="/wiki/Halocaridina_rubra" title="Halocaridina rubra">Halocaridina rubra</a></i>, <i><a href="/wiki/Palaemon_debilis" title="Palaemon debilis">Palaemon debilis</a></i>, and <i><a href="/wiki/Palaemon_pacificus" title="Palaemon pacificus">Palaemon pacificus</a></i>.<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> In the laboratory, <i>E. scolopes</i> has been reared on a varied diet of animals, including <a href="/wiki/Mysid" class="mw-redirect" title="Mysid">mysids</a> (<i><a href="/wiki/Anisomysis" title="Anisomysis">Anisomysis</a></i> sp.), <a href="/wiki/Brine_shrimp" title="Brine shrimp">brine shrimp</a> (<i><a href="/wiki/Artemia_salina" title="Artemia salina">Artemia salina</a></i>), <a href="/wiki/Mosquitofish" title="Mosquitofish">mosquitofish</a> (<i><a href="/wiki/Gambusia_affinis" class="mw-redirect" title="Gambusia affinis">Gambusia affinis</a></i>), <a href="/wiki/Prawn" title="Prawn">prawns</a> (<i><a href="/w/index.php?title=Leander_debilis&action=edit&redlink=1" class="new" title="Leander debilis (page does not exist)">Leander debilis</a></i>), and <a href="/wiki/Octopus" title="Octopus">octopuses</a> (<i><a href="/wiki/Octopus_cyanea" title="Octopus cyanea">Octopus cyanea</a></i>).<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> </p><p>The <a href="/wiki/Hawaiian_monk_seal" title="Hawaiian monk seal">Hawaiian monk seal</a> (<i>Monachus schauinslandi</i>) preys on <i>E. scolopes</i> in northwestern Hawaiian waters.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p><p>On June 3, 2021, <a href="/wiki/SpaceX_CRS-22" title="SpaceX CRS-22">SpaceX CRS-22</a> launched <i>E. scolopes,</i> along with <a href="/wiki/Tardigrade" title="Tardigrade">tardigrades</a>, to the <a href="/wiki/International_Space_Station" title="International Space Station">International Space Station</a>. The squid were launched as hatchlings and will be studied to see if they can incorporate their symbiotic bacteria into their light organ while in space.<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> </p> <div class="mw-heading mw-heading2"><h2 id="Symbiosis">Symbiosis</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Euprymna_scolopes&action=edit&section=1" title="Edit section: Symbiosis"><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">Further information: <a href="/wiki/Counter-illumination" title="Counter-illumination">Counter-illumination</a></div> <p><i>Euprymna scolopes</i> lives in a <a href="/wiki/Symbiosis" title="Symbiosis">symbiotic</a> relationship with the <a href="/wiki/Bioluminescent" class="mw-redirect" title="Bioluminescent">bioluminescent</a> <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> <i><a href="/wiki/Aliivibrio_fischeri" title="Aliivibrio fischeri">Aliivibrio fischeri</a></i>, which inhabits a special light organ in the squid's mantle. To allows this symbiotic relationship, <i>Crumbs protein</i> must first induce <i>Apopstosis</i>, which kills superficial epithelial tissue found in <i>Euprymna scolopes.</i> Apopstosis then helps create <i>crypt</i> epithelial cells, these cells directly take in the bioluminescence bacteria received from the <i>Aliivibrio fischeri</i>.<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> The bacteria are fed a <a href="/wiki/Sugar" title="Sugar">sugar</a> and <a href="/wiki/Amino_acid" title="Amino acid">amino acid</a> solution by the squid and in return hide the squid's silhouette when viewed from below by matching the amount of light hitting the top of the mantle (<a href="/wiki/Counter-illumination" title="Counter-illumination">counter-illumination</a>).<sup id="cite_ref-Young_12-0" class="reference"><a href="#cite_note-Young-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> <i>E. scolopes</i> serves as a <a href="/wiki/Model_organism" title="Model organism">model organism</a> for animal-bacterial symbiosis and its relationship with <i>A. fischeri</i> has been carefully studied.<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><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><sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Acquisition">Acquisition</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Euprymna_scolopes&action=edit&section=2" title="Edit section: Acquisition"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The bioluminescent bacterium, <i>A. fischeri</i>, is horizontally transmitted throughout the <i>E. scolopes</i> population. Hatchlings lack these necessary bacteria and must carefully select for them in a marine world saturated with other microorganisms.<sup id="cite_ref-Chun_21-0" class="reference"><a href="#cite_note-Chun-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p><p>To effectively capture these cells, <i>E. scolopes</i> secretes <a href="/wiki/Mucus" title="Mucus">mucus</a> in response to <a href="/wiki/Peptidoglycan" title="Peptidoglycan">peptidoglycan</a> (a major <a href="/wiki/Cell_wall" title="Cell wall">cell wall</a> component of <a href="/wiki/Bacteria" title="Bacteria">bacteria</a>).<sup id="cite_ref-Nyholm_22-0" class="reference"><a href="#cite_note-Nyholm-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> The mucus inundates the <a href="/wiki/Cilia" class="mw-redirect" title="Cilia">ciliated</a> fields in the immediate area around the six pores of the light organ and captures a large variety of bacteria. However, by some unknown mechanism, <i>A. fischeri</i> is able to outcompete other bacteria in the mucus.<sup id="cite_ref-Nyholm_22-1" class="reference"><a href="#cite_note-Nyholm-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Hawaiian_Bobtail_squid.tiff" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f9/Hawaiian_Bobtail_squid.tiff/lossless-page1-300px-Hawaiian_Bobtail_squid.tiff.png" decoding="async" width="300" height="128" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f9/Hawaiian_Bobtail_squid.tiff/lossless-page1-450px-Hawaiian_Bobtail_squid.tiff.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f9/Hawaiian_Bobtail_squid.tiff/lossless-page1-600px-Hawaiian_Bobtail_squid.tiff.png 2x" data-file-width="2222" data-file-height="951" /></a><figcaption>Adult <i>Euprymna scolopes</i> with scale.</figcaption></figure> <p>As <i>A. fischeri</i> cells aggregate in the mucus, they must use their <a href="/wiki/Flagella" class="mw-redirect" title="Flagella">flagella</a> to migrate through the <a href="https://en.wiktionary.org/wiki/pore" class="extiw" title="wiktionary:pore">pores</a> and down into the ciliated ducts of the light organ and endure another barrage of host factors meant to ensure only <i>A. fischeri</i> colonization.<sup id="cite_ref-Nyholm_22-2" class="reference"><a href="#cite_note-Nyholm-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> Besides the relentless host-derived current that forces <a href="/wiki/Motility" title="Motility">motility</a>-challenged bacteria out of the pores, a number of reactive oxygen species makes the environment unbearable.<sup id="cite_ref-Nyholm_22-3" class="reference"><a href="#cite_note-Nyholm-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> Squid <a href="/w/index.php?title=Halide_peroxidase&action=edit&redlink=1" class="new" title="Halide peroxidase (page does not exist)">halide peroxidase</a> is the main enzyme responsible for crafting this <a href="/wiki/Microbicide" class="mw-redirect" title="Microbicide">microbiocidal</a> environment, using <a href="/wiki/Hydrogen_peroxide" title="Hydrogen peroxide">hydrogen peroxide</a> as a substrate, but <i>A. fischeri</i> has evolved a brilliant counterattack. <i>A. fischeri</i> possesses a periplasmic <a href="/wiki/Catalyst" class="mw-redirect" title="Catalyst">catalase</a> that captures hydrogen peroxide before it can be used by the squid halide peroxidase, thus inhibiting the enzyme indirectly.<sup id="cite_ref-Nyholm_22-4" class="reference"><a href="#cite_note-Nyholm-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> Once through these ciliated ducts, <i>A. fischeri</i> cells swim on towards the antechamber, a large <a href="/wiki/Epithelium" title="Epithelium">epithelial</a>-lined space, and colonize the narrow epithelial crypts.<sup id="cite_ref-Nyholm_22-5" class="reference"><a href="#cite_note-Nyholm-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p><p>The bacteria thrive on the host-derived amino acids and sugars in the <a href="/wiki/Antechamber" class="mw-redirect" title="Antechamber">antechamber</a> and quickly fill the crypt spaces within 10 to 12 hours after hatching.<sup id="cite_ref-Visick_23-0" class="reference"><a href="#cite_note-Visick-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Ongoing_relationship">Ongoing relationship</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Euprymna_scolopes&action=edit&section=3" title="Edit section: Ongoing relationship"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Every second, a juvenile squid ventilates about 2.6 ml (0.092 imp fl oz; 0.088 US fl oz) of ambient <a href="/wiki/Seawater" title="Seawater">seawater</a> through its mantle cavity. Only a single <i>A. fischeri</i> cell, one/1-millionth of the total volume, is present with each ventilation.<sup id="cite_ref-Nyholm_22-6" class="reference"><a href="#cite_note-Nyholm-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p><p>The increased amino acids and sugars feed the <a href="/wiki/Metabolism" title="Metabolism">metabolically</a> demanding <a href="/wiki/Bioluminescence" title="Bioluminescence">bioluminescence</a> of the <i>A. fischeri</i>, and in 12 hours, the bioluminescence peaks and the juvenile squid is able to counterilluminate less than a day after hatching.<sup id="cite_ref-Visick_23-1" class="reference"><a href="#cite_note-Visick-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> Bioluminescence demands a substantial amount of energy from a bacterial cell. It is estimated to demand 20% of a cell's metabolic potential.<sup id="cite_ref-Visick_23-2" class="reference"><a href="#cite_note-Visick-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p><p>Nonluminescent strains of <i>A. fischeri</i> would have a definite competitive advantage over the luminescent wild-type, however nonluminescent mutants are never found in the light organ of the <i>E. scolopes</i>.<sup id="cite_ref-Visick_23-3" class="reference"><a href="#cite_note-Visick-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> In fact, experimental procedures have shown that removing the <a href="/wiki/Gene" title="Gene">genes</a> responsible for light production in <i>A. fischeri</i> drastically reduces colonization efficiency.<sup id="cite_ref-Visick_23-4" class="reference"><a href="#cite_note-Visick-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> Luminescent cells, with functioning <a href="/wiki/Luciferase" title="Luciferase">luciferase</a>, may have a higher affinity for <a href="/wiki/Oxygen" title="Oxygen">oxygen</a> than for <a href="/wiki/Peroxidases" class="mw-redirect" title="Peroxidases">peroxidases</a>, thereby negating the toxic effects of the peroxidases.<sup id="cite_ref-Timmins_24-0" class="reference"><a href="#cite_note-Timmins-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> For this reason, bioluminescence is thought to have evolved as an ancient oxygen detoxification mechanism in bacteria.<sup id="cite_ref-Timmins_24-1" class="reference"><a href="#cite_note-Timmins-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Venting">Venting</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Euprymna_scolopes&action=edit&section=4" title="Edit section: Venting"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Despite all the effort that goes into obtaining luminescent <i>A. fischeri</i>, the host squid jettisons most of the cells daily. This process, known as "venting", is responsible for the disposal of up to 95% of <i>A. fischeri</i> in the light organ every morning at dawn.<sup id="cite_ref-Schuster_25-0" class="reference"><a href="#cite_note-Schuster-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> The bacteria gain no benefit from this behavior and the upside for the squid itself is not clearly understood. One reasonable explanation points to the large energy expenditure in maintaining a colony of bioluminescent bacteria.<sup id="cite_ref-Jones_26-0" class="reference"><a href="#cite_note-Jones-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p><p>During the day when the squid are inactive and hidden, bioluminescence is unnecessary, and expelling the <i>A. fischeri</i> conserves energy. Another, more evolutionarily important reason may be that daily venting ensures selection for <i>A. fischeri</i> that have evolved specificity for a particular host, but can survive outside of the light organ.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> </p><p>Since <i>A. fischeri</i> is transmitted horizontally in <i>E. scolopes</i>, maintaining a stable population of them in the open ocean is essential in supplying future generations of squid with functioning light organs. </p> <div class="mw-heading mw-heading2"><h2 id="Light_organ">Light organ</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Euprymna_scolopes&action=edit&section=5" title="Edit section: Light organ"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The light organ has an electrical response when stimulated by light, which suggests the organ functions as a <a href="/wiki/Simple_eyes_in_invertebrates" class="mw-redirect" title="Simple eyes in invertebrates">photoreceptor</a> that enables the host squid to respond to <i>A. fischeri'</i>s luminescence.<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p><p>Extraocular <a href="/wiki/Vesicle_(biology)" class="mw-redirect" title="Vesicle (biology)">vesicles</a> collaborate with the eyes to monitor the down-welling light and light created from counterillumination, so as the squid moves to various depths, it can maintain the proper level of output light.<sup id="cite_ref-Jones_26-1" class="reference"><a href="#cite_note-Jones-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> Acting on this information, the squid can then adjust the intensity of the <a href="/wiki/Bioluminescence" title="Bioluminescence">bioluminescence</a> by modifying the <a href="/wiki/Ink_sac" title="Ink sac">ink sac</a>, which functions as a diaphragm around the light organ.<sup id="cite_ref-Jones_26-2" class="reference"><a href="#cite_note-Jones-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> Furthermore, the light organ contains a network of unique reflector and lens tissues that help reflect and focus the light ventrally through the <a href="/wiki/Mantle_(mollusc)" title="Mantle (mollusc)">mantle</a>.<sup id="cite_ref-Jones_26-3" class="reference"><a href="#cite_note-Jones-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p><p>The light organ of <a href="/wiki/Embryo" title="Embryo">embryonic</a> and juvenile squids has a striking anatomical similarity to an eye and <a href="/wiki/Gene_expression" title="Gene expression">expresses</a> several genes similar to those involved in eye development in mammalian embryos (e.g. <i><a href="/wiki/EYA1" class="mw-redirect" title="EYA1">eya</a></i>, <i><a href="/wiki/DACH1" title="DACH1">dac</a></i>) which indicate that squid eyes and squid light organs may be formed using the same <a href="/wiki/Developmental_biology" title="Developmental biology">developmental</a> "toolkit".<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> </p><p>As the down-welling light increases or decreases, the squid is able to adjust luminescence accordingly, even over multiple cycles of light intensity.<sup id="cite_ref-Jones_26-4" class="reference"><a href="#cite_note-Jones-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Euprymna_scolopes&action=edit&section=6" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Reflectin" title="Reflectin">Reflectin</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=Euprymna_scolopes&action=edit&section=7" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width reflist-columns-2"> <ol class="references"> <li id="cite_note-IUCN-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-IUCN_1-0">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFBarratt,_I.Allcock,_L.2012" class="citation journal cs1">Barratt, I.; <a href="/wiki/Louise_Allcock" title="Louise Allcock">Allcock, L.</a> (2012). <a rel="nofollow" class="external text" href="https://doi.org/10.2305%2FIUCN.UK.2012-1.RLTS.T162598A925206.en">"<i>Euprymna scolopes</i>"</a>. <i><a href="/wiki/The_IUCN_Red_List_of_Threatened_Species" class="mw-redirect" title="The IUCN Red List of Threatened Species">The IUCN Red List of Threatened Species</a></i>. <b>2012</b>: e.T162598A925206. <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/10.2305%2FIUCN.UK.2012-1.RLTS.T162598A925206.en">10.2305/IUCN.UK.2012-1.RLTS.T162598A925206.en</a></span>.</cite><span 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Roper 1976. Bioluminescent countershading in midwater animals: evidence from living squid. <i>Science</i> <b>191</b>(4231): 1046–1048. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscience.1251214">10.1126/science.1251214</a></span> </li> <li id="cite_note-13"><span class="mw-cite-backlink"><b><a href="#cite_ref-13">^</a></b></span> <span class="reference-text">DeLoney, C.R., T.M. Bartley & K.L. Visick 2002. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20040328114942/http://www.cephbase.utmb.edu/refdb/pdf/7468.pdf">"Role for phosphoglucomutase in <i>Aliivibrio fischeri</i>-<i>Euprymna scolopes</i> symbiosis"</a> <span class="cs1-format">(PDF)</span>. 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Archived from <a rel="nofollow" class="external text" href="http://www.cephbase.utmb.edu/refdb/pdf/7360.pdf">the original</a> <span class="cs1-format">(PDF)</span> on March 28, 2004.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Aliivibrio+fischeri+lipopolysaccharide+induces+developmental+apoptosis%2C+but+not+complete+morphogenesis%2C+of+the+Euprymna+scolopes+light+organ.&rft_id=http%3A%2F%2Fwww.cephbase.utmb.edu%2Frefdb%2Fpdf%2F7360.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEuprymna+scolopes" class="Z3988"></span> <span style="font-size:85%;">(610 KB)</span> <i>Developmental Biology</i> <b>226</b>(2): 242-254.</span> </li> <li id="cite_note-16"><span class="mw-cite-backlink"><b><a href="#cite_ref-16">^</a></b></span> <span class="reference-text">Hanlon, R.T., M.F. Claes, S.E. Ashcraft & P.V. Dunlap 1997. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20040328183009/http://www.cephbase.utmb.edu/refdb/pdf/7724.pdf">"Laboratory culture of the sepiolid squid <i>Euprymna scolopes</i>: A model system for bacteria-animal symbiosis"</a> <span class="cs1-format">(PDF)</span>. Archived from <a rel="nofollow" class="external text" href="http://www.cephbase.utmb.edu/refdb/pdf/7724.pdf">the original</a> <span class="cs1-format">(PDF)</span> on March 28, 2004.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Laboratory+culture+of+the+sepiolid+squid+Euprymna+scolopes%3A+A+model+system+for+bacteria-animal+symbiosis.&rft_id=http%3A%2F%2Fwww.cephbase.utmb.edu%2Frefdb%2Fpdf%2F7724.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEuprymna+scolopes" class="Z3988"></span> <span style="font-size:85%;">(2.38 MB)</span> <i>Biological Bulletin</i> <b>192</b>(3): 364-374.</span> </li> <li id="cite_note-17"><span class="mw-cite-backlink"><b><a href="#cite_ref-17">^</a></b></span> <span class="reference-text">Lee, K.-H. & E.G. Ruby 1995. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20040328145721/http://www.cephbase.utmb.edu/refdb/pdf/4338.pdf">"Symbiotic role of the viable but nonculturable state of <i>Aliivibrio fischeri</i> in Hawaiian coastal seawater"</a> <span class="cs1-format">(PDF)</span>. Archived from <a rel="nofollow" class="external text" href="http://www.cephbase.utmb.edu/refdb/pdf/4338.pdf">the original</a> <span class="cs1-format">(PDF)</span> on March 28, 2004.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Symbiotic+role+of+the+viable+but+nonculturable+state+of+Aliivibrio+fischeri+in+Hawaiian+coastal+seawater.&rft_id=http%3A%2F%2Fwww.cephbase.utmb.edu%2Frefdb%2Fpdf%2F4338.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEuprymna+scolopes" class="Z3988"></span> <span style="font-size:85%;">(249 KB)</span> <i>Applied and Environmental Microbiology</i> <b>61</b>(1): 278-283.</span> </li> <li id="cite_note-18"><span class="mw-cite-backlink"><b><a href="#cite_ref-18">^</a></b></span> <span class="reference-text">Lemus, J.D. & M.J. McFall-Ngai 2000. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20040328174755/http://www.cephbase.utmb.edu/refdb/pdf/7610.pdf">"Alterations in the protoeme of the <i>Euprymna scolopes</i> light organ in response to symbiotic <i>Aliivibrio fischeri</i>"</a> <span class="cs1-format">(PDF)</span>. Archived from <a rel="nofollow" class="external text" href="http://www.cephbase.utmb.edu/refdb/pdf/7610.pdf">the original</a> <span class="cs1-format">(PDF)</span> on March 28, 2004.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Alterations+in+the+protoeme+of+the+Euprymna+scolopes+light+organ+in+response+to+symbiotic+Aliivibrio+fischeri.&rft_id=http%3A%2F%2Fwww.cephbase.utmb.edu%2Frefdb%2Fpdf%2F7610.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEuprymna+scolopes" class="Z3988"></span> <span style="font-size:85%;">(2.10 MB)</span> <i>Applied and Environmental Microbiology</i> <b>66</b>: 4091-4097.</span> </li> <li id="cite_note-19"><span class="mw-cite-backlink"><b><a href="#cite_ref-19">^</a></b></span> <span class="reference-text">Millikan, D.S. & E.G. Ruby 2003. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20040328200403/http://www.cephbase.utmb.edu/refdb/pdf/7850.pdf">"FlrA, a s<sup>54</sup>-Dependent Transcriptional Activator in <i>Aliivibrio fischeri</i>, is required for motility and symbiotic light-organ colonization"</a> <span class="cs1-format">(PDF)</span>. Archived from <a rel="nofollow" class="external text" href="http://www.cephbase.utmb.edu/refdb/pdf/7850.pdf">the original</a> <span class="cs1-format">(PDF)</span> on March 28, 2004.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=FlrA%2C+a+s%3Csup%3E54%3C%2Fsup%3E-Dependent+Transcriptional+Activator+in+Aliivibrio+fischeri%2C+is+required+for+motility+and+symbiotic+light-organ+colonization.&rft_id=http%3A%2F%2Fwww.cephbase.utmb.edu%2Frefdb%2Fpdf%2F7850.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEuprymna+scolopes" class="Z3988"></span> <span style="font-size:85%;">(382 KB)</span> <i>Journal of Bacteriology</i> (American Society for Microbiology) <b>185</b>(12): 3547-3557.</span> </li> <li id="cite_note-20"><span class="mw-cite-backlink"><b><a href="#cite_ref-20">^</a></b></span> <span class="reference-text">Montgomery, M.K. & M. McFall-Ngai 1998. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20040328163337/http://www.cephbase.utmb.edu/refdb/pdf/5803.pdf">"Late postembryonic development of the symbiotic light organ of <i>Euprymna scolopes</i> (Cephalopoda: Sepiolidae)"</a> <span class="cs1-format">(PDF)</span>. Archived from <a rel="nofollow" class="external text" href="http://www.cephbase.utmb.edu/refdb/pdf/5803.pdf">the original</a> <span class="cs1-format">(PDF)</span> on March 28, 2004.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Late+postembryonic+development+of+the+symbiotic+light+organ+of+Euprymna+scolopes+%28Cephalopoda%3A+Sepiolidae%29.&rft_id=http%3A%2F%2Fwww.cephbase.utmb.edu%2Frefdb%2Fpdf%2F5803.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEuprymna+scolopes" class="Z3988"></span> <span style="font-size:85%;">(6.10 MB)</span> <i>Biological Bulletin</i> <b>195</b>: 326-336.</span> </li> <li id="cite_note-Chun-21"><span class="mw-cite-backlink"><b><a href="#cite_ref-Chun_21-0">^</a></b></span> <span class="reference-text">Effects of colonization, luminescence, and autoinducer on host transcription during development of the squid-vibrio association.<i><a href="/wiki/Proceedings_of_the_National_Academy_of_Sciences_of_the_United_States_of_America" title="Proceedings of the National Academy of Sciences of the United States of America">Proceedings of the National Academy of Sciences of the United States of America</a></i> <b>105</b>(32): 11323-11328. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.0802369105">10.1073/pnas.0802369105</a></span> </li> <li id="cite_note-Nyholm-22"><span class="mw-cite-backlink">^ <a href="#cite_ref-Nyholm_22-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Nyholm_22-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Nyholm_22-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Nyholm_22-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Nyholm_22-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-Nyholm_22-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-Nyholm_22-6"><sup><i><b>g</b></i></sup></a></span> <span class="reference-text">The evolutionary ecology of a sepiolid squid-<i>Aliivibrio</i> association: from cell to environment. <i><a href="/w/index.php?title=Vie_et_Milieu&action=edit&redlink=1" class="new" title="Vie et Milieu (page does not exist)">Vie et Milieu</a> <b>58</b>(2): 175-184. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://www.worldcat.org/search?fq=x0:jrnl&q=n2:0240-8759">0240-8759</a></i></span> </li> <li id="cite_note-Visick-23"><span class="mw-cite-backlink">^ <a href="#cite_ref-Visick_23-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Visick_23-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Visick_23-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Visick_23-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Visick_23-4"><sup><i><b>e</b></i></sup></a></span> <span class="reference-text">An exclusive contract: Specificity in the <i>Aliivibrio fischeri</i> <i>Euprymna scolopes</i> partnership. <i><a href="/wiki/Journal_of_Bacteriology" title="Journal of Bacteriology">Journal of Bacteriology</a></i> <b>182</b>(7): 1779-1787. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://www.worldcat.org/search?fq=x0:jrnl&q=n2:0021-9193">0021-9193</a></span> </li> <li id="cite_note-Timmins-24"><span class="mw-cite-backlink">^ <a href="#cite_ref-Timmins_24-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Timmins_24-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text">The evolution of bioluminescent oxygen consumption as an ancient oxygen detoxification mechanism.<i><a href="/wiki/Journal_of_Molecular_Evolution" title="Journal of Molecular Evolution">Journal of Molecular Evolution</a></i> <b>52</b>(4): 321-332. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://www.worldcat.org/search?fq=x0:jrnl&q=n2:0022-2844">0022-2844</a></span> </li> <li id="cite_note-Schuster-25"><span class="mw-cite-backlink"><b><a href="#cite_ref-Schuster_25-0">^</a></b></span> <span class="reference-text">Breaking the language barrier: experimental evolution of non-native Aliivibrio fischeri in squid tailors luminescence to the host. <i>Symbiosis</i> <b>51</b>(1): 85-96. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fs13199-010-0074-2">10.1007/s13199-010-0074-2</a></span> </li> <li id="cite_note-Jones-26"><span class="mw-cite-backlink">^ <a href="#cite_ref-Jones_26-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Jones_26-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Jones_26-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Jones_26-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Jones_26-4"><sup><i><b>e</b></i></sup></a></span> <span class="reference-text">Counterillumination in the Hawaiian bobtail squid, <i>Euprymna scolopes</i> Berry (Mollusca : Cephalopoda). <i><a href="/wiki/Marine_Biology_(journal)" title="Marine Biology (journal)">Marine Biology</a></i> <b>144</b>(6): 1151-1155. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fs00227-003-1285-3">10.1007/s00227-003-1285-3</a></span> </li> <li id="cite_note-27"><span class="mw-cite-backlink"><b><a href="#cite_ref-27">^</a></b></span> <span class="reference-text">Differentially expressed genes reveal adaptations between free-living and symbiotic niches of <i>Aliivibrio fischeri</i> in a fully established mutualism. <i><a href="/wiki/Canadian_Journal_of_Microbiology" title="Canadian Journal of Microbiology">Canadian Journal of Microbiology</a></i> <b>52</b>(12): 1218-1227. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1139%2Fw06-088">10.1139/w06-088</a></span> </li> <li id="cite_note-28"><span class="mw-cite-backlink"><b><a href="#cite_ref-28">^</a></b></span> <span class="reference-text">Tong, D., N.S. Rozas, T.H. Oakley, J. Mitchell, N.J. Colley & M.J. McFall-Ngai 2009. Evidence for light perception in a bioluminescent organ. <i>PNAS</i> <b>106</b>(24): 9836–9841. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.0904571106">10.1073/pnas.0904571106</a></span> </li> <li id="cite_note-29"><span class="mw-cite-backlink"><b><a href="#cite_ref-29">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPeyerPankeyOakleyMcFall-Ngai2014" class="citation journal cs1">Peyer, Suzanne M.; Pankey, M. Sabrina; Oakley, Todd H.; McFall-Ngai, Margaret J. (February 2014). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4000693">"Eye-specification genes in the bacterial light organ of the bobtail squid Euprymna scolopes, and their expression in response to symbiont cues"</a>. <i>Mechanisms of Development</i>. <b>131</b>: 111–126. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.mod.2013.09.004">10.1016/j.mod.2013.09.004</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4000693">4000693</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/24157521">24157521</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Mechanisms+of+Development&rft.atitle=Eye-specification+genes+in+the+bacterial+light+organ+of+the+bobtail+squid+Euprymna+scolopes%2C+and+their+expression+in+response+to+symbiont+cues&rft.volume=131&rft.pages=111-126&rft.date=2014-02&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4000693%23id-name%3DPMC&rft_id=info%3Apmid%2F24157521&rft_id=info%3Adoi%2F10.1016%2Fj.mod.2013.09.004&rft.aulast=Peyer&rft.aufirst=Suzanne+M.&rft.au=Pankey%2C+M.+Sabrina&rft.au=Oakley%2C+Todd+H.&rft.au=McFall-Ngai%2C+Margaret+J.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4000693&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEuprymna+scolopes" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Euprymna_scolopes&action=edit&section=8" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li>Callaerts, P., P.N. Lee, B. Hartmann, C. Farfan, D.W.Y. Choy, K. Ikeo, K.F. Fischbach, W.J. Gehring & G. de Couet 2002. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20040328104246/http://www.cephbase.utmb.edu/refdb/pdf/7087.pdf">"HOX genes in the sepiolid squid <i>Euprymna scolopes</i>: Implications for the evolution of complex body plans"</a> <span class="cs1-format">(PDF)</span>. Archived from the original on 2004-03-28.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=HOX+genes+in+the+sepiolid+squid+Euprymna+scolopes%3A+Implications+for+the+evolution+of+complex+body+plans.&rft_id=http%3A%2F%2Fwww.cephbase.utmb.edu%2Frefdb%2Fpdf%2F7087.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEuprymna+scolopes" class="Z3988"></span><span class="cs1-maint citation-comment"><code class="cs1-code">{{<a href="/wiki/Template:Cite_web" title="Template:Cite web">cite web</a>}}</code>: CS1 maint: bot: original URL status unknown (<a href="/wiki/Category:CS1_maint:_bot:_original_URL_status_unknown" title="Category:CS1 maint: bot: original URL status unknown">link</a>)</span> <span style="font-size:85%;">(465 KB)</span> <i>PNAS</i> <b>99</b>(4): 2088–2093.</li></ul> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Euprymna_scolopes&action=edit&section=9" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20050817053958/http://www.cephbase.utmb.edu/spdb/speciesc.cfm?CephID=140">"CephBase: Euprymna scolopes"</a>. Archived from <a rel="nofollow" class="external text" href="http://www.cephbase.utmb.edu/spdb/speciesc.cfm?CephID=140">the original</a> on 2005-08-17.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=CephBase%3A+Euprymna+scolopes&rft_id=http%3A%2F%2Fwww.cephbase.utmb.edu%2Fspdb%2Fspeciesc.cfm%3FCephID%3D140&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEuprymna+scolopes" class="Z3988"></span></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20070426122929/http://web.uconn.edu/mcbstaff/graf/VfEs/VfEssym.htm">The Light-Organ Symbiosis of <i>Vibrio fischeri</i> and the Hawaiian squid, <i>Euprymna scolopes</i></a></li> <li><a rel="nofollow" class="external text" href="http://feedthedatamonster.com/home/2013/9/11/mutualism-of-the-month-hawaiian-bobtail-squid">Mutualism of the Month: Hawai‘ian bobtail squid</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20190730161135/http://feedthedatamonster.com/home/2013/9/11/mutualism-of-the-month-hawaiian-bobtail-squid">Archived</a> 2019-07-30 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl ul,.mw-parser-output .hlist ol dl,.mw-parser-output .hlist ol ol,.mw-parser-output .hlist ol ul,.mw-parser-output .hlist ul dl,.mw-parser-output .hlist ul ol,.mw-parser-output .hlist ul ul{display:inline}.mw-parser-output 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title="wikispecies:Euprymna scolopes">Euprymna scolopes</a></span></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Animal_Diversity_Web" title="Animal Diversity Web">ADW</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://animaldiversity.org/accounts/Euprymna_scolopes/">Euprymna_scolopes</a></span></span></li> <li><span style="white-space:nowrap;">BioLib: <span class="uid"><a rel="nofollow" class="external text" href="https://www.biolib.cz/en/taxon/id134486">134486</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Barcode_of_Life_Data_System" title="Barcode of Life Data System">BOLD</a>: <span class="uid"><a rel="nofollow" class="external text" href="http://www.boldsystems.org/index.php/TaxBrowser_TaxonPage?taxid=30214">30214</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Catalogue_of_Life" title="Catalogue of Life">CoL</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.catalogueoflife.org/data/taxon/6HDV4">6HDV4</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Encyclopedia_of_Life" title="Encyclopedia of Life">EoL</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://eol.org/pages/491890">491890</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Global_Biodiversity_Information_Facility" title="Global Biodiversity Information Facility">GBIF</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.gbif.org/species/2289092">2289092</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/INaturalist" title="INaturalist">iNaturalist</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://inaturalist.org/taxa/143624">143624</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Interim_Register_of_Marine_and_Nonmarine_Genera" title="Interim Register of Marine and Nonmarine Genera">IRMNG</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.irmng.org/aphia.php?p=taxdetails&id=10210411">10210411</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Integrated_Taxonomic_Information_System" title="Integrated Taxonomic Information System">ITIS</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=556020">556020</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/IUCN_Red_List" title="IUCN Red List">IUCN</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://apiv3.iucnredlist.org/api/v3/taxonredirect/162598">162598</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/National_Center_for_Biotechnology_Information" title="National Center for Biotechnology Information">NCBI</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Info&id=6613">6613</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Ocean_Biodiversity_Information_System" title="Ocean Biodiversity Information System">OBIS</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://obis.org/taxon/342322">342322</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Open_Tree_of_Life" title="Open Tree of Life">Open Tree of Life</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://tree.opentreeoflife.org/taxonomy/browse?id=528814">528814</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Plazi" title="Plazi">Plazi</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://treatment.plazi.org/id/4A2A723C-8796-9418-60D7-39848970FDAA">4A2A723C-8796-9418-60D7-39848970FDAA</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/SeaLifeBase" 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