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You could also send an email to the <a href="https://meta.wikimedia.org/wiki/Special:MyLanguage/Stewards" class="extiw" title="m:Special:MyLanguage/Stewards">stewards</a> <a href="https://meta.wikimedia.org/wiki/Special:MyLanguage/VRT" class="extiw" title="m:Special:MyLanguage/VRT">VRT</a> queue at <kbd>stewards@wikimedia.org</kbd> including all above details. </p> </div> <p>Other useful links: <a href="https://meta.wikimedia.org/wiki/Global_blocks" class="extiw" title="m:Global blocks">Global blocks</a> · <a href="/wiki/Help:I_have_been_blocked" title="Help:I have been blocked">Help:I have been blocked</a> </p> </div></li></ul><hr /> <div id="viewsourcetext">You can view and copy the source of this page:</div><textarea readonly="" accesskey="," id="wpTextbox1" cols="80" rows="25" style="" class="mw-editfont-monospace" lang="en" dir="ltr" name="wpTextbox1">==Symbiosis== {{Further|Counter-illumination}} ''Euprymna scolopes'' lives in a [[symbiosis|symbiotic]] relationship with the [[bioluminescent]] [[bacteria]] ''[[Aliivibrio fischeri]]'', which inhabits a special light organ in the squid's mantle. To allows this symbiotic relationship, ''Crumbs protein'' must first induce ''Apopstosis'', which kills superficial epithelial tissue found in ''Euprymna scolopes.'' Apopstosis then helps create ''crypt'' epithelial cells, these cells directly take in the bioluminescence bacteria received from the ''Aliivibrio fischeri''.<ref>{{Cite journal |last1=Peyer |first1=Suzanne M. |last2=Heath-Heckman |first2=Elizabeth A. C. |last3=McFall-Ngai |first3=Margaret J. |date=2017-11-01 |title=Characterization of the cell polarity gene crumbs during the early development and maintenance of the squid–vibrio light organ symbiosis |journal=Development Genes and Evolution |language=en |volume=227 |issue=6 |pages=375–387 |doi=10.1007/s00427-017-0576-5 |issn=1432-041X |pmc=5519459 |pmid=28105525}}</ref> The bacteria are fed a [[sugar]] and [[amino acid]] 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 ([[counter-illumination]]).<ref name="Young">Young, R.E. & C.F. Roper 1976. Bioluminescent countershading in midwater animals: evidence from living squid. ''Science'' '''191'''(4231): 1046–1048. {{doi|10.1126/science.1251214}}</ref> ''E. scolopes'' serves as a [[model organism]] for animal-bacterial symbiosis and its relationship with ''A. fischeri'' has been carefully studied.<ref>DeLoney, C.R., T.M. Bartley & K.L. Visick 2002. {{cite web |url= http://www.cephbase.utmb.edu/refdb/pdf/7468.pdf |title= Role for phosphoglucomutase in ''Aliivibrio fischeri''-''Euprymna scolopes'' symbiosis. |url-status= dead |archive-url= https://web.archive.org/web/20040328114942/http://www.cephbase.utmb.edu/refdb/pdf/7468.pdf |archive-date= March 28, 2004 }}&nbsp;{{small|(221&nbsp;KB)}} ''Journal of Bacteriology'' '''184'''(18): 5121-5129.</ref><ref>Dunlap, P.V., K. Kitatsukamoto, J.B. Waterbury & S.M. Callahan 1995. {{cite web |url= http://www.cephbase.utmb.edu/refdb/pdf/7159.pdf |title= Isolation and characterization of a visibly luminous variant of ''Aliivibrio fischeri'' strain ES114 form the sepiolid squid ''Euprymna scolopes''. |url-status= dead |archive-url= https://web.archive.org/web/20040328141759/http://www.cephbase.utmb.edu/refdb/pdf/7159.pdf |archive-date= March 28, 2004 }}&nbsp;{{small|(105&nbsp;KB)}} ''Archives of Microbiology'' '''164'''(3): 194-202.</ref><ref>Foster, J.S., M.A. Apicella & M.J. McFall-Ngai 2000. {{cite web |url= http://www.cephbase.utmb.edu/refdb/pdf/7360.pdf |title= ''Aliivibrio fischeri'' lipopolysaccharide induces developmental apoptosis, but not complete morphogenesis, of the ''Euprymna scolopes'' light organ. |url-status= dead |archive-url= https://web.archive.org/web/20040328150636/http://www.cephbase.utmb.edu/refdb/pdf/7360.pdf |archive-date= March 28, 2004 }}&nbsp;{{small|(610&nbsp;KB)}} ''Developmental Biology'' '''226'''(2): 242-254.</ref><ref>Hanlon, R.T., M.F. Claes, S.E. Ashcraft & P.V. Dunlap 1997. {{cite web |url= http://www.cephbase.utmb.edu/refdb/pdf/7724.pdf |title= Laboratory culture of the sepiolid squid ''Euprymna scolopes'': A model system for bacteria-animal symbiosis. |url-status= dead |archive-url= https://web.archive.org/web/20040328183009/http://www.cephbase.utmb.edu/refdb/pdf/7724.pdf |archive-date= March 28, 2004 }}&nbsp;{{small|(2.38&nbsp;MB)}} ''Biological Bulletin'' '''192'''(3): 364-374.</ref><ref>Lee, K.-H. & E.G. Ruby 1995. {{cite web |url= http://www.cephbase.utmb.edu/refdb/pdf/4338.pdf |title= Symbiotic role of the viable but nonculturable state of ''Aliivibrio fischeri'' in Hawaiian coastal seawater. |url-status= dead |archive-url= https://web.archive.org/web/20040328145721/http://www.cephbase.utmb.edu/refdb/pdf/4338.pdf |archive-date= March 28, 2004 }}&nbsp;{{small|(249&nbsp;KB)}} ''Applied and Environmental Microbiology'' '''61'''(1): 278-283.</ref><ref>Lemus, J.D. & M.J. McFall-Ngai 2000. {{cite web |url= http://www.cephbase.utmb.edu/refdb/pdf/7610.pdf |title= Alterations in the protoeme of the ''Euprymna scolopes'' light organ in response to symbiotic ''Aliivibrio fischeri''. |url-status= dead |archive-url= https://web.archive.org/web/20040328174755/http://www.cephbase.utmb.edu/refdb/pdf/7610.pdf |archive-date= March 28, 2004 }}&nbsp;{{small|(2.10&nbsp;MB)}} ''Applied and Environmental Microbiology'' '''66''': 4091-4097.</ref><ref>Millikan, D.S. & E.G. Ruby 2003. {{cite web |url= http://www.cephbase.utmb.edu/refdb/pdf/7850.pdf |title= FlrA, a s<sup>54</sup>-Dependent Transcriptional Activator in ''Aliivibrio fischeri'', is required for motility and symbiotic light-organ colonization. |url-status= dead |archive-url= https://web.archive.org/web/20040328200403/http://www.cephbase.utmb.edu/refdb/pdf/7850.pdf |archive-date= March 28, 2004 }}&nbsp;{{small|(382&nbsp;KB)}} ''Journal of Bacteriology'' (American Society for Microbiology) '''185'''(12): 3547-3557.</ref><ref>Montgomery, M.K. & M. McFall-Ngai 1998. {{cite web |url= http://www.cephbase.utmb.edu/refdb/pdf/5803.pdf |title= Late postembryonic development of the symbiotic light organ of ''Euprymna scolopes'' (Cephalopoda: Sepiolidae). |url-status= dead |archive-url= https://web.archive.org/web/20040328163337/http://www.cephbase.utmb.edu/refdb/pdf/5803.pdf |archive-date= March 28, 2004 }}&nbsp;{{small|(6.10&nbsp;MB)}} ''Biological Bulletin'' '''195''': 326-336.</ref> ===Acquisition=== The bioluminescent bacterium, ''A. fischeri'', is horizontally transmitted throughout the ''E. scolopes'' population. Hatchlings lack these necessary bacteria and must carefully select for them in a marine world saturated with other microorganisms.<ref name="Chun">Effects of colonization, luminescence, and autoinducer on host transcription during development of the squid-vibrio association.''[[Proceedings of the National Academy of Sciences of the United States of America]]'' '''105'''(32): 11323-11328. {{doi|10.1073/pnas.0802369105}}</ref> To effectively capture these cells, ''E. scolopes'' secretes [[mucus]] in response to [[peptidoglycan]] (a major [[cell wall]] component of [[bacteria]]).<ref name="Nyholm">The evolutionary ecology of a sepiolid squid-''Aliivibrio'' association: from cell to environment. ''[[Vie et Milieu]] '''58'''(2): 175-184. {{ISSN|0240-8759}}''</ref> The mucus inundates the [[cilia]]ted 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, ''A. fischeri'' is able to outcompete other bacteria in the mucus.<ref name="Nyholm" /> [[File:Hawaiian Bobtail squid.tiff|thumb|left|300px|Adult ''Euprymna scolopes'' with scale.]] As ''A. fischeri'' cells aggregate in the mucus, they must use their [[flagella]] to migrate through the [[wiktionary:pore|pore]]s and down into the ciliated ducts of the light organ and endure another barrage of host factors meant to ensure only ''A. fischeri'' colonization.<ref name="Nyholm" /> Besides the relentless host-derived current that forces [[motility]]-challenged bacteria out of the pores, a number of reactive oxygen species makes the environment unbearable.<ref name="Nyholm" /> Squid [[halide peroxidase]] is the main enzyme responsible for crafting this [[microbicide|microbiocidal]] environment, using [[hydrogen peroxide]] as a substrate, but ''A. fischeri'' has evolved a brilliant counterattack. ''A. fischeri'' possesses a periplasmic [[catalyst|catalase]] that captures hydrogen peroxide before it can be used by the squid halide peroxidase, thus inhibiting the enzyme indirectly.<ref name="Nyholm" /> Once through these ciliated ducts, ''A. fischeri'' cells swim on towards the antechamber, a large [[epithelium|epithelial]]-lined space, and colonize the narrow epithelial crypts.<ref name="Nyholm" /> The bacteria thrive on the host-derived amino acids and sugars in the [[antechamber]] and quickly fill the crypt spaces within 10 to 12 hours after hatching.<ref name="Visick">An exclusive contract: Specificity in the ''Aliivibrio fischeri'' ''Euprymna scolopes'' partnership. ''[[Journal of Bacteriology]]'' '''182'''(7): 1779-1787. {{ISSN|0021-9193}}</ref> ===Ongoing relationship=== Every second, a juvenile squid ventilates about {{convert|2.6|ml|abbr=on}} of ambient [[seawater]] through its mantle cavity. Only a single ''A. fischeri'' cell, one/1-millionth of the total volume, is present with each ventilation.<ref name="Nyholm" /> The increased amino acids and sugars feed the [[metabolism|metabolically]] demanding [[bioluminescence]] of the ''A. fischeri'', and in 12 hours, the bioluminescence peaks and the juvenile squid is able to counterilluminate less than a day after hatching.<ref name="Visick" /> Bioluminescence demands a substantial amount of energy from a bacterial cell. It is estimated to demand 20% of a cell's metabolic potential.<ref name="Visick" /> Nonluminescent strains of ''A. fischeri'' would have a definite competitive advantage over the luminescent wild-type, however nonluminescent mutants are never found in the light organ of the ''E. scolopes''.<ref name="Visick" /> In fact, experimental procedures have shown that removing the [[gene]]s responsible for light production in ''A. fischeri'' drastically reduces colonization efficiency.<ref name="Visick" /> Luminescent cells, with functioning [[luciferase]], may have a higher affinity for [[oxygen]] than for [[peroxidases]], thereby negating the toxic effects of the peroxidases.<ref name="Timmins">The evolution of bioluminescent oxygen consumption as an ancient oxygen detoxification mechanism.''[[Journal of Molecular Evolution]]'' '''52'''(4): 321-332. {{ISSN|0022-2844}}</ref> For this reason, bioluminescence is thought to have evolved as an ancient oxygen detoxification mechanism in bacteria.<ref name="Timmins"/> ===Venting=== Despite all the effort that goes into obtaining luminescent ''A. fischeri'', the host squid jettisons most of the cells daily. This process, known as "venting", is responsible for the disposal of up to 95% of ''A. fischeri'' in the light organ every morning at dawn.<ref name="Schuster">Breaking the language barrier: experimental evolution of non-native Aliivibrio fischeri in squid tailors luminescence to the host. ''Symbiosis'' '''51'''(1): 85-96. {{doi|10.1007/s13199-010-0074-2}}</ref> 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.<ref name="Jones">Counterillumination in the Hawaiian bobtail squid, ''Euprymna scolopes'' Berry (Mollusca : Cephalopoda). ''[[Marine Biology (journal)|Marine Biology]]'' '''144'''(6): 1151-1155. {{doi|10.1007/s00227-003-1285-3}}</ref> During the day when the squid are inactive and hidden, bioluminescence is unnecessary, and expelling the ''A. fischeri'' conserves energy. Another, more evolutionarily important reason may be that daily venting ensures selection for ''A. fischeri'' that have evolved specificity for a particular host, but can survive outside of the light organ.<ref>Differentially expressed genes reveal adaptations between free-living and symbiotic niches of ''Aliivibrio fischeri'' in a fully established mutualism. ''[[Canadian Journal of Microbiology]]'' '''52'''(12): 1218-1227. {{doi|10.1139/w06-088}}</ref> Since ''A. fischeri'' is transmitted horizontally in ''E. scolopes'', maintaining a stable population of them in the open ocean is essential in supplying future generations of squid with functioning light organs. </textarea><div class="templatesUsed"></div><p id="mw-returnto">Return to <a href="/wiki/Euprymna_scolopes" title="Euprymna scolopes">Euprymna scolopes</a>.</p> <noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1&useformat=desktop" alt="" width="1" height="1" style="border: none; position: absolute;"></noscript> <div class="printfooter" data-nosnippet="">Retrieved from "<a dir="ltr" href="https://en.wikipedia.org/wiki/Euprymna_scolopes">https://en.wikipedia.org/wiki/Euprymna_scolopes</a>"</div></div> <div id="catlinks" class="catlinks catlinks-allhidden" data-mw="interface"></div> </div> </main> </div> <div class="mw-footer-container"> <footer id="footer" class="mw-footer" > <ul id="footer-info"> </ul> <ul id="footer-places"> <li id="footer-places-privacy"><a href="https://foundation.wikimedia.org/wiki/Special:MyLanguage/Policy:Privacy_policy">Privacy policy</a></li> <li id="footer-places-about"><a href="/wiki/Wikipedia:About">About Wikipedia</a></li> <li id="footer-places-disclaimers"><a href="/wiki/Wikipedia:General_disclaimer">Disclaimers</a></li> <li id="footer-places-contact"><a href="//en.wikipedia.org/wiki/Wikipedia:Contact_us">Contact Wikipedia</a></li> <li id="footer-places-wm-codeofconduct"><a href="https://foundation.wikimedia.org/wiki/Special:MyLanguage/Policy:Universal_Code_of_Conduct">Code of Conduct</a></li> <li id="footer-places-developers"><a href="https://developer.wikimedia.org">Developers</a></li> <li id="footer-places-statslink"><a href="https://stats.wikimedia.org/#/en.wikipedia.org">Statistics</a></li> <li id="footer-places-cookiestatement"><a href="https://foundation.wikimedia.org/wiki/Special:MyLanguage/Policy:Cookie_statement">Cookie statement</a></li> <li id="footer-places-mobileview"><a href="//en.m.wikipedia.org/w/index.php?title=Euprymna_scolopes&action=edit&section=1&mobileaction=toggle_view_mobile" class="noprint stopMobileRedirectToggle">Mobile view</a></li> </ul> <ul id="footer-icons" class="noprint"> <li id="footer-copyrightico"><a href="https://wikimediafoundation.org/" class="cdx-button cdx-button--fake-button cdx-button--size-large cdx-button--fake-button--enabled"><img src="/static/images/footer/wikimedia-button.svg" width="84" height="29" alt="Wikimedia Foundation" loading="lazy"></a></li> <li id="footer-poweredbyico"><a href="https://www.mediawiki.org/" class="cdx-button cdx-button--fake-button cdx-button--size-large cdx-button--fake-button--enabled"><img src="/w/resources/assets/poweredby_mediawiki.svg" alt="Powered by MediaWiki" width="88" height="31" loading="lazy"></a></li> </ul> </footer> </div> </div> </div> <div class="vector-settings" id="p-dock-bottom"> <ul></ul> </div><script>(RLQ=window.RLQ||[]).push(function(){mw.config.set({"wgHostname":"mw-web.codfw.main-5c59558b9d-flhfz","wgBackendResponseTime":211,"wgPageParseReport":{"limitreport":{"cputime":"0.054","walltime":"0.069","ppvisitednodes":{"value":418,"limit":1000000},"postexpandincludesize":{"value":17740,"limit":2097152},"templateargumentsize":{"value":6556,"limit":2097152},"expansiondepth":{"value":9,"limit":100},"expensivefunctioncount":{"value":0,"limit":500},"unstrip-depth":{"value":0,"limit":20},"unstrip-size":{"value":469,"limit":5000000},"entityaccesscount":{"value":0,"limit":400},"timingprofile":["100.00% 47.365 1 -total"," 99.86% 47.298 2 Template:Blocked_text"," 41.03% 19.435 2 Template:Replace"," 40.58% 19.221 1 Template:Colocationwebhost"," 36.63% 17.348 1 Template:Hidden"," 15.74% 7.457 1 Template:Hidden_begin"," 14.93% 7.071 1 Template:Tlx"," 2.09% 0.988 1 Template:Hidden_end"," 1.95% 0.924 1 MediaWiki:Wikimedia-globalblocking-blockedtext-mistake"," 1.59% 0.754 1 MediaWiki:Wikimedia-globalblocking-blockedtext-mistake-email-steward"]},"scribunto":{"limitreport-timeusage":{"value":"0.011","limit":"10.000"},"limitreport-memusage":{"value":1043320,"limit":52428800}},"cachereport":{"origin":"mw-web.codfw.main-5c59558b9d-flhfz","timestamp":"20241130023851","ttl":2592000,"transientcontent":false}}});});</script> </body> </html>