View source for Bioluminescence

{{Short description|Emission of light by a living organism}}
{{Distinguish |Iridescence |Structural coloration |Phosphorescence}}
{{Use dmy dates |date=September 2020}}
{{Use American English|date=January 2021}}<!-- ?? -->
[[File:Photinus pyralis Firefly glowing.jpg|thumb|Flying and glowing [[firefly]], ''[[Photinus pyralis]]'']]
[[File:Lampyris noctiluca.jpg|thumb|upright |Female [[glowworm]], ''[[Lampyris noctiluca]]'']]
'''Bioluminescence''' is the emission of [[light]] during a [[chemiluminescence]] reaction by living [[organism]]s.<ref name=":0">{{Cite journal |last=Schramm |first=Stefan |last2=Weiß |first2=Dieter |date=2024 |title=Bioluminescence – The Vibrant Glow of Nature and its Chemical Mechanisms |url=https://chemistry-europe.onlinelibrary.wiley.com/doi/10.1002/cbic.202400106 |journal=ChemBioChem |language=en |volume=25 |issue=9 |pages=e202400106 |doi=10.1002/cbic.202400106 |issn=1439-7633|doi-access=free }}</ref> Bioluminescence occurs in diverse organisms ranging from marine [[vertebrate]]s and [[invertebrate]]s, as well as in some [[Fungus|fungi]], microorganisms including some [[bioluminescent bacteria]], [[Dinoflagellate|dinoflagellates]] and terrestrial arthropods such as [[Firefly|fireflies]]. In some animals, the light is bacteriogenic, produced by [[symbiosis|symbiotic]] bacteria such as those from the genus ''[[Vibrio]]'';<ref>{{Cite web |last=Ples |first=Marek |date=2021-11-11 |title=Lab Snapshots by Marek Ples; Microbiology - The biology on a different level |url=https://weirdscience.eu/Mikrobio/?id=1 |access-date=2023-07-02 |website=weirdscience.eu}}</ref> in others, it is autogenic, produced by the animals themselves.

In most cases, the principal chemical reaction in bioluminescence involves the reaction of a substrate called [[luciferin]] and an [[enzyme]], called [[luciferase]]. Because these are generic names, luciferins and luciferases are often distinguished by the species or group, e.g. [[firefly luciferin]] or [[Vargulin|cypridina luciferin]]. In all characterized cases, the enzyme [[Catalysis|catalyzes]] the [[Redox|oxidation]] of the luciferin resulting in [[excited state]] oxyluciferin, which is the light emitter of the reaction. Upon their decay to the [[ground state]] they emit visible light. In all known cases of bioluminescence the production of the excited state molecules involves the decomposition of [[organic peroxides]].<ref name=":0" />

In some species, the luciferase requires other [[Cofactor (biochemistry)|cofactors]], such as [[calcium]] or [[magnesium]] ions, and sometimes also the energy-carrying molecule [[adenosine triphosphate]] (ATP). In [[evolution]], luciferins vary little: one in particular, [[coelenterazine]], is found in 11 different animal [[phylum|phyla]], though in some of these, the animals obtain it through their diet. Conversely, luciferases vary widely between different species. Bioluminescence has arisen over 40 times in [[Evolutionary history of life|evolutionary history]].

Both [[Aristotle]] and [[Pliny the Elder]] mentioned that damp wood sometimes gives off a glow. Many centuries later [[Robert Boyle]] showed that oxygen was involved in the process, in both wood and glowworms. It was not until the late nineteenth century that bioluminescence was properly investigated. The phenomenon is widely distributed among animal groups, especially in marine environments. On land it occurs in fungi, bacteria and some groups of [[invertebrate]]s, including [[insect]]s.

The uses of bioluminescence by animals include [[counterillumination]] camouflage, [[mimicry]] of other animals, for example to lure prey, and [[signalling theory|signaling]] to other individuals of the same species, such as to attract mates. In the laboratory, luciferase-based systems are used in genetic engineering and biomedical research. Researchers are also investigating the possibility of using bioluminescent systems for street and decorative lighting, and a bioluminescent plant has been created.<ref>Callaway, E. 2013. Glowing plants spark debate. ''Nature'', 498:15–16, 4 June 2013. http://www.nature.com/news/glowing-plants-spark-debate-1.13131</ref>

==History==
Before the development of the [[safety lamp]] for use in coal mines, dried fish skins were used in Britain and Europe as a weak source of light.<ref>{{cite book | last=Smiles | first=Samuel | author-link=Samuel Smiles | title=Lives of the Engineers | volume=III (George and Robert Stephenson) | page=107 | year=1862 | publisher=John Murray | location=London | isbn=978-0-7153-4281-7}} (ISBN refers to the David & Charles reprint of 1968 with an introduction by [[L. T. C. Rolt]])</ref> This experimental form of illumination avoided the necessity of using candles which risked sparking explosions of [[firedamp]].<ref name="Freese2006">{{cite book |last=Freese |first=Barbara |title=Coal: A Human History |url=https://books.google.com/books?id=p4z7QjaBrCkC&pg=PA51 |year=2006 |publisher=Arrow |isbn=978-0-09-947884-3 |page=51}}</ref> In 1920, the American zoologist [[E. Newton Harvey]] published a monograph, ''The Nature of Animal Light'', summarizing early work on bioluminescence. Harvey notes that [[Aristotle]] mentions light produced by dead fish and flesh, and that both Aristotle and [[Pliny the Elder]] (in his ''[[Natural History (Pliny)|Natural History]]'') mention light from damp wood. He records that [[Robert Boyle]] experimented on these light sources, and showed that both they and the glowworm require air for light to be produced. Harvey notes that in 1753, J. Baker identified the flagellate ''[[Noctiluca]]'' "as a luminous animal" "just visible to the naked eye",<ref>Harvey cites this as Baker, J.: 1743–1753, ''The Microscope Made Easy and Employment for the Microscope''.</ref> and in 1854 [[Johann Florian Heller]] (1813–1871) identified strands ([[hypha]]e) of fungi as the source of light in dead wood.<ref>{{cite book |last=Harvey |first=E. Newton |year=1920 |url=https://www.biodiversitylibrary.org/bibliography/57246#/summary |title=The Nature of Animal Light |location=Philadelphia & London |publisher=J. B. Lippencott |page=1}}</ref>

[[James Hingston Tuckey|Tuckey]], in his posthumous 1818 ''Narrative of the Expedition to the Zaire'', described catching the animals responsible for luminescence. He mentions pellucids, crustaceans (to which he ascribes the milky whiteness of the water), and cancers (shrimps and crabs). Under the microscope he described the "luminous property" to be in the brain, resembling "a most brilliant amethyst about the size of a large pin's head".<ref>{{cite journal |last=Tuckey |first=James Hingston |author-link=James Hingston Tuckey |title=Narrative of the Expedition to the Zaire |journal=[[Annals of Philosophy]] |editor-last=Thomson |editor-first=Thomas |editor-link=Thomas Thomson (chemist) |volume=11 |date=May 1818 |issue=65 |page=392 |url=https://www.biodiversitylibrary.org/item/53908#page/412/mode/1up}}</ref>

[[Charles Darwin]] noticed bioluminescence in the sea, describing it in his ''Journal'':

<blockquote>While sailing in these latitudes on one very dark night, the sea presented a wonderful and most beautiful spectacle. There was a fresh breeze, and every part of the surface, which during the day is seen as foam, now glowed with a pale light. The vessel drove before her bows two billows of liquid phosphorus, and in her wake she was followed by a milky train. As far as the eye reached, the crest of every wave was bright, and the sky above the horizon, from the reflected glare of these livid flames, was not so utterly obscure, as over the rest of the heavens.<ref name=Journal>{{cite book |last=Darwin |first=Charles |author-link=Charles Darwin |title=Narrative of the surveying voyages of His Majesty's Ships Adventure and Beagle between the years 1826 and 1836, describing their examination of the southern shores of South America, and the Beagle's circumnavigation of the globe. Journal and remarks. 1832–1836. |date=1839 |publisher=Henry Colburn |pages=190–192 |url=http://darwin-online.org.uk/content/frameset?viewtype=side&itemID=F10.3&pageseq=30}}</ref></blockquote>

Darwin also observed a luminous "jelly-fish of the genus Dianaea",<ref name=Journal/> noting that: "When the waves scintillate with bright green sparks, I believe it is generally owing to minute crustacea. But there can be no doubt that very many other pelagic animals, when alive, are phosphorescent."<ref name=Journal/> He guessed that "a disturbed electrical condition of the atmosphere"<ref name=Journal/> was probably responsible. Daniel Pauly comments that Darwin "was lucky with most of his guesses, but not here",<ref name=Pauly2004>{{cite book |last=Pauly |first=Daniel |title=Darwin's Fishes: An Encyclopedia of Ichthyology, Ecology, and Evolution |url=https://books.google.com/books?id=lTDa0fM4vQIC&pg=PA16 |date=13 May 2004 |publisher=Cambridge University Press |isbn=978-1-139-45181-9 |pages=15–16}}</ref> noting that biochemistry was too little known, and that the complex evolution of the marine animals involved "would have been too much for comfort".<ref name=Pauly2004/>

[[File:Aequorea4.jpg|thumb|[[Osamu Shimomura]] isolated the [[photoprotein]] [[aequorin]] and its cofactor [[coelenterazine]] from the crystal jelly ''[[Aequorea victoria]]'' in 1961.<ref name="pmid7654844">{{cite journal |last=Shimomura |first=O. |title=A short story of aequorin |journal=The Biological Bulletin |volume=189 |issue=1 |pages=1–5 |date=August 1995 |pmid=7654844 |doi=10.2307/1542194 |url=http://www.biolbull.org/cgi/pmidlookup?view=long&pmid=7654844 |jstor=1542194}}</ref>]]

Bioluminescence attracted the attention of the [[United States Navy]] in the [[Cold War]], since submarines in some waters can create a bright enough wake to be detected; a German submarine was sunk in the [[First World War]], having been detected in this way. The Navy was interested in predicting when such detection would be possible, and hence guiding their own submarines to avoid detection.<ref name=Economist/>

Among the anecdotes of navigation by bioluminescence is one recounted by the [[Apollo 13]] astronaut [[Jim Lovell]], who as a Navy pilot had found his way back to his aircraft carrier [[USS Shangri-La|USS ''Shangri-La'']] when his navigation systems failed. Turning off his cabin lights, he saw the glowing wake of the ship, and was able to fly to it and land safely.<ref name="Huth2013">{{cite book |last=Huth |first=John Edward |title=The Lost Art of Finding Our Way |url=https://books.google.com/books?id=daT6AAAAQBAJ&pg=PA423 |date=15 May 2013 |publisher=Harvard University Press |isbn=978-0-674-07282-4 |page=423}}</ref>

The [[France|French]] pharmacologist [[Raphaël Dubois]] carried out work on bioluminescence in the late nineteenth century. He studied [[Pyrophorus (beetle)|click beetles]] (''Pyrophorus'') and the marine [[Bivalvia|bivalve]] mollusc ''[[Pholas dactylus]]''. He refuted the old idea that bioluminescence came from phosphorus,<ref>{{cite web |last=Reshetiloff |first=Kathy |title=Chesapeake Bay night-lights add sparkle to woods, water |url=http://www.bayjournal.com/article/chesapeake_bay_nightlights_add_sparkle_to_woods_water |publisher=Bay Journal |access-date=16 December 2014 |date=1 July 2001 |archive-date=1 May 2019 |archive-url=https://web.archive.org/web/20190501200443/https://www.bayjournal.com/article/chesapeake_bay_nightlights_add_sparkle_to_woods_water |url-status=dead }}</ref>{{efn |However, the name 'phosphorus', as used in the 17th century, did not necessarily mean the modern element; any substance that glowed by itself could be given this name, meaning "light bearer".<ref>{{cite web |title=Luminescence |url=https://www.britannica.com/EBchecked/topic/351229/luminescence/68942/Early-investigations |publisher=Encyclopædia Britannica |access-date=16 December 2014}}</ref>}} and demonstrated that the process was related to the oxidation of a specific compound, which he named [[luciferin]], by an [[enzyme]].<ref>{{cite journal |last=Poisson |first=Jacques |date=April 2010 | title=Raphaël Dubois, from pharmacy to bioluminescence |journal=Rev Hist Pharm (Paris) |volume=58 |issue=365 |pages=51–56 | issn=0035-2349 | pmid=20533808 | language=fr<!--so where's the French title then?--> |doi=10.3406/pharm.2010.22136 }}</ref> He sent Harvey [[Siphon (mollusc)|siphons]] from the mollusc preserved in sugar. Harvey had become interested in bioluminescence as a result of visiting the South Pacific and Japan and observing phosphorescent organisms there. He studied the phenomenon for many years. His research aimed to demonstrate that luciferin, and the enzymes that act on it to produce light, were interchangeable between species, showing that all bioluminescent organisms had a common ancestor. However, he found this hypothesis to be false, with different organisms having major differences in the composition of their light-producing proteins. He spent the next 30 years purifying and studying the components, but it fell to the young Japanese chemist [[Osamu Shimomura]] to be the first to obtain crystalline luciferin. He used the [[Vargula hilgendorfii|sea firefly]] ''Vargula hilgendorfii'', but it was another ten years before he discovered the chemical's structure and published his 1957 paper ''Crystalline Cypridina Luciferin''.<ref name=Pieribone/> Shimomura, [[Martin Chalfie]], and [[Roger Y. Tsien]] won the 2008 [[Nobel Prize in Chemistry]] for their 1961 discovery and development of [[green fluorescent protein]] as a tool for biological research.<ref name=Nobel>{{cite news | title=The Nobel Prize in Chemistry 2008 | date=8 October 2008 | url=http://nobelprize.org/nobel_prizes/chemistry/laureates/2008/ | access-date=23 November 2014}}</ref>

Harvey wrote a detailed historical account on all forms of luminescence in 1957.<ref>{{Cite book |title=A History of Luminescence: From the Earliest Times Until 1900 |last=Harvey |first=E. Newton |publisher=American Philosophical Society |year=1957 |location=Philadelphia}}</ref> An updated book on bioluminescence covering also the twentieth and early twenty-first century was published recently.<ref>{{Cite book |title=Luminous Creatures: The History and Science of Light Production in Living Organisms |last=Anctil |first=Michel |publisher=McGill-Queen's University Press |year=2018 |isbn=978-0-7735-5312-5 |location=Montreal & Kingston, London, Chicago}}</ref><ref>{{cite web | url=http://www.tennessee.gov/environment/conservationist/docs/bluelights.pdf | title=Lovely and Dangerous Lights | last=Fulcher |first=Bob | work=Tennessee Conservationist Magazine | access-date=28 November 2014 | url-status=dead | archive-url=https://web.archive.org/web/20140814055028/http://www.tennessee.gov/environment/conservationist/docs/bluelights.pdf | archive-date=14 August 2014}}</ref>

== Evolution ==

In 1932 E. N. Harvey was among the first to propose how bioluminescence could have evolved.<ref>{{Cite journal |last=Harvey |first=E.N. |title=The evolution of bioluminescence and its relation to cell respiration |date=1932 |journal=Proceedings of the American Philosophical Society |volume=71 |pages=135–141}}</ref> In this early paper, he suggested that proto-bioluminescence could have arisen from respiratory chain proteins that hold fluorescent groups. This hypothesis has since been disproven, but it did lead to considerable interest in the origins of the phenomenon. Today, the two prevailing hypotheses (both concerning marine bioluminescence) are those put forth by [[Howard Harold Seliger|Howard Seliger]] in 1993 and Rees et al. in 1998.<ref name=Seliger>{{cite journal |last=Seliger |first=H.H. |title=Bioluminescence: excited states under cover of darkness |journal=Naval Research Reviews |date=1993 |volume=45 }}</ref><ref name=Rees>{{Cite journal |last=Rees |first=J. F. |display-authors=etal |title=The origins of marine bioluminescence: Turning oxygen defence mechanisms into deep-sea communication tools |journal=Journal of Experimental Biology |date=1998 |volume=201 |issue=Pt 8 |pages=1211–1221 |doi=10.1242/jeb.201.8.1211 |pmid=9510532 }}</ref>

Seliger's theory identifies luciferase enzymes as the catalyst for the evolution of bioluminescent systems. It suggests that the original purpose of luciferases was as mixed-function oxygenases. As the early ancestors of many species moved into deeper and darker waters natural selection favored the development of increased eye sensitivity and enhanced visual signals.<ref>{{Cite book |last=Widder |first=Edith A. |title=Bioluminescence |series=Adaptive Mechanisms in the Ecology of Vision |date=1999 |publisher=Springer |editor=Archer, S. |editor2=Djamgoz, M.B. |editor3=Loew, E. |editor4=Partridge, J.C. |editor5=Vallerga, S. |pages=555–581 }}</ref> If selection were to favor a mutation in the oxygenase enzyme required for the breakdown of pigment molecules (molecules often associated with spots used to attract a mate or distract a predator) it could have eventually resulted in external luminescence in tissues.<ref name=Seliger/>

Rees et al. use evidence gathered from the marine luciferin coelenterazine to suggest that selection acting on luciferins may have arisen from pressures to protect oceanic organisms from potentially deleterious reactive oxygen species (e.g. H<sub>2</sub>O<sub>2</sub> and O<sub>2</sub><sup>−</sup> ). The functional shift from antioxidation to bioluminescence probably occurred when the strength of selection for antioxidation defense decreased as early species moved further down the water column. At greater depths exposure to ROS is significantly lower, as is the endogenous production of ROS through metabolism.<ref name=Rees/>

While popular at first, Seliger's theory has been challenged, particularly on the biochemical and genetic evidence that Rees examines. What remains clear, however, is that bioluminescence has evolved independently at least 40 times.<ref>{{Cite journal |last=Haddock |first=S. H. D. |s2cid=3872860 |author-link=Steven Haddock |display-authors=etal |title=Bioluminescence in the Sea |journal=Annual Review of Marine Science |date=2010 |volume=2 |pages=443–493 |doi=10.1146/annurev-marine-120308-081028 |pmid=21141672 |bibcode=2010ARMS....2..443H }}</ref> Bioluminescence in fish began at least by the [[Cretaceous]] period. About 1,500 fish species are known to be bioluminescent; the capability evolved independently at least 27 times. Of these, 17 involved the taking up of bioluminous bacteria from the surrounding water while in the others, the intrinsic light evolved through chemical synthesis. These fish have become surprisingly diverse in the deep ocean and control their light with the help of their nervous system, using it not just to lure prey or hide from predators, but also for communication.<ref name="Thuesen Davis 2016"/><ref name="Yong">{{Cite web |url=http://phenomena.nationalgeographic.com/2016/06/08/the-many-origins-of-glowing-fish/ |archive-url=https://web.archive.org/web/20160609142140/http://phenomena.nationalgeographic.com/2016/06/08/the-many-origins-of-glowing-fish/ |url-status=dead |archive-date=9 June 2016 |title=Surprising History of Glowing Fish |last=Yong |first=Ed |date=8 June 2016 |website=Phenomena |publisher=National Geographic |access-date=11 June 2016}}</ref>

All bioluminescent organisms have in common that the reaction of a "luciferin" and oxygen is catalyzed by a luciferase to produce light.<ref>{{Cite journal |last1=Wilson |first1=Thérèse |last2=Hastings |first2=J. Woodland |date=1998 |title=Bioluminescence |journal=Annual Review of Cell and Developmental Biology |volume=14 |issue=1 |pages=197–230 |doi=10.1146/annurev.cellbio.14.1.197 |pmid=9891783}}</ref> McElroy and Seliger proposed in 1962 that the bioluminescent reaction evolved to detoxify oxygen, in parallel with photosynthesis.<ref>{{Cite journal |last1=McElroy |first1=William D. |last2=Seliger |first2=Howard H. |date=December 1962 |title=Biological Luminescence |journal=[[Scientific American]] |volume=207 |issue=6 |pages=76–91 |doi=10.1038/scientificamerican1262-76 |bibcode=1962SciAm.207f..76M |issn=0036-8733}}</ref>

Thuesen, Davis et al. showed in 2016 that bioluminescence has evolved independently 27 times within 14 fish clades across ray-finned fishes.<ref name="Thuesen Davis 2016">{{cite journal |last1=Thuesen |first1=Erik V. |last2=Davis |first2=Matthew P. |last3=Sparks |first3=John S. |last4=Smith |first4=W. Leo |title=Repeated and Widespread Evolution of Bioluminescence in Marine Fishes |journal=PLOS ONE |volume=11 |issue=6 |year=2016 |page=e0155154 |issn=1932-6203 |doi=10.1371/journal.pone.0155154|pmid=27276229 |pmc=4898709 |bibcode=2016PLoSO..1155154D |doi-access=free }}</ref> The oldest of these appears to be Stomiiformes and Myctophidae.<ref>{{Cite journal |last1=Paitio |first1=José |last2=Oba |first2=Yuichi |date=2024 |title=Luminous fishes: Endocrine and neuronal regulation of bioluminescence |url=https://linkinghub.elsevier.com/retrieve/pii/S2468550X23000564 |journal=Aquaculture and Fisheries |language=en |volume=9 |issue=3 |pages=486–500 |doi=10.1016/j.aaf.2023.04.003|bibcode=2024AqFis...9..486P |doi-access=free }}</ref> In sharks, bioluminescence has evolved only once.<ref>{{Cite journal |last=Ferrón |first=Humberto G. |date=2023 |editor-last=Ballen |editor-first=Gustavo |title=Illuminating the evolution of bioluminescence in sharks |url=https://onlinelibrary.wiley.com/doi/10.1111/pala.12641 |journal=Palaeontology |volume=66 |issue=1 |page=12641 |doi=10.1111/pala.12641 |bibcode=2023Palgy..6612641F |issn=0031-0239|doi-access=free |hdl=10550/85586 |hdl-access=free }}</ref>
<!-- Bioluminescence is not only present in ray finned fishes but also [[bacteria]], [[Dinoflagellate |dinoflagellates]], [[Radiolaria |radiolarians]], [[Ctenophora |ctenophores]], [[Cnidaria |cnidarians]], [[Mollusca |molluscs]], [[Crustacean |crustaceans]], [[Echinoderm |echinoderms]] and [[Tunicate |tunicates]]. It is most common among fish, squid and [[zooplankton]]. Some of these co-evolved with bioluminescent symbionts, usually bacteria.{{cn}}-->
Genomic analysis of [[Octocorallia|octocorals]] indicates that their ancestor was bioluminescent as long as 540 million years ago.<ref>{{cite journal |last1=DeLeo |first1=Danielle |last2=Bessho-Uehara |first2=Manabu |last3=Haddock |first3=Steven |last4=McFadden |first4=Catherine |last5=Quattrini |first5=Andrea |title=Evolution of bioluminescence in Anthozoa with emphasis on Octocorallia |journal=Proceedings of the Royal Society B |date=April 30, 2024 |volume=291 |issue=2021 |doi=10.1098/rspb.2023.2626|pmc=11040251 }}</ref>

==Chemical mechanism==
{{main |Luciferase}}
[[File:Firefly Luciferase Crystal Structure.rsh.png|thumb|[[Protein structure]] of the [[firefly luciferase|luciferase]] of the [[firefly]] ''[[Photinus pyralis]]''. The enzyme is a much larger molecule than luciferin.]]

Bioluminescence is a form of [[chemiluminescence]] where light energy is released by a chemical reaction. This reaction involves a light-emitting pigment, the [[luciferin]], and a [[luciferase]], the enzyme component.<ref name=Haddock/> Because of the diversity of luciferin/luciferase combinations, there are very few commonalities in the chemical mechanism. From currently studied systems, the only unifying mechanism is the role of molecular [[oxygen]]; often there is a concurrent release of [[carbon dioxide]] (CO<sub>2</sub>). For example, the firefly luciferin/luciferase reaction requires [[magnesium]] and [[Adenosine triphosphate|ATP]] and produces CO<sub>2</sub>, [[adenosine monophosphate]] (AMP) and [[pyrophosphate]] (PP) as waste products. Other cofactors may be required, such as calcium (Ca<sup>2+</sup>) for the [[photoprotein]] [[aequorin]], or [[magnesium]] (Mg<sup>2+</sup>) ions and ATP for the [[firefly luciferase]].<ref name="pmid6358519">{{cite journal | last=Hastings |first=J. W. | title=Biological diversity, chemical mechanisms, and the evolutionary origins of bioluminescent systems | journal=Journal of Molecular Evolution | volume=19 | issue=5 | pages=309–21 | year=1983 | pmid=6358519 | doi=10.1007/BF02101634 | issn=1432-1432 | bibcode=1983JMolE..19..309H | s2cid=875590}}</ref> Generically, this reaction can be described as:

:Luciferin + O<sub>2</sub><chem>->[\text{Luciferase}][\text{other cofactors}]</chem>Oxyluciferin + light energy

[[Image:Coelenterazine.svg|thumb|[[Coelenterazine]] is a [[luciferin]] found in many different marine phyla from [[Ctenophora|comb jellies]] to [[vertebrate]]s. Like all luciferins, it is oxidised to produce light.]]

Instead of a luciferase, the jellyfish ''[[Aequorea victoria]]'' makes use of another type of protein called a [[photoprotein]], in this case specifically [[aequorin]].<ref name=Shimomura_1962>{{cite journal |last=Shimomura |first=O. |author2=Johnson, F. H. |author3=Saiga, Y. | title=Extraction, purification and properties of aequorin, a bioluminescent protein from the luminous hydromedusan, Aequorea | journal=J Cell Comp Physiol | volume=59 | issue=3 | pages=223–39 | year=1962 | pmid=13911999 | doi=10.1002/jcp.1030590302 }}</ref> When calcium ions are added, rapid [[catalysis]] creates a brief flash quite unlike the prolonged glow produced by luciferase. In a second, much slower step, luciferin is regenerated from the oxidized (oxyluciferin) form, allowing it to recombine with aequorin, in preparation for a subsequent flash. Photoproteins are thus [[enzyme]]s, but with unusual reaction kinetics.<ref>{{cite journal |author1=Shimomura, O. |author2=Johnson, F.H. | title=Regeneration of the photoprotein aequorin | journal=Nature | volume=256 | pages=236–238 | year=1975 | doi=10.1038/256236a0 | pmid=239351 | issue=5514 |bibcode=1975Natur.256..236S |s2cid=4176627 }}</ref> Furthermore, some of the blue light released by aequorin in contact with calcium ions is absorbed by a [[green fluorescent protein]], which in turn releases green light in a process called [[Förster resonance energy transfer#BRET|resonant energy transfer]].<ref name=Morise_1974>{{cite journal |author1=Morise, H. |author2=Shimomura, O. |author3=Johnson, F.H. |author4=Winant, J. | title=Intermolecular energy transfer in the bioluminescent system of Aequorea | journal=Biochemistry | volume=13 | issue=12 | pages=2656–2662 | year=1974 | pmid=4151620 | doi=10.1021/bi00709a028}}</ref>

Overall, bioluminescence has arisen over 40 times in evolutionary history.<ref name=Haddock/> In [[evolution]], luciferins tend to vary little: one in particular, [[coelenterazine]], is the light emitting pigment for nine [[phylum|phyla]] (groups of very different organisms), including polycystine [[radiolaria]], [[Cercozoa]] ([[Phaeodaria]]), [[protozoa]], [[Ctenophora|comb jellies]], [[cnidaria]] including [[jellyfish]] and [[coral]]s, [[crustacean]]s, [[Mollusca|molluscs]], [[Chaetognatha|arrow worms]] and [[vertebrate]]s ([[ray-finned fish]]). Not all these organisms synthesise coelenterazine: some of them obtain it through their diet.<ref name=Haddock/> Conversely, luciferase enzymes vary widely and tend to be different in each species.<ref name=Haddock/>

==Distribution==
{{further |List of bioluminescent organisms}}
[[File:Surfing_on_bioluminescent_wave.webm|thumb|left |upright=1.6<!--width for very low image--> |Huge numbers of [[dinoflagellate]]s creating bioluminescence<!--don't wikilink, it comes back here--> in breaking waves]]

Bioluminescence occurs widely among animals, especially in the open sea, including [[fish]], [[jellyfish]], [[Ctenophora|comb jellies]], [[crustacean]]s, and [[cephalopod]] molluscs; in some [[List of bioluminescent fungi|fungi]] and [[bioluminescent bacteria|bacteria]]; and in various terrestrial invertebrates, nearly all of which are [[beetle]]s. In marine coastal habitats, about 2.5% of organisms are estimated to be bioluminescent, whereas in pelagic habitats in the eastern Pacific, about 76% of the main taxa of [[deep sea creature|deep-sea animals]] have been found to be capable of producing light.<ref>{{Cite journal |last1=Martini |first1=Séverine |last2=Haddock |first2=Steven H. D. |author2-link=Steven Haddock |date=April 2017 |title=Quantification of bioluminescence from the surface to the deep sea demonstrates its predominance as an ecological trait |journal=Scientific Reports |volume=7 |pages=45750 |doi=10.1038/srep45750 |pmc=5379559 |pmid=28374789 |bibcode=2017NatSR...745750M}}</ref> More than 700 animal genera have been recorded with light-producing species.<ref>{{Cite journal |last1=Kanie |first1=Shusei |last2=Miura |first2=Daisuke |last3=Jimi |first3=Naoto |last4=Hayashi |first4=Taro |last5=Nakamura |first5=Koji |last6=Sakata |first6=Masahiko |last7=Ogoh |first7=Katsunori |last8=Ohmiya |first8=Yoshihiro |last9=Mitani |first9=Yasuo |date=2021-09-27 |title=Violet bioluminescent Polycirrus sp. (Annelida: Terebelliformia) discovered in the shallow coastal waters of the Noto Peninsula in Japan |journal=Scientific Reports |language=en |volume=11 |issue=1 |pages=19097 |doi=10.1038/s41598-021-98105-6 |pmid=34580316 |pmc=8476577 |bibcode=2021NatSR..1119097K |issn=2045-2322}}</ref> Most marine light-emission is in the blue and green [[electromagnetic radiation|light spectrum]]. However, some [[Stoplight loosejaw|loose-jawed fish]] emit red and [[infrared]] light, and the genus ''[[Tomopteris]]'' emits yellow light.<ref name=Haddock/><ref>{{Cite journal |title=The Covert World of Fish Biofluorescence: A Phylogenetically Widespread and Phenotypically Variable Phenomenon |journal=PLOS ONE |date=8 January 2014 |pmc=3885428 |pmid=24421880 |pages=e83259 |volume=9 |issue=1 |doi=10.1371/journal.pone.0083259 |first1=John S. |last1=Sparks |first2=Robert C. |last2=Schelly |first3=W. Leo |last3=Smith |first4=Matthew P. |last4=Davis |first5=Dan |last5=Tchernov |first6=Vincent A. |last6=Pieribone |first7=David F. |last7=Gruber |bibcode=2014PLoSO...983259S |doi-access=free }}</ref>

The most frequently encountered bioluminescent organisms may be the [[dinoflagellate]]s in the surface layers of the sea, which are responsible for the sparkling luminescence sometimes seen at night in disturbed water. At least 18 genera of these [[phytoplankton]] exhibit luminosity.<ref name=Haddock/> Luminescent dinoflagellate ecosystems are present in warm water lagoons and bays with narrow openings to the ocean.<ref>{{Cite web |title=Bioluminescence {{!}} National Geographic Society |url=https://education.nationalgeographic.org/resource/bioluminescence/ |access-date=2023-01-28 |website=education.nationalgeographic.org}}</ref> A different effect is the thousands of square miles of the ocean which shine with the light produced by bioluminescent bacteria, known as [[Milky seas effect|mareel or the milky seas effect]].<ref>{{cite news | url=http://news.bbc.co.uk/1/hi/sci/tech/3760124.stm | title='Milky seas' detected from space | publisher=BBC | date=27 September 2005 | access-date=13 March 2013 | last=Ross |first=Alison}}</ref> 
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Non-marine bioluminescence is less widely distributed, the two best-known cases being in [[Firefly |fireflies]] and [[Phengodidae |glowworms]]. Other invertebrates including insect [[larva]]e, [[annelid]]s and [[arachnid]]s possess bioluminescent abilities. Some forms of bioluminescence are brighter (or exist only) at night, following a [[circadian rhythm]].{{cn}}-->

=== Pelagic zone ===
Bioluminescence is abundant in the pelagic zone, with the most concentration at depths devoid of light and surface waters at night. These organisms participate in diurnal vertical migration from the dark depths to the surface at night, dispersing the population of bioluminescent organisms across the pelagic water column. The dispersal of bioluminescence across different depths in the pelagic zone has been attributed to the selection pressures imposed by predation and the lack of places to hide in the open sea. In depths where sunlight never penetrates, often below 200m, the significance of bioluminescent is evident in the retainment of functional eyes for organisms to detect bioluminescence.<ref name="Widder 2002">{{Cite journal |last=Widder |first=Edith |date=January 2002 |title=Bioluminescence and the Pelagic Visual Environment |url=http://dx.doi.org/10.1080/10236240290025581 |journal=Marine and Freshwater Behaviour and Physiology |volume=35 |issue=1–2 |pages=1–26 |doi=10.1080/10236240290025581 |bibcode=2002MFBP...35....1W |s2cid=85259393 |issn=1023-6244}}</ref>

==== Bacterial symbioses ====

Organisms often produce bioluminescence themselves, rarely do they generate it from outside phenomena. However, there are occasions where bioluminescence is produced by bacterial symbionts that have a symbiotic relationship with the host organism. Although many luminous bacteria in the marine environment are free-living, a majority are found in symbiotic relationships that involve fish, squids, crustaceans etc. as hosts. Most luminous bacteria inhabit the sea, dominated by ''[[Photobacterium]]'' and ''[[Vibrio]]''.<ref>{{Cite journal |last1=Miyamoto |first1=C. |last2=Skouris |first2=N. |last3=Hosseinkhani |first3=S |last4=Lin |first4=L. Y. |last5=Meighen |first5=E. A. |date=November 2002 |title=Common Features of the Quorum Sensing Systems in Vibrio Species |url=http://dx.doi.org/10.1142/9789812776624_0021 |journal=Bioluminescence and Chemiluminescence |pages=97–100 |publisher=World Scientific |doi=10.1142/9789812776624_0021 |isbn=978-981-238-156-9}}</ref>

In the symbiotic relationship, bacterium benefit from having a source of nourishment and a refuge to grow. Hosts obtain these bacterial symbionts either from the environment, [[Spawn (biology)|spawning]], or the luminous bacterium is evolving with their host.<ref>{{Cite journal |last1=Baker |first1=Lydia J. |last2=Freed |first2=Lindsay L. |last3=Easson |first3=Cole G |last4=Lopez |first4=Jose V |last5=Fenolio |first5=Danté |last6=Sutton |first6=Tracey T. |last7=Nyholm |first7=Spencer V. |last8=Hendry |first8=Tory A |date=2019-10-01 |title=Diverse deep-sea anglerfishes share a genetically reduced luminous symbiont that is acquired from the environment |journal=eLife |language=en |volume=8 |pages=e47606 |doi=10.7554/eLife.47606 |pmid=31571583 |pmc=6773444 |issn=2050-084X |doi-access=free}}</ref> Coevolutionary interactions are suggested as host organisms' anatomical adaptations have become specific to only certain luminous bacteria, to suffice ecological dependence of bioluminescence.<ref>{{Cite journal |last1=Dunlap |first1=Paul V. |last2=Ast |first2=Jennifer C. |last3=Kimura |first3=Seishi |last4=Fukui |first4=Atsushi |last5=Yoshino |first5=Tetsuo |last6=Endo |first6=Hiromitsu |date=October 2007 |title=Phylogenetic analysis of host?symbiont specificity and codivergence in bioluminescent symbioses |url=http://dx.doi.org/10.1111/j.1096-0031.2007.00157.x |journal=Cladistics |volume=23 |issue=5 |pages=507–532 |doi=10.1111/j.1096-0031.2007.00157.x |issn=0748-3007 |hdl=2027.42/73754 |s2cid=31107773 |hdl-access=free}}</ref>

=== Benthic zone ===

Bioluminescence is widely studied amongst species located in the mesopelagic zone, but the [[benthic zone]] at mesopelagic depths has remained widely unknown. Benthic habitats at depths beyond the mesopelagic are also poorly understood due to the same constraints. Unlike the pelagic zone where the emission of light is undisturbed in the open sea, the occurrence of bioluminescence in the benthic zone is less common. It has been attributed to the blockage of emitted light by a number of sources such as the sea floor, and inorganic and organic structures. Visual signals and communication that is prevalent in the pelagic zone such as counter-illumination may not be functional or relevant in the benthic realm. Bioluminescence in bathyal benthic species still remains poorly studied due to difficulties of the collection of species at these depths.<ref name="Johnsen 2012">{{Cite journal |last1=Johnsen |first1=S. |last2=Frank |first2=T. M. |last3=Haddock |first3=S. H. D. |last4=Widder |first4=E. A. |last5=Messing |first5=C. G. |date=September 2012 |title=Light and vision in the deep-sea benthos: I. Bioluminescence at 500-1000 m depth in the Bahamian Islands |journal=Journal of Experimental Biology |volume=215 |issue=19 |pages=3335–3343 |doi=10.1242/jeb.072009 |pmid=22956246 |issn=0022-0949 |doi-access=free}}</ref>

==Uses in nature==
[[File:Mycena chlorophos.jpg|thumb|''[[Mycena chlorophos]]'', a [[List of bioluminescent fungi|bioluminescent mushroom]]]]
Bioluminescence has several functions in different taxa. [[Steven Haddock]] et al. (2010) list as more or less definite functions in marine organisms the following: defensive functions of startle, counterillumination (camouflage), misdirection (smoke screen), distractive body parts, burglar alarm (making predators easier for higher predators to see), and warning to deter settlers; offensive functions of lure, stun or confuse prey, illuminate prey, and mate attraction/recognition. It is much easier for researchers to detect that a species is able to produce light than to analyze the chemical mechanisms or to prove what function the light serves.<ref name=Haddock/> In some cases the function is unknown, as with species in three families of earthworm ([[Oligochaeta]]), such as ''[[Diplocardia longa]],'' where the [[Coelom#Coelomic fluid|coelomic fluid]] produces light when the animal moves.<ref name=Viviani/> The following functions are reasonably well established in the named organisms.

===Counterillumination camouflage===
[[File:Squid Counterillumination.png|thumb|Principle of [[counterillumination]] [[camouflage]] in firefly squid, ''[[Watasenia scintillans]]''. When seen from below by a predator, the bioluminescence helps to match the squid's brightness and color to the sea surface above.]]
{{Further |Counterillumination}}

In many animals of the deep sea, including several [[squid]] species, bacterial bioluminescence is used for [[camouflage]] by [[counterillumination]], in which the animal matches the overhead environmental light as seen from below.<ref name="Young">{{Cite journal | doi=10.1126/science.1251214 | last1=Young | first1=R. E. | last2=Roper | first2=C. F. | s2cid=11284796 | year=1976 | title=Bioluminescent countershading in midwater animals: evidence from living squid | journal=Science | volume=191 | issue=4231 | pages=1046–8 | pmid=1251214 |bibcode=1976Sci...191.1046Y}}</ref> In these animals, photoreceptors control the illumination to match the brightness of the background.<ref name="Young"/> These light organs are usually separate from the tissue containing the bioluminescent bacteria. However, in one species, ''[[Euprymna scolopes]]'', the bacteria are an integral component of the animal's light organ.<ref>{{cite journal | last1=Tong | first1=D. | last2=Rozas | first2=N. S. | last3=Oakley | first3=T. H. | last4=Mitchell | first4=J. | last5=Colley | first5=N. J. | last6=McFall-Ngai | first6=M.J. | title=Evidence for light perception in a bioluminescent organ | journal=Proceedings of the National Academy of Sciences of the United States of America | volume=106 | issue=24 | pages=9836–41 | year=2009 | pmid=19509343 | pmc=2700988 | doi=10.1073/pnas.0904571106 | bibcode=2009PNAS..106.9836T| doi-access=free }}</ref>

===Attraction===
[[File:Stauroteuthis.jpg|thumb|upright|''Stauroteuthis syrtensis'' bioluminescent photophores]]

Bioluminescence is used in a variety of ways and for different purposes. The cirrate octopod ''Stauroteuthis syrtensis'' uses emits bioluminescence from its sucker like structures.<ref>{{Cite journal |last1=Johnsen |first1=S. |last2=Balser |first2=E. J. |last3=Fisher |first3=E. C. |last4=Widder |first4=E. A. |date=1999-08-01 |title=Bioluminescence in the Deep-Sea Cirrate Octopod Stauroteuthis syrtensis Verrill (Mollusca: Cephalopoda) |url=https://www.journals.uchicago.edu/doi/10.2307/1542994 |journal=The Biological Bulletin |volume=197 |issue=1 |pages=26–39 |doi=10.2307/1542994 |jstor=1542994 |pmid=28296499 |issn=0006-3185}}</ref> These structures are believed to have evolved from what are more commonly known as octopus suckers. They do not have the same function as the normal suckers because they no longer have any handling or grappling ability due its evolution of [[photophore]]s. The placement of the photophores are within the animals oral reach, which leads researchers to suggest that it uses it bioluminescence to capture and lure prey.<ref name="Haddock 2009">{{Cite journal |last1=Haddock |first1=Steven H.D. |last2=Moline |first2=Mark A. |last3=Case |first3=James F. |date=2009-12-14 |title=Bioluminescence in the Sea |url=https://www.annualreviews.org/doi/10.1146/annurev-marine-120308-081028 |journal=Annual Review of Marine Science |volume=2 |issue=1 |pages=443–493 |doi=10.1146/annurev-marine-120308-081028 |pmid=21141672 |bibcode=2010ARMS....2..443H |issn=1941-1405}}</ref>

[[Firefly|Fireflies]] use light to attract [[mating|mates]]. Two systems are involved according to species; in one, females emit light from their abdomens to attract males; in the other, flying males emit signals to which the sometimes sedentary females respond.<ref name=Viviani/><ref>{{cite journal | last1=Stanger-Hall | first1=K. F. | last2=Lloyd | first2=J.E. | last3=Hillis | first3=D.M. | title=Phylogeny of North American fireflies (Coleoptera: Lampyridae): implications for the evolution of light signals | journal=Molecular Phylogenetics and Evolution |volume=45 |issue=1 |pages=33–49 |doi=10.1016/j.ympev.2007.05.013 | year=2007 | pmid=17644427| bibcode=2007MolPE..45...33S }}</ref> [[Click beetle]]s emit an orange light from the abdomen when flying and a green light from the thorax when they are disturbed or moving about on the ground. The former is probably a sexual attractant but the latter may be defensive.<ref name=Viviani/> Larvae of the click beetle ''[[Pyrophorus nyctophanus]]'' live in the surface layers of termite mounds in Brazil. They light up the mounds by emitting a bright greenish glow which attracts the flying insects on which they feed.<ref name=Viviani/>

In the marine environment, use of luminescence for mate attraction is chiefly known among [[ostracod]]s, small shrimp-like [[crustacean]]s, especially in the family [[Cyprididae]]. [[Pheromone]]s may be used for long-distance communication, with bioluminescence used at close range to enable mates to "home in".<ref name=Haddock/> A [[polychaete]] worm, the [[Odontosyllis enopla|Bermuda fireworm]] creates a brief display, a few nights after the full moon, when the female lights up to attract males.<ref>{{cite book |last=Shimomura |first=Osamu |author-link=Osamu Shimomura |title=Bioluminescence: Chemical Principles and Methods |url=https://books.google.com/books?id=pGyto9ctbVoC&pg=PA234 |year=2012 |publisher=World Scientific |isbn=978-981-4366-08-3 |page=234}}</ref>

===Defense===

[[File:The depths of the ocean; a general account of the modern science of oceanography based largely on the scientific researches of the Norwegian steamer Michael Sars in the North Atlantic (1912) (20674653149).jpg|thumb|upright|''Acanthephyra purpurea'' has photophores along its body which it uses in defense against predators.]]

The defense mechanisms for bioluminescent organisms can come in multiple forms; startling prey, counter-illumination, smoke screen or misdirection, distractive body parts, burglar alarm, sacrificial tag or warning coloration. The shrimp family Oplophoridae Dana use their bioluminescence as a way of startling the predator that is after them.<ref name="sciencedirect.com">{{cite journal |last1=Wong |first1=Juliet M. |last2=Pérez-Moreno |first2=Jorge L. |last3=Chan |first3=Tin-Yam |last4=Frank |first4=Tamara M. |last5=Bracken-Grissom |first5=Heather D. |date=2015-02-01 |title=Phylogenetic and transcriptomic analyses reveal the evolution of bioluminescence and light detection in marine deep-sea shrimps of the family Oplophoridae (Crustacea: Decapoda) |url=http://www.sciencedirect.com/science/article/pii/S1055790314004151 |journal=Molecular Phylogenetics and Evolution |language=en |volume=83 |pages=278–292 |doi=10.1016/j.ympev.2014.11.013 |pmid=25482362 |bibcode=2015MolPE..83..278W |issn=1055-7903}}</ref> ''Acanthephyra purpurea'', within the Oplophoridae family, uses its photophores to emit light, and can secrete a bioluminescent substance when in the presence of a predator. This secretory mechanism is common among prey fish.<ref name="sciencedirect.com"/>

Many [[cephalopod]]s, including at least 70 [[genus (biology)|genera]] of [[squid]], are bioluminescent.<ref name=Haddock/> Some squid and small [[crustacean]]s use bioluminescent chemical mixtures or bacterial slurries in the same way as many squid use [[Cephalopod ink|ink]]. A cloud of luminescent material is expelled, distracting or repelling a potential predator, while the animal escapes to safety.<ref name=Haddock>{{cite journal |last1=Haddock |first1=Steven H.D. |author-link=Steven Haddock |last2=Moline |first2=Mark A. |last3=Case |first3=James F. |s2cid=3872860 |title=Bioluminescence in the Sea |journal=Annual Review of Marine Science |date=2010 |volume=2 |pages=443–493 |doi=10.1146/annurev-marine-120308-081028 |pmid=21141672 |bibcode=2010ARMS....2..443H }}</ref> The deep sea squid ''[[Octopoteuthis deletron]]'' may [[Autotomy|autotomize]] portions of its arms which are luminous and continue to twitch and flash, thus distracting a predator while the animal flees.<ref name=Haddock/>

[[Dinoflagellate]]s may use bioluminescence for defense against [[predator]]s. They shine when they detect a predator, possibly making the predator itself more vulnerable by attracting the attention of predators from higher trophic levels.<ref name=Haddock/> Grazing copepods release any phytoplankton cells that flash, unharmed; if they were eaten they would make the copepods glow, attracting predators, so the phytoplankton's bioluminescence is defensive. The problem of shining stomach contents is solved (and the explanation corroborated) in predatory deep-sea fishes: their stomachs have a black lining able to keep the light from any bioluminescent fish prey which they have swallowed from attracting larger predators.<ref name=Pauly2004/>

The [[Vargula hilgendorfii|sea-firefly]] is a small crustacean living in sediment. At rest it emits a dull glow but when disturbed it darts away leaving a cloud of shimmering blue light to confuse the predator. During World War II it was gathered and dried for use by the Japanese army as a source of light during clandestine operations.<ref name=Pieribone>{{cite book |author1=Pieribone, Vincent |author2=Gruber, David F. |title=Aglow in the Dark: The Revolutionary Science of Biofluorescence |url=https://archive.org/details/aglowindarkrevol00pier |url-access=registration |year=2005 |publisher=Harvard University Press |isbn=978-0-674-01921-8 |pages=[https://archive.org/details/aglowindarkrevol00pier/page/35 35]–41}}</ref>

The larvae of [[railroad worm]]s (''Phrixothrix'') have paired photic organs on each body segment, able to glow with green light; these are thought to have a defensive purpose.<ref>{{cite web |url=http://entnemdept.ufl.edu/creatures/misc/beetles/glow-worms.htm |title=Glow-worms, railroad-worms (Insecta: Coleoptera: Phengodidae) |last=Branham |first=Marc |work=Featured Creatures |publisher=University of Florida |access-date=29 November 2014}}</ref> They also have organs on the head which produce red light; they are the only terrestrial organisms to emit light of this color.<ref>{{cite journal |last=Viviani |first=Vadim R. |author2=Bechara, Etelvino J.H. |year=1997 |title=Bioluminescence and Biological Aspects of Brazilian Railroad-Worms (Coleoptera: Phengodidae) |journal=Annals of the Entomological Society of America |volume=90 |issue=3 |pages=389–398 |url=https://www.researchgate.net/publication/233631656 |doi=10.1093/aesa/90.3.389}}</ref>

===Warning===
<!--[[File:Glowing millipede.jpg|thumb|Some [[millipede]]s glow to warn off predators. Model of ''Motyxia sequoiae'' at the American Museum of Natural History.]]-->
[[Aposematism]] is a widely used function of bioluminescence, providing a warning that the creature concerned is unpalatable. It is suggested that many firefly larvae glow to repel predators; some [[millipede]]s glow for the same purpose.<ref>{{cite journal |author1=Marek, Paul |author2=Papaj, Daniel |author3=Yeager, Justin |author4=Molina, Sergio |author5=Moore, Wendy |title=Bioluminescent aposematism in millipedes | journal=Current Biology | doi=10.1016/j.cub.2011.08.012 | volume=21 | issue=18 | pages=R680–R681 | pmid=21959150 | pmc=3221455 |year=2011|bibcode=2011CBio...21.R680M }}</ref> Some marine organisms are believed to emit light for a similar reason. These include [[Polynoidae|scale worms]], [[jellyfish]] and [[brittle star]]s but further research is needed to fully establish the function of the luminescence. Such a mechanism would be of particular advantage to soft-bodied [[cnidaria]]ns if they were able to deter predation in this way.<ref name=Haddock/> The [[limpet]] ''[[Latia neritoides]]'' is the only known freshwater [[Gastropoda|gastropod]] that emits light. It produces greenish luminescent [[mucus]] which may have an anti-predator function.<ref>{{cite journal |last=Meyer-Rochow |first=V. B. |author2=Moore, S. |year=1988 |title=Biology of ''Latia neritoides'' Gray 1850 (Gastropoda, Pulmonata, Basommatophora): the Only Light-producing Freshwater Snail in the World |journal=Internationale Revue der Gesamten Hydrobiologie und Hydrographie |volume=73 |issue=1 |pages=21–42 |doi=10.1002/iroh.19880730104 }}</ref> The marine snail ''[[Hinea brasiliana]]'' uses flashes of light, probably to deter predators. The blue-green light is emitted through the translucent shell, which functions as an efficient diffuser of light.<ref name=Deheyn>{{cite journal |title=Bioluminescent signals spatially amplified by wavelength-specific diffusion through the shell of a marine snail |last1=Deheyn |first1=Dimitri D. |last2=Wilson |first2=Nerida G. |journal=Proceedings of the Royal Society |year=2010 |doi=10.1098/rspb.2010.2203 |pmid=21159673 |volume=278 |issue=1715 |pages=2112–2121 |pmc=3107627 }}</ref>

===Communication===
[[File:Tunicate off Atauro island.jpg|thumb|''[[Pyrosoma]]'', a colonial [[Tunicata|tunicate]]; each individual zooid in the colony flashes a blue-green light.]]
Communication in the form of [[quorum sensing]] plays a role in the regulation of luminescence in many species of bacteria. Small extracellularly secreted molecules stimulate the bacteria to turn on genes for light production when cell density, measured by concentration of the secreted molecules, is high.<ref name=Haddock/>

[[Pyrosome]]s are colonial [[Tunicata|tunicates]] and each [[zooid]] has a pair of luminescent organs on either side of the inlet siphon. When stimulated by light, these turn on and off, causing rhythmic flashing. No neural pathway runs between the zooids, but each responds to the light produced by other individuals, and even to light from other nearby colonies.<ref>{{cite journal |last1=Bowlby |first1=Mark R. |last2=Widder |first2=Edith |last3=Case |first3=James |year=1990 |title=Patterns of stimulated bioluminescence in two pyrosomes (Tunicata: Pyrosomatidae) |journal=Biological Bulletin |volume=179 |issue=3 |pages=340–350 |jstor=1542326 |doi=10.2307/1542326 |pmid=29314963 |s2cid=42380533 |url=https://www.biodiversitylibrary.org/part/24992 }}</ref> Communication by light emission between the zooids enables coordination of colony effort, for example in swimming where each zooid provides part of the propulsive force.<ref>{{cite book |title=Encyclopedia of the Aquatic World |url=https://books.google.com/books?id=8UfQr_CqFOUC&pg=PA1115 |date=January 2004 |publisher=Marshall Cavendish |isbn=978-0-7614-7418-0 |page=1115}}</ref>

Some bioluminous bacteria infect [[nematode]]s that parasitize [[Lepidoptera]] larvae. When these [[caterpillar]]s die, their luminosity may attract predators to the dead insect thus assisting in the dispersal of both bacteria and nematodes.<ref name=Viviani>{{cite web |url=http://www.photobiology.info/Viviani.html |title=Terrestrial bioluminescence |last=Viviani |first=Vadim |date=17 February 2009 |access-date=26 November 2014}}</ref> A similar reason may account for the many species of fungi that emit light. Species in the genera ''[[Armillaria]]'', ''[[Mycena]]'', ''[[Omphalotus]]'', ''[[Panellus]]'', ''[[Pleurotus]]'' and others do this, emitting usually greenish light from the [[mycelium]], [[Pileus (mycology)|cap]] and [[Lamella (mycology)|gills]]. This may attract night-flying insects and aid in spore dispersal, but other functions may also be involved.<ref name=Viviani/>

''[[Quantula striata]]'' is the only known bioluminescent terrestrial mollusk. Pulses of light are emitted from a gland near the front of the foot and may have a communicative function, although the adaptive significance is not fully understood.<ref>{{cite journal |last1=Copeland |first1=J. |last2=Daston |first2=M. M. |year=1989 |title=Bioluminescence in the terrestrial snail ''Quantula (Dyakia) striata'' |journal=Malacologia |volume=30 |issue=1–2 |pages=317–324 |url=https://archive.org/stream/malacologia301989inst#page/316/mode/2up }}</ref>

===Mimicry===
[[File:Bufoceratias.jpg|thumb|A [[deep sea fish|deep sea]] [[anglerfish]], ''[[Bufoceratias wedli]]'', showing the [[Fin ray|esca]] (lure)]]

Bioluminescence is used by a variety of animals to [[mimicry|mimic]] other species. Many species of [[deep sea fish]] such as the [[anglerfish]] and [[Stomiidae|dragonfish]] make use of [[aggressive mimicry]] to attract [[prey]]. They have an [[appendage]] on their heads called an [[Fin ray|esca]] that contains bioluminescent bacteria able to produce a long-lasting glow which the fish can control. The glowing esca is dangled or waved about to lure small animals to within striking distance of the fish.<ref name=Haddock/><ref>{{cite journal | url=http://www.ingentaconnect.com/content/umrsmas/bullmar/1983/00000033/00000004/art00003 | title=Oceanic Bioluminescence: an Overview of General Functions | last=Young |first=Richard Edward | journal=Bulletin of Marine Science |date=October 1983 | volume=33 | issue=4 | pages=829–845}}</ref>

The [[cookiecutter shark]] uses bioluminescence to camouflage its underside by counter-illumination, but a small patch near its pectoral fins remains dark, appearing as a small fish to large predatory fish like [[tuna]] and [[mackerel]] swimming beneath it. When such fish approach the lure, they are bitten by the shark.<ref>{{cite web | url=http://www.elasmo-research.org/education/ecology/deepsea-cookiecutter.htm | title=Biology of Sharks and Rays: Cookiecutter Shark | publisher=ReefQuest Centre for Shark Research | access-date=13 March 2013 | last=Martin |first=R. Aidan}}</ref><ref>{{cite web | url=http://www.sciencenews.org/pages/sn_arc98/8_1_98/fob5.htm | title=Glow-in-the-dark shark has killer smudge | publisher=Science News | date=1 August 1998 | access-date=13 March 2013 | last=Milius | first=S. | archive-date=7 December 2009 | archive-url=https://web.archive.org/web/20091207210338/http://www.sciencenews.org/pages/sn_arc98/8_1_98/fob5.htm | url-status=dead }}</ref>

Female ''[[Photuris]]'' fireflies sometimes mimic the light pattern of another firefly, ''Photinus'', to attract its males as prey. In this way they obtain both food and the defensive chemicals named [[lucibufagin]]s, which ''Photuris'' cannot synthesize.<ref>{{cite journal |last=Eisner |first=Thomas |author-link=Thomas Eisner |author2=Goetz, Michael A. |author3=Hill, David E. |author4=Smedley, Scott R. |author5=Meinwald, Jarrold | year=1997 | title=Firefly "femmes fatales" acquire defensive steroids (lucibufagins) from their firefly prey | journal=Proceedings of the National Academy of Sciences of the United States of America | volume=94 | issue=18 | pages=9723–9728 | doi=10.1073/pnas.94.18.9723 |pmid=9275191 |bibcode=1997PNAS...94.9723E |pmc=23257 |doi-access=free }}</ref>

South American giant cockroaches of the genus ''[[Lucihormetica]]'' were believed to be the first known example of defensive mimicry, emitting light in imitation of bioluminescent, poisonous click beetles.<ref>{{cite web |url=http://www.abc.net.au/science/photos/2014/07/16/4032762.htm?xml=4032762.mediarss.xml#bigpicturepos |title=Out of the darkness |last=Sullivan |first=Rachel |date=16 July 2014 |publisher=ABC Science |access-date=17 December 2014}}</ref> However, doubt has been cast on this assertion, and there is no conclusive evidence that the cockroaches are bioluminescent.<ref>{{cite journal |last=Greven |first=Hartmut |author1-link=Hartmut Greven |author2=Zwanzig, Nadine |year=2013 |title=Courtship, Mating, and Organisation of the Pronotum in the Glowspot Cockroach ''Lucihormetica verrucosa'' (Brunner von Wattenwyl, 1865) (Blattodea: Blaberidae) |journal=Entomologie Heute |volume=25 |pages=77–97 |url=https://www.researchgate.net/publication/258845997 }}</ref><ref>{{cite journal |author=Merritt, David J. |year=2013 |title=Standards of evidence for bioluminescence in cockroaches |journal=Naturwissenschaften |volume=100 |issue=7 |pages=697–698 |url=https://www.researchgate.net/publication/237060004 |doi=10.1007/s00114-013-1067-9 |bibcode=2013NW....100..697M |pmid=23740173 |s2cid=33240197 }}</ref>

[[File:Malacosteus niger cam.jpg|thumb|upright=1.3 |Flashing of photophores of black dragonfish, ''[[Malacosteus niger]]'', showing red fluorescence]]

=== Illumination ===

While most marine bioluminescence is green to blue, some deep sea [[Stomiidae|barbeled dragonfishes]] in the genera ''[[Aristostomias]]'', ''[[Pachystomias]]'' and ''[[Malacosteus]]''<!--Not a list: do not add more examples here, thanks--> emit a red glow. This adaptation allows the fish to see red-pigmented prey, which are normally invisible to other organisms in the deep ocean environment where red light has been filtered out by the water column.<ref name="Herring-2005">{{Cite journal |last1=Herring |first1=Peter J. |last2=Cope |first2=Celia |date=December 2005 |title=Red bioluminescence in fishes: on the suborbital photophores of Malacosteus, Pachystomias and Aristostomias |url=http://link.springer.com/10.1007/s00227-005-0085-3 |journal=Marine Biology |volume=148 |issue=2 |pages=383–394 |doi=10.1007/s00227-005-0085-3 |bibcode=2005MarBi.148..383H |s2cid=86463272 |issn=0025-3162}}</ref> These fish are able to utilize the longer wavelength to act as a spotlight for its prey that only they can see.<ref name="Herring-2005"/> The fish may also use this light to communicate with each other to find potential mates.<ref>{{Cite journal |last1=Widder |first1=Edith A. |last2=Latz |first2=Michael I. |last3=Herring |first3=Peter J. |last4=Case |first4=James F. |date=1984-08-03 |title=Far Red Bioluminescence from Two Deep-Sea Fishes |url=https://www.science.org/doi/10.1126/science.225.4661.512 |journal=Science |language=en |volume=225 |issue=4661 |pages=512–514 |doi=10.1126/science.225.4661.512 |pmid=17750854 |bibcode=1984Sci...225..512W |s2cid=31510972 |issn=0036-8075}}</ref> The ability of the fish to see this light is explained by the presence of specialized rhodopsin pigment.<ref name="Herring-2005" /> The mechanism of light creation is through a suborbital photophore that utilizes gland cells which produce exergonic chemical reactions that produce light with a longer, red wavelength.<ref name="Haddock-2005" /> The dragonfish species which produce the red light also produce blue light in photophore on the dorsal area.<ref name="Haddock-2005" /> The main function of this is to alert the fish to the presence of its prey.<ref name="Bone-2008">{{Cite book |last=Bone |first=Q. |url=https://www.worldcat.org/oclc/244632464 |title=Biology of fishes |date=2008 |publisher=Taylor & Francis |others=Richard H. Moore |isbn=978-0-203-88522-2 |edition=3rd |location=New York |oclc=244632464}}</ref> The additional pigment is thought to be assimilated from [[chlorophyll]] derivatives found in the [[copepod]]s which form part of its diet.<ref name="Bone-2008" />

The angler siphonophore (''[[Erenna]]'') utilizes red bioluminescence in appendages to lure fish.<ref name="Haddock-2005">{{Cite journal |last1=Haddock |first1=Steven H. D. |last2=Dunn |first2=Casey W. |last3=Pugh |first3=Philip R. |last4=Schnitzler |first4=Christine E. |date=2005-07-08 |title=Bioluminescent and Red-Fluorescent Lures in a Deep-Sea Siphonophore |url=https://www.science.org/doi/10.1126/science.1110441 |journal=Science |language=en |volume=309 |issue=5732 |pages=263 |doi=10.1126/science.1110441 |pmid=16002609 |s2cid=29284690 |issn=0036-8075}}</ref>

==Biotechnology==

===Biology and medicine===
{{further|optogenetics}}

Bioluminescent organisms are a target for many areas of research. Luciferase systems are widely used in [[genetic engineering]] as [[reporter gene]]s, each producing a different color by fluorescence,<ref name="reporter">{{cite journal |last=Koo |first=J. |author2=Kim, Y. |author3=Kim, J. |author4=Yeom, M. |author5=Lee, I. C. |author6=Nam, H. G. | date=2007 | title=A GUS/Luciferase Fusion Reporter for Plant Gene Trapping and for Assay of Promoter Activity with Luciferin-Dependent Control of the Reporter Protein Stability | journal=Plant and Cell Physiology | volume=48 | issue=8 | pages=1121–31 | doi=10.1093/pcp/pcm081 | pmid=17597079 | doi-access=free }}</ref><ref>{{cite journal |last=Nordgren |first=I. K. |author2=Tavassoli, A. | date=2014 | title=A bidirectional fluorescent two-hybrid system for monitoring protein-protein interactions | journal=Molecular BioSystems | volume=10 | issue=3 | pages=485–490 | doi=10.1039/c3mb70438f | pmid=24382456 | url=http://pubs.rsc.org/en/content/articlepdf/2014/mb/c3mb70438f | doi-access=free }}</ref> and for biomedical research using [[bioluminescence imaging]].<ref>{{cite journal |pages=380–7 |doi=10.1128/AAC.49.1.380-387.2005 |title=Real-Time in Vivo Bioluminescent Imaging for Evaluating the Efficacy of Antibiotics in a Rat Staphylococcus aureus Endocarditis Model |year=2004 |last1=Xiong |first1=Yan Q. |last2=Willard |first2=Julie |last3=Kadurugamuwa |first3=Jagath L. |last4=Yu |first4=Jun |last5=Francis |first5=Kevin P. |last6=Bayer |first6=Arnold S. |journal=Antimicrobial Agents and Chemotherapy |volume=49 |pmid=15616318 |issue=1 |pmc=538900}}</ref><ref>{{cite journal |last=Di Rocco |first=Giuliana |author2=Gentile, Antonietta |author3=Antonini, Annalisa |author4=Truffa, Silvia |author5=Piaggio, Giulia |author6=Capogrossi, Maurizio C. |author7=Toietta, Gabriele |title=Analysis of biodistribution and engraftment into the liver of genetically modified mesenchymal stromal cells derived from adipose tissue |journal=Cell Transplantation |date=1 September 2012 |volume=21 |issue=9 |pages=1997–2008 |doi=10.3727/096368911X637452 |pmid=22469297 |s2cid=21603693 |url=http://opendepot.org/2024/1/DiRocco2012.pdf |doi-access=free }}{{Dead link|date=October 2023 |bot=InternetArchiveBot |fix-attempted=yes }}</ref><ref>{{cite journal |pages=2445–51 |doi=10.1096/fj.07-103713 |url=http://www.newswise.com/articles/view/541186/ |title=Antivascular effects of combretastatin A4 phosphate in breast cancer xenograft assessed using dynamic bioluminescence imaging and confirmed by MRI |year=2008 |last1=Zhao |first1=Dawen |last2=Richer |first2=Edmond |last3=Antich |first3=Peter P. |last4=Mason |first4=Ralph P. |journal=The FASEB Journal |volume=22 |issue=7 |doi-access=free |pmid=18263704 |pmc=4426986 }}</ref> For example, the firefly luciferase gene was used as early as 1986 for research using transgenic tobacco plants.<ref>{{cite journal | title=Transient and stable expression of the firefly luciferase gene in plant cells and transgenic plants |last=Ow |first=D. W. |author2=Wood, K. V. |author3=DeLuca, M. |author4=de Wet, J. R. |author5=Helinski, D. R. |author6=Howell, S.H. | journal=Science | volume=234 | issue=4778 | pages=856–859 | issn=0036-8075 | year=1986 | publisher=American Association for the Advancement of Science | doi=10.1126/science.234.4778.856 |pmid=17758108 |bibcode=1986Sci...234..856O |s2cid=32603977 }}</ref> ''Vibrio'' bacteria symbiose with [[marine invertebrates]] such as the [[Euprymna scolopes|Hawaiian bobtail squid]] (''Euprymna scolopes''), are key [[model organism|experimental models]] for bioluminescence.<ref>{{cite journal |last=Altura |first=M.A. |author2=Heath-Heckman, E.A. |author3=Gillette, A. |author4=Kremer, N. |author5=Krachler, A.M. |author6=Brennan, C. |author7=Ruby, E.G. |author8=Orth, K. |author9=McFall-Ngai, M.J. | title=The first engagement of partners in the Euprymna scolopes-Vibrio fischeri symbiosis is a two-step process initiated by a few environmental symbiont cells | journal=Environmental Microbiology | date=2013 | doi=10.1111/1462-2920.12179 | pmid=23819708 | pmc=3937295 | volume=15 |issue=11 | pages=2937–50|bibcode=2013EnvMi..15.2937A }}</ref><ref>{{cite web |title=Comprehensive Squid-Vibrio Publications List |url=http://labs.medmicro.wisc.edu/mcfall-ngai/allpublications.html |archive-url=https://web.archive.org/web/20141019165545/http://labs.medmicro.wisc.edu/mcfall-ngai/allpublications.html |url-status=dead |archive-date=19 October 2014 |publisher=University of Wisconsin-Madison}}</ref> [[Bioluminescent activated destruction]] is an experimental cancer treatment.<ref>{{cite web |author=Ludwig Institute for Cancer Research |title=Firefly Light Helps Destroy Cancer Cells; Researchers Find That The Bioluminescence Effects of Fireflies May Kill Cancer Cells From Within |url=https://www.sciencedaily.com/releases/2003/04/030421084227.htm |publisher=Science Daily |access-date=4 December 2014 |date=21 April 2003}}</ref>

''In Vivo'' luminescence cell and animal imaging uses dyes and fluorescent proteins as [[chromophore]]s. The characteristics of each chromophore determine which cell area(s) will be targeted and illuminated.<ref>{{cite book|first1=Jorge H.S.K. |last1=Monteiro|first2=Josiane A.|last2=Sobrinho|first3=Ana|last3=de Bettancourt-Dias|title=Metal Ions in Bio-Imaging Techniques|publisher=Springer|year=2021|pages=371–401 |chapter=Chapter 13. Luminescence Imaging of Cancer Cells|doi=10.1515/9783110685701-019|s2cid=233628296}}</ref>

===Light production===

[[File:Firefly_petunia.jpg|thumb|upright|A "Firefly" [[petunia]], genetically engineered to produce luciferase.]]

The structures of [[photophore]]s, the light producing organs in bioluminescent organisms, are being investigated by [[industrial design]]ers. Engineered bioluminescence could perhaps one day be used to reduce the need for street lighting, or for decorative purposes if it becomes possible to produce light that is both bright enough and can be sustained for long periods at a workable price.<ref name=Economist>{{cite news |title=How illuminating |url=http://www.economist.com/node/18304146 |newspaper=The Economist | date=10 March 2011 |access-date=6 December 2014}}</ref><ref name=QandA>[http://siobiolum.ucsd.edu/Biolum_q&a.html Bioluminescence Questions and Answers]. Siobiolum.ucsd.edu. Retrieved on 20 October 2011.</ref><ref>(4 May 2013) [https://www.newscientist.com/article/mg21829156.500-one-per-cent.html One Per Cent: Grow your own living lights] The New Scientist, Issue 2915, Retrieved 7 May 2013</ref> The gene that makes the tails of [[Firefly|fireflies]] glow has been added to mustard plants. The plants glow faintly for an hour when touched, but a sensitive camera is needed to see the glow.<ref>[http://news.bbc.co.uk/2/hi/science/nature/751069.stm Dr. Chris Riley, "Glowing plants reveal touch sensitivity", BBC 17 May 2000.]</ref> [[University of Wisconsin–Madison]] is researching the use of genetically engineered bioluminescent [[Escherichia coli|E. coli]] bacteria, for use as [[bioluminescent bacteria]] in a [[light bulb]].<ref>{{cite news |url=http://news.discovery.com/tech/alternative-power-sources/bacteria-powered-light-bulb-is-electricity-free-130815.htm |title=Bacteria-Powered Light Bulb Is Electricity-Free |date=15 August 2013 |last=Halverson |first=Nic |access-date=29 September 2013 |archive-date=25 May 2016 |archive-url=https://web.archive.org/web/20160525132633/http://news.discovery.com/tech/alternative-power-sources/bacteria-powered-light-bulb-is-electricity-free-130815.htm |url-status=dead }}</ref> In 2011, [[Philips]] launched a microbial system for ambience lighting in the home.<ref>{{cite web |last=Swaminathan |first=Miep |url=http://www.philips.com/a-w/about/news/archive/design/news/press/2011/Philips_launches%20_Microbial_Home_new_forward_looking_design_concepts.html |title=Philips launches 'Microbial Home' new forward looking design concepts |access-date=8 May 2017 }}{{dead link |date=January 2018 |bot=InternetArchiveBot |fix-attempted=yes }}</ref><ref>{{cite web |last=Cha |first=Bonnie |url=https://www.cnet.com/news/philips-bio-light-creates-mood-lighting-with-bacteria/ |title=Philips Bio-light creates mood lighting with bacteria |date=28 November 2011}}</ref> An [[International Genetically Engineered Machine|iGEM]] team from Cambridge (England) has started to address the problem that luciferin is consumed in the light-producing reaction by developing a genetic biotechnology part that codes for a luciferin regenerating enzyme from the North American firefly.<ref>{{cite web |title=E.glowli Cambridge: Parts submitted |url=http://2010.igem.org/Team:Cambridge/BioBricks |publisher=iGEM |access-date=6 December 2014}}</ref> In 2016, Glowee, a French company started selling bioluminescent lights for shop fronts and street signs,<ref name="Glowee">{{cite news |last=Marcellin |first=Frances |title=Glow-in-the-dark bacterial lights could illuminate shop windows 2016 |work=[[New Scientist]] |date=26 February 2016}}</ref> for use between 1 and 7 in the morning when the law forbids use of electricity for this purpose.<ref name="EDF">{{cite web |title=Glowee: A vision of night-time lighting |url=http://pulse.edf.com/en/glowee-a-vision-of-night-time-lighting |publisher=Electricite de France |access-date=4 March 2016 |date=2015 |archive-date=10 March 2016 |archive-url=https://web.archive.org/web/20160310001140/http://pulse.edf.com/en/glowee-a-vision-of-night-time-lighting |url-status=dead }}</ref><ref name="Glowee New Scientist">{{cite news |url=https://www.newscientist.com/article/2078921-glow-in-the-dark-bacterial-lights-could-illuminate-shop-windows/ |title=Glow-in-the-dark bacterial lights could illuminate shop windows |date=26 February 2016 |work=New Scientist}}</ref> They used the bioluminescent bacterium ''[[Aliivibrio fischeri]]'', but the maximum lifetime of their product was three days.<ref name="Glowee"/> In April 2020, plants were genetically engineered to glow more brightly using genes from the bioluminescent mushroom ''[[Neonothopanus nambi]]'' to convert [[caffeic acid]] into luciferin.<ref name="Glowee New Scientist"/><ref>{{cite journal |last1=Mitiouchkina |first1=Tatiana |last2=Mishin |first2=Alexander S. |last3=Somermeyer |first3=Louisa Gonzalez |last4=Markina |first4=Nadezhda M. |last5=Chepurnyh |first5=Tatiana V. |last6=Guglya |first6=Elena B. |last7=Karataeva |first7=Tatiana A. |last8=Palkina |first8=Kseniia A. |last9=Shakhova |first9=Ekaterina S. |last10=Fakhranurova |first10=Liliia I. |last11=Chekova |first11=Sofia V. |last12=Tsarkova |first12=Aleksandra S. |last13=Golubev |first13=Yaroslav V. |last14=Negrebetsky |first14=Vadim V. |last15=Dolgushin |first15=Sergey A. |last16=Shalaev |first16=Pavel V. |last17=Shlykov |first17=Dmitry |last18=Melnik |first18=Olesya A. |last19=Shipunova |first19=Victoria O. |last20=Deyev |first20=Sergey M. |last21=Bubyrev |first21=Andrey I. |last22=Pushin |first22=Alexander S. |last23=Choob |first23=Vladimir V. |last24=Dolgov |first24=Sergey V. |last25=Kondrashov |first25=Fyodor A. |last26=Yampolsky |first26=Ilia V. |last27=Sarkisyan |first27=Karen S. |title=Plants with genetically encoded autoluminescence |journal=Nature Biotechnology |date=27 April 2020 |volume=38 |issue=8 |pages=944–946 |doi=10.1038/s41587-020-0500-9 |pmid=32341562 |s2cid=216559981 |pmc=7610436 }}</ref>
Another possible application is to replace chemiluminescence with bioluminescent enzymes. A Canadian company, Lux Bio, is developing long-duration bioluminescent enzymes for this purpose.<ref>https://thetyee.ca/WhatWorks/2024/05/21/Glow-Stick-Uses-Nature-Bioluminescence/</ref><ref>http://www.luxbio.ca</ref>

=== ATP bioluminescence ===

ATP bioluminescence is the process in which ATP is used to generate luminescence in an organism, in conjunction with other compounds such as luciferin. It proves to be a very good [[biosensor]] to test for the presence of living [[microbe]]s in water.<ref name="Arroyo-2017">{{Cite journal |last1=Arroyo |first1=Máira Gazzola |last2=Ferreira |first2=Adriano Menis |last3=Frota |first3=Oleci Pereira |last4=Rigotti |first4=Marcelo Alessandro |last5=de Andrade |first5=Denise |last6=Brizzotti |first6=Natalia Seron |last7=Peresi |first7=Jacqueline Tanury Macruz |last8=Castilho |first8=Elza Maria |last9=de Almeida |first9=Margarete Teresa Gottardo |display-authors=6 |date=2017-06-30 |title=Effectiveness of ATP bioluminescence assay for presumptive identification of microorganisms in hospital water sources |journal=BMC Infectious Diseases |volume=17 |issue=1 |page=458 |doi=10.1186/s12879-017-2562-y |pmid=28666419 |pmc=5493863 |issn=1471-2334 |doi-access=free }}</ref><ref>{{Cite journal |last1=Eed |first1=Heba Ramadan |last2=Abdel-Kader |first2=Nora S. |last3=El Tahan |first3=Mahmoud Helmy |last4=Dai |first4=Tianhong |last5=Amin |first5=Rehab |date=2016 |title=Bioluminescence-Sensing Assay for Microbial Growth Recognition |journal=Journal of Sensors |volume=2016 |pages=1–5 |doi=10.1155/2016/1492467 |issn=1687-725X |doi-access=free }}</ref> Different types of microbial populations are determined through different sets of ATP assays using other substrates and reagents. ''[[Renilla]]''- and ''Gaussia''-based cell viability assays use the substrate coelenterazine.<ref>{{Cite journal |last=Tannous |first=Bakhos A. |date=April 2009 |title=Gaussia luciferase reporter assay for monitoring biological processes in culture and in vivo |url=http://dx.doi.org/10.1038/nprot.2009.28 |journal=Nature Protocols |volume=4 |issue=4 |pages=582–591 |doi=10.1038/nprot.2009.28 |pmid=19373229 |pmc=2692611 |issn=1754-2189}}</ref>

==See also==
*[[Animal coloration]]
*[[Biophoton]]
*''[[Life That Glows]]'', 2016 full-length documentary

==Notes==
{{notelist}}

==References==
{{reflist|26em}}

==Further reading==
* Schramm, Stefan; Weiß, Dieter (2024). "Bioluminescence – The Vibrant Glow of Nature and its Chemical Mechanisms". ''ChemBioChem''. '''25''' (9): e202400106. [https://chemistry-europe.onlinelibrary.wiley.com/doi/10.1002/cbic.202400106 doi:10.1002/cbic.202400106.]
* Victor Benno Meyer-Rochow (2009) Bioluminescence in Focus – a collection of illuminating essays Research Signpost: {{ISBN |978-81-308-0357-9}}
* [[Osamu Shimomura|Shimomura, Osamu]] (2006). ''Bioluminescence: Chemical Principles and Methods.'' Word Scientific Publishing. {{ISBN |981-256-801-8}}.
* Lee, John (2016). "Bioluminescence, the Nature of the Light." The University of Georgia Libraries. http://hdl.handle.net/10724/20031
* {{cite journal | last1=Wilson | first1=T. | last2=Hastings | first2=J.W. | year=1998 | title=Bioluminescence | journal=Annual Review of Cell and Developmental Biology | volume=14 | pages=197–230 | doi=10.1146/annurev.cellbio.14.1.197 | pmid=9891783 |ref=none }}
*Anctil, Michel (2018). ''Luminous Creatures: The History and Science of Light Production in Living Organisms''. McGill-Queen's University Press. {{ISBN |978-0-7735-5312-5}}

==External links==
{{Commons category |Bioluminescence}}
<!--please don't add Wikisource of 100yr old texts here, no use-->
{{Scholia |topic}}

<!--Please do not add your favourite forums, blogs, image galleries etc here as they will be removed-->
* [https://www.bbc.co.uk/news/av/world-us-canada-44076902/red-tide-electric-blue-waves-wash-california-shore BBC: Red tide: Electric blue waves wash California shore]
* [https://web.archive.org/web/20150214014827/http://www.mbari.org/staff/conn/botany/dinos/alimon/biolumin.htm MBARI: Gonyaulax Bioluminescence]
* [http://entomology.ifas.ufl.edu/creatures/misc/beetles/glow-worms.htm UF/IFAS: glow-worms]
* [http://www.ted.com/talks/edith_widder_glowing_life_in_an_underwater_world.html TED: Glowing life in an underwater world] (video)
* [http://ocean.si.edu/slideshow/bioluminescent-animals-photo-gallery Smithsonian Ocean Portal: Bioluminescent animals photo gallery]
* [https://web.archive.org/web/20130104165411/http://education.nationalgeographic.com/education/encyclopedia/bioluminescence/?ar_a=1 National Geographic: Bioluminescence]
* [http://www.annualreviews.org/eprint/pbWcm4DyqvQGJ7F2qGkc/full/10.1146/annurev-marine-120308-081028 Annual Review of Marine Science: Bioluminescence in the Sea]
* [https://www.canon.com.au/explore/bioluminescence-photography-tips Canon Australia – Tips on How to Photograph Bioluminescence]
* The [[New York City]] [[American Natural History Museum]]'s "Creatures of Light: Nature's Bioluminescence" 2022 Featured Exhibit (in concert with the [[Ottawa, Canada]]-based [[Canadian Museum of Nature]] and [[Chicago]]'s [[Field Museum of Natural History]]) webpage: [https://www.amnh.org/exhibitions/creatures-of-light]

{{aquatic ecosystem topics |state=expand}}
{{modelling ecosystems |expanded=none}}

{{Authority control}}

{{good article}}

[[Category:Bioluminescence| ]]
[[Category:Fisheries science]]
[[Category:Light sources]]
[[Category:Counter-illumination camouflage]]
[[Category:Mimicry]]
[[Category:Bioelectromagnetics]]