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Bioluminescence

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Not to be confused withIridescence,Structural coloration,orPhosphorescence.

Flying and glowingfirefly,Photinus pyralis
Femaleglowworm,Lampyris noctiluca

Bioluminescenceis the production and emission oflightby livingorganisms.It is a form ofchemiluminescence.Bioluminescence occurs widely in marinevertebratesandinvertebrates,as well as in somefungi,microorganisms including somebioluminescent bacteria,and terrestrial arthropods such asfireflies.In some animals, the light is bacteriogenic, produced bysymbioticbacteria such as those from the genusVibrio;[1]in others, it is autogenic, produced by the animals themselves.

In a general sense, the principal chemical reaction in bioluminescence involves a light-emitting molecule and anenzyme,generally calledluciferinandluciferase,respectively. Because these are generic names, luciferins and luciferases are often distinguished by the species or group, e.g.firefly luciferin.In all characterized cases, the enzymecatalyzestheoxidationof the luciferin.

In some species, the luciferase requires othercofactors,such ascalciumormagnesiumions, and sometimes also the energy-carrying moleculeadenosine triphosphate(ATP). Inevolution,luciferins vary little: one in particular,coelenterazine,is found in 11 different animalphyla,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 inevolutionary history.

BothAristotleandPliny the Eldermentioned that damp wood sometimes gives off a glow. Many centuries laterRobert Boyleshowed 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 ofinvertebrates,includinginsects.

The uses of bioluminescence by animals includecounterilluminationcamouflage,mimicryof other animals, for example to lure prey, andsignalingto 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.[2]

History

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Before the development of thesafety lampfor use in coal mines, dried fish skins were used in Britain and Europe as a weak source of light.[3]This experimental form of illumination avoided the necessity of using candles which risked sparking explosions offiredamp.[4]In 1920, the American zoologistE. Newton Harveypublished a monograph,The Nature of Animal Light,summarizing early work on bioluminescence. Harvey notes thatAristotlementions light produced by dead fish and flesh, and that both Aristotle andPliny the Elder(in hisNatural History) mention light from damp wood. He records thatRobert Boyleexperimented 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 flagellateNoctiluca"as a luminous animal" "just visible to the naked eye",[5]and in 1854Johann Florian Heller(1813–1871) identified strands (hyphae) of fungi as the source of light in dead wood.[6]

Tuckey,in his posthumous 1818Narrative 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".[7]

Charles Darwinnoticed bioluminescence in the sea, describing it in hisJournal:

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.[8]

Darwin also observed a luminous "jelly-fish of the genus Dianaea",[8]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."[8]He guessed that "a disturbed electrical condition of the atmosphere"[8]was probably responsible. Daniel Pauly comments that Darwin "was lucky with most of his guesses, but not here",[9]noting that biochemistry was too little known, and that the complex evolution of the marine animals involved "would have been too much for comfort".[9]

Osamu Shimomuraisolated thephotoproteinaequorinand its cofactorcoelenterazinefrom the crystal jellyAequorea victoriain 1961.[10]

Bioluminescence attracted the attention of theUnited States Navyin theCold War,since submarines in some waters can create a bright enough wake to be detected; a German submarine was sunk in theFirst 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.[11]

Among the anecdotes of navigation by bioluminescence is one recounted by theApollo 13astronautJim Lovell,who as a navy pilot had found his way back to his aircraft carrierUSSShangri-Lawhen 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.[12]

TheFrenchpharmacologistRaphaël Duboiscarried out work on bioluminescence in the late nineteenth century. He studiedclick beetles(Pyrophorus) and the marinebivalvemolluscPholas dactylus.He refuted the old idea that bioluminescence came from phosphorus,[13][a]and demonstrated that the process was related to the oxidation of a specific compound, which he namedluciferin,by anenzyme.[15]He sent Harveysiphonsfrom 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 chemistOsamu Shimomurato be the first to obtain crystalline luciferin. He used thesea fireflyVargula hilgendorfii,but it was another ten years before he discovered the chemical's structure and published his 1957 paperCrystalline Cypridina Luciferin.[16]Shimomura,Martin Chalfie,andRoger Y. Tsienwon the 2008Nobel Prize in Chemistryfor their 1961 discovery and development ofgreen fluorescent proteinas a tool for biological research.[17]

Harvey wrote a detailed historical account on all forms of luminescence in 1957.[18]An updated book on bioluminescence covering also the twentieth and early twenty-first century was published recently.[19][20]

Evolution

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In 1932 E. N. Harvey was among the first to propose how bioluminescence could have evolved.[21]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 byHoward Seligerin 1993 and Rees et al. in 1998.[22][23]

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.[24]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.[22]

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. H2O2and O2−). 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.[23]

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.[25]Bioluminescence in fish began at least by theCretaceousperiod. 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.[26][27]

All bioluminescent organisms have in common that the reaction of a "luciferin" and oxygen is catalyzed by a luciferase to produce light.[28]McElroy and Seliger proposed in 1962 that the bioluminescent reaction evolved to detoxify oxygen, in parallel with photosynthesis.[29]

Thuesen, Davis et al. showed in 2016 that bioluminescence has evolved independently 27 times within 14 fish clades across ray-finned fishes.[26]The oldest of these appears to be Stomiiformes and Myctophidae.[30]In sharks, bioluminescence has evolved only once.[31] Genomic analysis ofoctocoralsindicates that their ancestor was bioluminescent as long as 540 million years ago.[32]

Chemical mechanism

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Main article:Luciferase
Protein structureof theluciferaseof thefireflyPhotinus pyralis.The enzyme is a much larger molecule than luciferin.

Bioluminescence is a form ofchemiluminescencewhere light energy is released by a chemical reaction. This reaction involves a light-emitting pigment, theluciferin,and aluciferase,the enzyme component.[33]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 molecularoxygen;often there is a concurrent release ofcarbon dioxide(CO2). For example, the firefly luciferin/luciferase reaction requiresmagnesiumandATPand produces CO2,adenosine monophosphate(AMP) andpyrophosphate(PP) as waste products. Other cofactors may be required, such as calcium (Ca2+) for thephotoproteinaequorin,ormagnesium(Mg2+) ions and ATP for thefirefly luciferase.[34]Generically, this reaction can be described as:

Luciferin + O2Oxyluciferin + light energy
Coelenterazineis aluciferinfound in many different marine phyla fromcomb jelliestovertebrates.Like all luciferins, it is oxidised to produce light.

Instead of a luciferase, the jellyfishAequorea victoriamakes use of another type of protein called aphotoprotein,in this case specificallyaequorin.[35]When calcium ions are added, rapidcatalysiscreates 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 thusenzymes,but with unusual reaction kinetics.[36]Furthermore, some of the blue light released by aequorin in contact with calcium ions is absorbed by agreen fluorescent protein,which in turn releases green light in a process calledresonant energy transfer.[37]

Overall, bioluminescence has arisen over 40 times in evolutionary history.[33]Inevolution,luciferins tend to vary little: one in particular,coelenterazine,is the light emitting pigment for ninephyla(groups of very different organisms), including polycystineradiolaria,Cercozoa(Phaeodaria),protozoa,comb jellies,cnidariaincludingjellyfishandcorals,crustaceans,molluscs,arrow wormsandvertebrates(ray-finned fish). Not all these organisms synthesise coelenterazine: some of them obtain it through their diet.[33]Conversely, luciferase enzymes vary widely and tend to be different in each species.[33]

Distribution

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Further information:List of bioluminescent organisms
Huge numbers ofdinoflagellatescreating bioluminescence in breaking waves

Bioluminescence occurs widely among animals, especially in the open sea, includingfish,jellyfish,comb jellies,crustaceans,andcephalopodmolluscs; in somefungiandbacteria;and in various terrestrial invertebrates, nearly all of which arebeetles.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 ofdeep-sea animalshave been found to be capable of producing light.[38]More than 700 animal genera have been recorded with light-producing species.[39]Most marine light-emission is in the blue and greenlight spectrum.However, someloose-jawed fishemit red andinfraredlight, and the genusTomopterisemits yellow light.[33][40]

The most frequently encountered bioluminescent organisms may be thedinoflagellatesin 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 thesephytoplanktonexhibit luminosity.[33]Luminescent dinoflagellate ecosystems are present in warm water lagoons and bays with narrow openings to the ocean.[41]A different effect is the thousands of square miles of the ocean which shine with the light produced by bioluminescent bacteria, known asmareel or the milky seas effect.[42]

Pelagic zone

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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.[43]

Bacterial symbioses

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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 byPhotobacteriumandVibrio.[44]

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,spawning,or the luminous bacterium is evolving with their host.[45]Coevolutionary interactions are suggested as host organisms' anatomical adaptations have become specific to only certain luminous bacteria, to suffice ecological dependence of bioluminescence.[46]

Benthic zone

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Bioluminescence is widely studied amongst species located in the mesopelagic zone, but thebenthic zoneat 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.[47]

Uses in nature

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Mycena chlorophos,abioluminescent mushroom

Bioluminescence has several functions in different taxa.Steven Haddocket 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.[33]In some cases the function is unknown, as with species in three families of earthworm (Oligochaeta), such asDiplocardia longa,where thecoelomic fluidproduces light when the animal moves.[48]The following functions are reasonably well established in the named organisms.

Counterillumination camouflage

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Principle ofcounterilluminationcamouflagein 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 information:Counterillumination

In many animals of the deep sea, including severalsquidspecies, bacterial bioluminescence is used forcamouflagebycounterillumination,in which the animal matches the overhead environmental light as seen from below.[49]In these animals, photoreceptors control the illumination to match the brightness of the background.[49]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.[50]

Attraction

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Stauroteuthis syrtensisbioluminescent photophores

Bioluminescence is used in a variety of ways and for different purposes. The cirrate octopodStauroteuthis syrtensisuses emits bioluminescence from its sucker like structures.[51]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 ofphotophores.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.[52]

Firefliesuse light to attractmates.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.[48][53]Click beetlesemit 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.[48]Larvae of the click beetlePyrophorus nyctophanuslive 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.[48]

In the marine environment, use of luminescence for mate attraction is chiefly known amongostracods,small shrimp-likecrustaceans,especially in the familyCyprididae.Pheromonesmay be used for long-distance communication, with bioluminescence used at close range to enable mates to "home in".[33]Apolychaeteworm, theBermuda firewormcreates a brief display, a few nights after the full moon, when the female lights up to attract males.[54]

Defense

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Acanthephyra purpureahas 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.[55]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.[55]

Manycephalopods,including at least 70generaofsquid,are bioluminescent.[33]Some squid and smallcrustaceansuse bioluminescent chemical mixtures or bacterial slurries in the same way as many squid useink.A cloud of luminescent material is expelled, distracting or repelling a potential predator, while the animal escapes to safety.[33]The deep sea squidOctopoteuthis deletronmayautotomizeportions of its arms which are luminous and continue to twitch and flash, thus distracting a predator while the animal flees.[33]

Dinoflagellatesmay use bioluminescence for defense againstpredators.They shine when they detect a predator, possibly making the predator itself more vulnerable by attracting the attention of predators from higher trophic levels.[33]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.[9]

Thesea-fireflyis 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.[16]

The larvae ofrailroad worms(Phrixothrix) have paired photic organs on each body segment, able to glow with green light; these are thought to have a defensive purpose.[56]They also have organs on the head which produce red light; they are the only terrestrial organisms to emit light of this color.[57]

Warning

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Aposematismis 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; somemillipedesglow for the same purpose.[58]Some marine organisms are believed to emit light for a similar reason. These includescale worms,jellyfishandbrittle starsbut further research is needed to fully establish the function of the luminescence. Such a mechanism would be of particular advantage to soft-bodiedcnidariansif they were able to deter predation in this way.[33]ThelimpetLatia neritoidesis the only known freshwatergastropodthat emits light. It produces greenish luminescentmucuswhich may have an anti-predator function.[59]The marine snailHinea brasilianauses 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.[60]

Communication

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Pyrosoma,a colonialtunicate;each individual zooid in the colony flashes a blue-green light.

Communication in the form ofquorum sensingplays 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.[33]

Pyrosomesare colonialtunicatesand eachzooidhas 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.[61]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.[62]

Some bioluminous bacteria infectnematodesthat parasitizeLepidopteralarvae. When thesecaterpillarsdie, their luminosity may attract predators to the dead insect thus assisting in the dispersal of both bacteria and nematodes.[48]A similar reason may account for the many species of fungi that emit light. Species in the generaArmillaria,Mycena,Omphalotus,Panellus,Pleurotusand others do this, emitting usually greenish light from themycelium,capandgills.This may attract night-flying insects and aid in spore dispersal, but other functions may also be involved.[48]

Quantula striatais 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.[63]

Mimicry

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Adeep seaanglerfish,Bufoceratias wedli,showing theesca(lure)

Bioluminescence is used by a variety of animals tomimicother species. Many species ofdeep sea fishsuch as theanglerfishanddragonfishmake use ofaggressive mimicryto attractprey.They have anappendageon their heads called anescathat 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.[33][64]

Thecookiecutter sharkuses 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 liketunaandmackerelswimming beneath it. When such fish approach the lure, they are bitten by the shark.[65][66]

FemalePhoturisfireflies 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 namedlucibufagins,whichPhoturiscannot synthesize.[67]

South American giant cockroaches of the genusLucihormeticawere believed to be the first known example of defensive mimicry, emitting light in imitation of bioluminescent, poisonous click beetles.[68]However, doubt has been cast on this assertion, and there is no conclusive evidence that the cockroaches are bioluminescent.[69][70]

Flashing of photophores of black dragonfish,Malacosteus niger,showing red fluorescence

Illumination

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While most marine bioluminescence is green to blue, some deep seabarbeled dragonfishesin the generaAristostomias,PachystomiasandMalacosteusemit 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.[71]These fish are able to utilize the longer wavelength to act as a spotlight for its prey that only they can see.[71]The fish may also use this light to communicate with each other to find potential mates.[72]The ability of the fish to see this light is explained by the presence of specialized rhodopsin pigment.[71]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.[73]The dragonfish species which produce the red light also produce blue light in photophore on the dorsal area.[73]The main function of this is to alert the fish to the presence of its prey.[74]The additional pigment is thought to be assimilated fromchlorophyllderivatives found in thecopepodswhich form part of its diet.[74]

The angler siphonophore (Erenna) utilizes red bioluminescence in appendages to lure fish.[73]

Biotechnology

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Biology and medicine

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Further information:optogenetics

Bioluminescent organisms are a target for many areas of research. Luciferase systems are widely used ingenetic engineeringasreporter genes,each producing a different color by fluorescence,[75][76]and for biomedical research usingbioluminescence imaging.[77][78][79]For example, the firefly luciferase gene was used as early as 1986 for research using transgenic tobacco plants.[80]Vibriobacteria symbiose withmarine invertebratessuch as theHawaiian bobtail squid(Euprymna scolopes), are keyexperimental modelsfor bioluminescence.[81][82]Bioluminescent activated destructionis an experimental cancer treatment.[83]

In Vivoluminescence cell and animal imaging uses dyes and fluorescent proteins aschromophores.The characteristics of each chromophore determine which cell area(s) will be targeted and illuminated.[84]

Light production

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A "Firefly"petunia,genetically engineered to produce luciferase.

The structures ofphotophores,the light producing organs in bioluminescent organisms, are being investigated byindustrial designers.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.[11][85][86]The gene that makes the tails offirefliesglow 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.[87]University of Wisconsin–Madisonis researching the use of genetically engineered bioluminescentE. colibacteria, for use asbioluminescent bacteriain alight bulb.[88]In 2011,Philipslaunched a microbial system for ambience lighting in the home.[89][90]AniGEMteam 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.[91]In 2016, Glowee, a French company started selling bioluminescent lights for shop fronts and street signs,[92]for use between 1 and 7 in the morning when the law forbids use of electricity for this purpose.[93][94]They used the bioluminescent bacteriumAliivibrio fischeri,but the maximum lifetime of their product was three days.[92]In April 2020, plants were genetically engineered to glow more brightly using genes from the bioluminescent mushroomNeonothopanus nambito convertcaffeic acidinto luciferin.[94][95]

ATP bioluminescence

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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 goodbiosensorto test for the presence of livingmicrobesin water.[96][97]Different types of microbial populations are determined through different sets of ATP assays using other substrates and reagents.Renilla- andGaussia-based cell viability assays use the substrate coelenterazine.[98]

See also

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Notes

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  1. ^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".[14]

References

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  1. ^Ples, Marek (11 November 2021)."Lab Snapshots by Marek Ples; Microbiology - The biology on a different level".weirdscience.eu.Retrieved2 July2023.
  2. ^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
  3. ^Smiles, Samuel(1862).Lives of the Engineers.Vol. III (George and Robert Stephenson). London: John Murray. p. 107.ISBN978-0-7153-4281-7.(ISBN refers to the David & Charles reprint of 1968 with an introduction byL. T. C. Rolt)
  4. ^Freese, Barbara (2006).Coal: A Human History.Arrow. p. 51.ISBN978-0-09-947884-3.
  5. ^Harvey cites this as Baker, J.: 1743–1753,The Microscope Made Easy and Employment for the Microscope.
  6. ^Harvey, E. Newton (1920).The Nature of Animal Light.Philadelphia & London: J. B. Lippencott. p. 1.
  7. ^Tuckey, James Hingston(May 1818).Thomson, Thomas(ed.)."Narrative of the Expedition to the Zaire".Annals of Philosophy.11(65): 392.
  8. ^abcdDarwin, Charles(1839).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.Henry Colburn. pp. 190–192.
  9. ^abcPauly, Daniel (13 May 2004).Darwin's Fishes: An Encyclopedia of Ichthyology, Ecology, and Evolution.Cambridge University Press. pp. 15–16.ISBN978-1-139-45181-9.
  10. ^Shimomura, O. (August 1995)."A short story of aequorin".The Biological Bulletin.189(1): 1–5.doi:10.2307/1542194.JSTOR1542194.PMID7654844.
  11. ^ab"How illuminating".The Economist.10 March 2011.Retrieved6 December2014.
  12. ^Huth, John Edward (15 May 2013).The Lost Art of Finding Our Way.Harvard University Press. p. 423.ISBN978-0-674-07282-4.
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