Bioluminescence

Bioluminescence is the production and emission of light by a living organism. It

is a form of chemiluminescence. Bioluminescence occurs widely in marine
vertebrates and invertebrates, as well as in some fungi, microorganisms
including some bioluminescent bacteria and terrestrial invertebrates such as
fireflies. In some animals, the light is bacteriogenic, produced by symbiotic
organisms such as Vibrio bacteria; in others, it is autogenic, produced by the
animals themselves.

In a general sense, the principal chemical reaction in bioluminescence involves

some light-emitting molecule and an enzyme, generally called the luciferin and
the luciferase, respectively. Because these are generic names, the luciferins and
Flying and glowing firefly, Photinus
luciferases are often distinguished by including the species or group, i.e. Firefly
pyralis
luciferin. In all characterized cases, the enzyme catalyzes the oxidation of the
luciferin.

In some species, the luciferase requires other 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 eleven
different animal (phyla), though in some of these, the animals obtain it through their diet.
Conversely, luciferases vary widely between different species, and consequently
bioluminescence has arisen over forty times in evolutionary history.

Both Aristotle and Pliny the Elder mentioned that damp wood sometimes gives off a glow
and many centuries later Robert Boyle showed that oxygen was involved in the process,
both in wood and in glow-worms. It was not until the late nineteenth century that
bioluminescence was properly investigated. The phenomenon is widely distributed among Female Glowworm,
animal groups, especially in marine environments where dinoflagellates cause Lampyris noctiluca
phosphorescence in the surface layers of water. On land it occurs in fungi, bacteria and
some groups of invertebrates, including insects.

The uses of bioluminescence by animals include counter-illumination

camouflage, mimicry of other animals, for example to lure prey, and signalling
to other individuals of the same species, such as to attract mates. In the
laboratory, luciferase-based systems are used in genetic engineering and for
biomedical research. Other researchers are investigating the possibility of using
bioluminescent systems for street and decorative lighting, and a bioluminescent Male and female of the species
plant has been created.[1] Lampyris noctiluca mating. The
female of this species has no wings,
she's a larviforme, unlike the male.

Contents
History
Evolution
 Chemical mechanism
Distribution
Uses in nature
Counterillumination camouflage
Attraction
Defense
Warning
Communication
Mimicry
Illumination
Biotechnology
Biology and medicine
Light production
See also
Notes
References
Further reading
External links

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.[2] This experimental form of illumination avoided the necessity of using candles which risked sparking
explosions of firedamp.[3] Another safe source of illumination in mines was bottles containing fireflies.[4] 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) mention light from damp wood. He also records that Robert Boyle experimented on these light sources, and
showed that both they and the glow-worm 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",[5] and in 1854 Johann Florian Heller (1813–1871)
identified strands (hyphae) of fungi as the source of light in dead wood.[6]

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".[7]

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

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] and noted 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]

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.[11]

Among the anecdotes of navigation by bioluminescence, the Apollo 13 astronaut

Jim Lovell recounted how as a navy pilot he had found his way back to his
aircraft carrier USS Shangri-La when his navigation systems failed. Turning off Osamu Shimomura isolated the
photoprotein aequorin and its
his cabin lights, he saw the glowing wake of the ship, and was able to fly to it
cofactor coelenterazine from the
and land safely.[12] crystal jelly Aequorea victoria in
1961.[10]
The French pharmacologist Raphaël Dubois carried out work on
bioluminescence in the late nineteenth century. He studied click beetles
(Pyrophorus) and the marine bivalve mollusc Pholas 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 named
luciferin, by an enzyme.[15] He sent Harvey 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 is 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 thirty 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 sea firefly Vargula hilgendorfii, but it was another ten years
before he discovered the chemical's structure and was able to publish his 1957 paper Crystalline Cypridina Luciferin.[16] More
recently, Martin Chalfie, Osamu Shimomura 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.[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]

In 2016, deep sea bioluminescent corals were captured for the first time in color HD Video.[21]

Evolution
E. N. Harvey (1932) was among the first to propose how bioluminescence could have evolved.[22] 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 the ones put forth by Seliger (1993) and Rees et al.
(1998).[23][24]
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 applied forces that favored the development of increased eye sensitivity and enhanced visual
signals.[25] 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.[23]

Rees et al. (1998) uses 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 (ROS) (e.g.
H2O2 and 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.[24]

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.[26]
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 27 occasions, 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.[27][28]

All bioluminescent organism have one thing in common. A "luciferin" and oxygen is always catalyzed by luciferase to light.[29]
McElroy and Seliger proposed 1962 that the bioluminescent reaction evolved to detoxify oxygen. So bioluminescence evolved
parallel with photosynthetic organism.[30] Today bioluminescence has other purposes.

Chemical mechanism
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.[31] 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, though many examples have a
concurrent release of carbon dioxide. For example, the firefly luciferin/luciferase
reaction requires magnesium and ATP and produces carbon dioxide (CO2),
adenosine monophosphate (AMP) and pyrophosphate (PP) as waste products.
Other cofactors may be required for the reaction, such as calcium (Ca2+) for the
Protein structure of the luciferase of
photoprotein aequorin, or magnesium (Mg2+) ions and ATP for the firefly the firefly Photinus pyralis. The
luciferase.[32] Generically, this reaction could be described as: enzyme is a much larger molecule
than luciferin.

Instead of a luciferase, the jellyfish Aequorea victoria makes use of another type of protein called a photoprotein, in this case
specifically aequorin.[33] When calcium ions are added, the 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 oxidised (oxyluciferin) form, allowing it
to recombine with aequorin, in readiness for a subsequent flash. Photoproteins are thus enzymes, but with unusual reaction
kinetics.[34] 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 resonant energy transfer.[35]
Overall, bioluminescence has arisen over forty times in evolutionary history.[31]
In evolution, luciferins tend to vary little: one in particular, coelenterazine, is the
light emitting pigment for nine phyla (groups of very different organisms),
including polycystine radiolaria, Cercozoa (Phaeodaria), protozoa, comb jellies,
cnidaria including jellyfish and corals, crustaceans, molluscs, arrow worms and
vertebrates (ray-finned fish). Not all these organisms synthesize coelenterazine:
some of them obtain it through their diet.[31] Conversely, luciferase enzymes
vary widely and tend to be different in each species.[31] Coelenterazine is a luciferin found in
many different marine phyla from

Distribution comb jellies to vertebrates. Like all

luciferins, it is oxidised to produce
light.

Bioluminescence occurs widely among animals,

Huge numbers of bioluminescent dinoflagellates creating
phosphorescence in breaking waves
light, and the genus Tomopteris emits yellow light.[31][37]
especially in the open sea, including fish, jellyfish,
comb jellies, crustaceans, and cephalopod molluscs; in
some fungi and bacteria; and in various terrestrial
invertebrates including insects. About 76% of the main
taxa of deep-sea animals produce light.[36] Most marine
light-emission is in the blue and green light spectrum.
However, some loose-jawed fish emit red and infrared

The most frequently encountered bioluminescent organisms may be the dinoflagellates present in the surface layers of the sea,
which are responsible for the sparkling phosphorescence sometimes seen at night in disturbed water. At least eighteen genera
exhibit luminosity.[31] A different effect is the thousands of square miles of the ocean which shine with the light produced by
bioluminescent bacteria, known as mareel or the milky seas effect.[38]

Non-marine bioluminescence is less widely distributed, the two best-known cases being in fireflies and glow worms. Other
invertebrates including insect larvae, annelids and arachnids possess bioluminescent abilities. Some forms of bioluminescence are
brighter (or exist only) at night, following a circadian rhythm.

Uses in nature
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 analyse the chemical mechanisms or to prove what function the light serves.[31] In
some cases the function is unknown, as with species in three families of earthworm (Oligochaeta), such as Diplocardia longa
where the coelomic fluid produces light when the animal moves.[39] The following functions are reasonably well established in
the named organisms.

Counterillumination camouflage
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.[40] In these animals,
photoreceptors control the illumination to match the brightness of the background.[40] 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.[41]

Attraction
A fungus gnat from New Zealand, Arachnocampa luminosa, lives in the
predator-free environment of caves and its larvae emit bluish-green light.[42]
They dangle silken threads that glow and attract flying insects, and wind in their
Principle of counterillumination
fishing-lines when prey becomes entangled.[43] The bioluminescence of the
camouflage in firefly squid,
larvae of another fungus gnat from North America which lives on streambanks
Watasenia scintillans. When seen
and under overhangs has a similar function. Orfelia fultoni builds sticky little from below by a predator, the
webs and emits light of a deep blue colour. It has an inbuilt biological clock and, bioluminescence helps to match the
even when kept in total darkness, turns its light on and off in a circadian squid's brightness and colour to the
rhythm.[44] sea surface above.

Fireflies use light to attract 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.[39][45] Click beetles 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.[39] 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.[39]

In the marine environment, use of luminescence for mate attraction is chiefly known among ostracods, small shrimplike
crustaceans, especially in the family Cyprididae. Pheromones may be used for long-distance communication, with
bioluminescence used at close range to enable mates to "home in".[31] A polychaete worm, the Bermuda fireworm creates a brief
display, a few nights after the full moon, when the female lights up to attract males.[46]

Defense
Many cephalopods, including at least 70 genera of squid, are bioluminescent.[31] Some squid and small crustaceans use
bioluminescent chemical mixtures or bacterial slurries in the same way as many squid use ink. A cloud of luminescent material is
expelled, distracting or repelling a potential predator, while the animal escapes to safety.[31] The deep sea squid Octopoteuthis
deletron may autotomise portions of its arms which are luminous and continue to twitch and flash, thus distracting a predator
while the animal flees.[31]

Dinoflagellates may use bioluminescence for defence against predators. They shine when they detect a predator, possibly making
the predator itself more vulnerable by attracting the attention of predators from higher trophic levels.[31] 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]

The 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
military as a source of light during clandestine operations.[16]
The larvae of railroad worms (Phrixothrix) have paired photic organs on each body segment, able to glow with green light; these
are thought to have a defensive purpose.[47] They also have organs on the head which produce red light; they are the only
terrestrial organisms to emit light of this colour.[48]

Warning
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 millipedes glow for the same purpose.[49] Some marine
organisms are believed to emit light for a similar reason. These include scale worms, jellyfish and brittle stars but further research
is needed to fully establish the function of the luminescence. Such a mechanism would be of particular advantage to soft-bodied
cnidarians if they were able to deter predation in this way.[31] The limpet Latia neritoides is the only known freshwater gastropod
that emits light. It produces greenish luminescent mucus which may have an anti-predator function.[50] 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.[51]

Communication
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.[31]

Pyrosomes are colonial 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
Pyrosoma, a colonial tunicate; each
from other nearby colonies.[52] Communication by light emission between the
individual zooid in the colony flashes
zooids enables coordination of colony effort, for example in swimming where
a blue-green light.
each zooid provides part of the propulsive force.[53]

Some bioluminous bacteria infect nematodes that parasitize Lepidoptera larvae.

When these caterpillars die, their luminosity may attract predators to the dead insect thus assisting in the dispersal of both bacteria
and nematodes.[39] 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, cap and gills.
This may attract night-flying insects and aid in spore dispersal, but other functions may also be involved.[39]

Quantula striata is the only known bioluminescent terrestrial mollusc. 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.[54]

Mimicry
Bioluminescence is used by a variety of animals to mimic other species. Many species of deep sea fish such as the anglerfish and
dragonfish make use of aggressive mimicry to attract prey. They have an appendage on their heads called an 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.[31][55]

The cookiecutter shark uses bioluminescence to camouflage its underside by counterillumination, 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.[56][57]
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 lucibufagins, which Photuris cannot synthesize.[58]

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.[59] However, doubt has been cast on
this assertion, and there is no conclusive evidence that the cockroaches are
bioluminescent.[60][61]
A deep sea anglerfish, Bufoceratias
wedli, showing the esca (lure)
Illumination
While most marine bioluminescence is green to blue, some deep
sea barbeled dragonfishes in the genera Aristostomias,
Pachystomias and Malacosteus emit a red glow. This adaptation
allows the fish to see red-pigmented prey, which are normally
invisible in the deep ocean environment where red light has been
filtered out by the water column.[62]

The black dragonfish (also called the northern stoplight loosejaw)

Malacosteus niger is believed to be one of the only fish to produce
a red glow. Its eyes, however, are insensitive to this wavelength; it
Flashing of photophores of black dragonfish,
has an additional retinal pigment which fluoresces blue-green when Malacosteus niger, showing red fluorescence
illuminated. This alerts the fish to the presence of its prey. The
additional pigment is thought to be assimilated from chlorophyll
derivatives found in the copepods which form part of its diet.[63]

Biotechnology

Biology and medicine

Bioluminescent organisms are a target for many areas of research. Luciferase systems are widely used in genetic engineering as
reporter genes, each producing a different colour by fluorescence,[64][65] and for biomedical research using bioluminescence
imaging.[66][67][68] For example, the firefly luciferase gene was used as early as 1986 for research using transgenic tobacco
plants.[69] Vibrio bacteria symbiose with marine invertebrates such as the Hawaiian bobtail squid (Euprymna scolopes), are key
experimental models for bioluminescence.[70][71] Bioluminescent activated destruction is an experimental cancer treatment.[72]
See also optogenetics which involves the use of light to control cells in living tissue, typically neurons, that have been genetically
modified to express light-sensitive ion channels, and also see biophoton, a photon of non-thermal origin in the visible and
ultraviolet spectrum emitted from a biological system.

Light production
The structures of photophores, the light producing organs in bioluminescent organisms, are being investigated by industrial
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][73][74] The gene that makes the tails of 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.[75] University of Wisconsin–Madison is researching the use
of genetically engineered bioluminescent E. coli bacteria, for use as bioluminescent bacteria in a light bulb.[76] In 2011, Philips
launched a microbial system for ambience lighting in the home.[77][78] An 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; this enzyme "helps to strengthen and sustain light
output".[79] In 2016, Glowee, a French company started selling bioluminescent lights, targeting shop fronts and municipal street
signs as their main markets.[80] France has a law that forbids retailers and offices from illuminating their windows between 1 and
7 in the morning in order to minimise energy consumption and pollution.[81][82] Glowee hoped their product would get around
this ban. They used bacteria called Aliivibrio fischeri which glow in the dark, but the maximum lifetime of their product was three
days.[80]

See also
Animal coloration

Notes
a. 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]

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Further reading
Victor Benno Meyer-Rochow (2009) Bioluminescence in Focus – a collection of illuminating essays Research
Signpost: ISBN 978-81-308-0357-9
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
Wilson, T.; Hastings, J.W. (1998). "Bioluminescence". Annual Review of Cell and Developmental Biology. 14:
197–230. doi:10.1146/annurev.cellbio.14.1.197 (https://doi.org/10.1146%2Fannurev.cellbio.14.1.197).
PMID 9891783 (https://www.ncbi.nlm.nih.gov/pubmed/9891783).
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
BBC: Red tide: Electric blue waves wash California shore (https://www.bbc.co.uk/news/av/world-us-canada-4407
6902/red-tide-electric-blue-waves-wash-california-shore)
MBARI: Gonyaulax Bioluminescence (https://web.archive.org/web/20150214014827/http://www.mbari.org/staff/co
nn/botany/dinos/alimon/biolumin.htm)
UF/IFAS: glow-worms (http://entomology.ifas.ufl.edu/creatures/misc/beetles/glow-worms.htm)
TED: Glowing life in an underwater world (http://www.ted.com/talks/edith_widder_glowing_life_in_an_underwater
_world.html) (video)
Smithsonian Ocean Portal: Bioluminescent animals photo gallery (http://ocean.si.edu/slideshow/bioluminescent-a
nimals-photo-gallery)
National Geographic: Bioluminescence (http://education.nationalgeographic.com/education/encyclopedia/biolumi
nescence/?ar_a=1)
 Annual Review of Marine Science: Bioluminescence in the Sea (http://www.annualreviews.org/eprint/pbWcm4Dy
qvQGJ7F2qGkc/full/10.1146/annurev-marine-120308-081028)

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