20th May, 2016

Sara Cauty

In what way is bioluminescence important?

The world is full of wonders. Even when there is no light, nature will light
up, filling the space with awe-inspiring beauty. These spectacular shows of
light are part of a phenomenon known as bioluminescence. This process has
brought the main inquiry question of this paper; in what way is
bioluminescence important? This questions are explained by the three inquiry
sub-questions; how is bioluminescence a useful trait, which chemical
processes cause bioluminescence and which adaptations caused
bioluminescence to evolve? It is a phenomenon that has much to be discovered
and understood, and yet presents so many opportunities to the world. It adds
beauty and light to nature, even where the light cannot touch, deep
underground, or in the depths of the ocean.

Biology: How is bioluminescence a useful trait?

What is bioluminescence? Bioluminescence is, the emission of visible

light by biological systems, which arises from enzyme-catalyzed chemical
reactions. (Meighen, 2016) There are many terms that are associated with
bioluminescence. Words such as luminescence and chemiluminescence. They
are not as complicated as they might seem, as they all share common root
words that imply their meanings. Luminescence is light that is produced by a
reaction that does not create heat. Chemiluminescence is similar to
luminescence, in which it is a light that is created by a chemical reaction.
It is an awe-striking natural occurrence that has kept scientists curious
for centuries. Bioluminescence is most commonly found in marine animals,
however, it is not selective to organisms that live in the ocean. Luminescent
species are found among marine and terrestrial bacteria, annelids or
segmented worms, beetles, algae, mollusk, coelenterates, bony fish, and
cartilaginous fish. (Meighen, 2016) Bioluminescence is not only found in
organisms that contain luminescent organs, but some animals collect

luminescent bacteria in a sac or light organ to produce bioluminescence for

them.

Bioluminescence occurs differently in each organism, there are three

main bioluminescent systems that have been studied and identified. These
systems are fireflies/beetles, dinoflagellates, and bacteria. Each system has
their own defining characteristics, and tend to glow in different ways that help
to identify them. Fireflies, for instance emit [colour] in the yellow to green
range, as part of a courtship process; click beetles emit green to orange light;
whereas railway worms emit red light, with green light being emitted on
movement. (Meighen, 2016) Dinoflagellates, or unicellular algae accounts
for many of the recorded observations that have described the apparent
"phosphorescence" of the sea. Dinoflagellates are very sensitive to motion
induced by ships or fish, and respond with rapid and brilliant flashes.
(Meighen, 2016) While Dinoflagellates are known to cause red tides when they
bloom, they do not glow red. They glow a more green-blue hue that is bright
and easy to see.
Bacteria provide a wide variety of bioluminescent organisms. Not only are
they able to produce their own light, but other animals such as the Angler fish
can become bioluminescent by containing the bacteria; much like trapping
fireflies in a jar to make a lantern. Requiring salt for growth and
luminescenceThe most common [bioluminescent bacteria] are Vibrio and
Photo bacterium species. While luminous bacteria come in various shapes,
they do not form clusters or chains, as do many other bacteria. The light of an
individual bacterium, of course, cannot be seen with the naked eye, but the
light from a liquid or agar culture containing billions of bacteria is readily
visible. The light is bluish and continuous. Many luminous bacteria live in the
light organs of fish and squids, without adversely affecting their hosts.
("bioluminescence | chemical reaction", 2016) Bioluminescent bacteria can
exist in the ocean as well as on land. These bacteria as well as bioluminescent
fungus are known as foxfire, a beautiful but harmless display of light during
the night. Foxfire is usually found on fungus that feed on decaying wood and
plants, and will emit a continuous glow rather than flashes. ("bioluminescence
| chemical reaction", 2016) Most bioluminescence is produced by their
light organs [that] flash for periods of less than a second to about 10 seconds.

These flashes can occur in specific spots, such as the dots on a squid. Other
flashes can illuminate the organism's entire body. (Evers, 2013)
There are many different types of bioluminescence, and many colours
alongside it. There are meanings for the colours, dependant on their
environment, species and purpose for the luminescence. In the ocean, the light
will only appear blue or green, as the water interferes with any other
wavelength of bioluminescence, and most marine organisms do not possess the
sight organs to see red, yellow or violet. Terrestrial organisms such as some
beetles, bacteria and fungus, will also exhibit the blue-green spectrum of light.
However, there are exceptions such as fireflies and some beetle larvae, who will
glow yellow.
Besides the environment the organisms live in, the cause for
bioluminescence is decided by five main functions: camouflage, repulsion,
illumination, attraction and communication. For camouflage, the
bioluminescence used is known as counterillumination. Many predators, such
as sharks, hunt from below. They look above, where sunlight creates shadows
beneath prey. Counterillumination is a type of camouflage against this
predatory behavior. (Evers, 2013)
Hatchet fish use counterillumination. Hatchet fish have light-producing
organs that point downward. They adjust the amount of light coming from their
undersides to match the light coming from above. By adjusting their
bioluminescence, they disguise their shadows and become virtually invisible to
predators looking up. (Evers, 2013) Repulsion can be used to indicate toxic or
foul-tasting chemicals that serve as a deterrent to predators, they will light up
to warn predators that they are toxic. Toads, birds, and other predators know
that consuming these larvae will result in illness and possible death. (Evers,
2013)
Illumination, without the intent of attraction, is mostly used in the ocean
that are below the twilight zone. The purpose is to produce light, so that
creatures can attract prey that are sensitive to light, or be able to detect
obstacles in their pathway, facilitating their movements throughout the water.
Some deep sea fish use this to their advantage. Loosejaws have adapted to
emit red light; most fish can only see blue light, so loosejaws have an enormous
advantage when they light up a surrounding area. They can see their prey, but
their prey can't see them. (Evers, 2013) However, illumination and attraction

and communication, are side by side. During mating rituals, the pattern of
glow is used as a source of communication among potential mates, or other
species, to inform on location of mates and food. In the case of fireflies, the
light is used as a signal that is visible throughout the night over long distances
to possible mates. The role of the flashing light of adult fireflies is more
complex, with some species relying purely on pheromones to attract a mate,
while others developing unique patterns and rates of flashing. The flash rate
may change to indicate to a flying male that a female on the ground is
interested in his advances. The length of time between the male flash and the
female response is also used to identify females of the same species. (Biggs,
McPhail, & Nassau, 2016)
The illumination and pattern of these bioluminescent organisms are
essential not only to mating rituals, but how they obtain food as well. For
example, The anglerfish uses bioluminescence to lure prey. [It] has a huge
head, sharp teeth, and a long, thin, fleshy growth (called a filament) on the top
of its head. On the end of the filament is a ball (called the esca) that the
anglerfish can light up. Smaller fish, curious about the spot of light, swim in
for a closer look. By the time the prey sees the enormous, dark jaws of the
anglerfish behind the bright esca, it may be too late. (Evers, 2013)
Bioluminescence has served itself very useful to all kinds of organisms. It
appears mostly in the ocean, however, it can appear in bacteria, fungus and
insects that live on land as well. Bioluminescence is not just visible in the
green-blue hues that are seen in the ocean, but can be emitted in reds, yellows
and violets. The main uses for bioluminescence are camouflage, repulsion,
illumination, communication and attraction. However, this leads to the next
question, which surrounds the process of making the luminescence that is
seen in these organisms.

Which chemical process causes bioluminescence?

Bioluminescence is known as cold light, because it does not produce any
heat, unlike other luminescent chemical or electrical reactions. Because no
heat is produced, the reaction is, by definition, nearly 100% efficient. This
chemical reaction changes per species needs, and the colour frequency it emits.
However, the foundation of a bioluminescent reaction requires, the one which
produces the light called a "luciferin" and the one that drives or catalyzes

the reaction called a "luciferase The luciferase catalyzes the oxidation of

luciferin, resulting in light and an inactive oxyluciferin." (Haddock, McDougall,
& Case, 2016). In each reaction, there are additional proteins that are added to
the luciferin. These proteins help decide the properties the reaction will have.
These proteins separate them into the five different classes of luciferins based
on what proteins are in each reaction. These proteins are; Aldehydes,
Benzothiazoles, Imidazolopyrazines [a.k.a Coelenterazine], Tetrapyrroles and
Flavins. (Lee, 2016) As the name implies, Coelenterazine is the luciferin found
in coelenterates, and in many of the marine organisms.
Luciferins is not made in every bioluminescent organism. Some will
acquire it by consuming a bioluminescent organism, others possess the
capability to create it themselves. In these organisms, there are luminescence
systems, each with their own individual chemical reaction sequence and
proteins. The basic reaction is luciferin and catalyzing protein (the equivalent
of a luciferase), as well as a co-factor such as oxygen, bound together to form a
single unit called a photoprotein. This molecule can be triggered to produce
light when a particular type of ion is added to the system (frequently calcium).
(Haddock, McDougall, & Case, 2016)
These systems are Bacterial, Dinoflagellate, Vargula, Coelenterazine,
Fireflies and other or unknown mechanism. (Haddock, McDougall, & Case,
2016) Bacterial luminescence is commonly found in Bacteria, Pyrosomes, and
some fish and squid. It is made of luciferin and luciferase. Bacterial luciferin is
a reduced riboflavin phosphate (FMNH2, pictured here) which is oxidized in
association with a long-chain aldehyde, oxygen, and a luciferase. (Haddock,
McDougall, & Case, 2016)

Bacterial Luciferin Molecule

Dinoflagellate luciferin is also made of luciferin and luciferase, and is

commonly found in, as the name suggests, dinoflagellates, and Euphausiid

shrimp. Dinoflagellate luciferin is thought to be derived from chlorophyll, and

has a very similar structure. In the genus Gonyaulax, at pH 8 the molecule is
"protected" from the luciferase by a "luciferin-binding protein", but when the pH
lowers to around 6, the free luciferin reacts and light is produced. A modified
form of this luciferin is also found in herbivorous Euphausiid shrimp, perhaps
indicating a dietary link for the acquisition of luciferin. (Haddock, McDougall,
& Case, 2016)

Dinoflagellate Luciferin Molecule

Vargula (Vargulin) luciferin is found in ostracods and in some fish. In

Vargula luminescence there is a clear dietary link, with fish losing their ability
to luminesce until they are fed with luciferin-bearing food. It has been shown
that ostracods synthesize this molecule from the amino acids tryptophan,
isoleucine, and arginine. (Haddock, McDougall, & Case, 2016) This is one of
the systems that shows how not every organism created the luminescence
entirely by themselves, some, if not most, consumed bioluminescent organisms
that would give them the necessary proteins to complete the reaction.

Vargula (Vargulin) luciferin is found in ostracods and in some fish. In Vargula

luminescence there is a clear dietary link, with fish losing their ability to luminesce
until they are fed with luciferin-bearing food. It has been shown that ostracods
synthesize this molecule from the amino acids tryptophan, isoleucine, and
arginine. (Haddock, McDougall, & Case, 2016) This is one of the systems that
shows how not every organism created the luminescence entirely by themselves,
some, if not most, consumed bioluminescent organisms that would give them the

necessary proteins to complete the reaction.

Vargulin Luciferin Molecule

Coelenterazine luciferin is different, in that it is composed of luciferin,

luciferase, and a photoprotein. As one of the most common bioluminescent
marine systems, it can be found in Radiolarians, Ctenophores, Cnidarians,
Squids, Cocepods, Decapod Shrimp, Mysid Shrimp Chaetognaths and some
fish. This molecule can occur in luciferin-luciferase systems, and is famous for
being the light emitter of the photoprotein "aequorin". (Haddock, McDougall, &
Case, 2016)

Coelenterazine Luciferin Molecule

Firefly luciferin, is like the others, made of luciferin and luciferase, and is
found in the organisms that give it the name, fireflies. They have been intensely
studied, therefore they have been given their own category. Firefly luciferin is
used in a luciferin-luciferase system that requires ATP as a cofactor. Because of
this, it can be used as a bio-indicator of the presence of energy or "life".
(Haddock, McDougall, & Case, 2016)

Firefly Luciferin Molecule

Light is composed of a variety of wavelengths, giving

off different colours of the colour spectrum. This is what can
cause specific colours to be produced by certain species.
Visible light is seen in wavelengths of 400-700 nanometers,
whereas bioluminescence is composed of broad bands that
are half the height (60-100 nanometers). In marine
bioluminescent organisms, the light will range from 450-510
nanometers, the blue-green zone. Terrestrial
bioluminescence leans towards the yellow spectrum at 560600 nanometers, of which they are most sensitive towards. (Lee, 2016)

Which adaptations caused bioluminescence to evolve?

While there are many causes that are theorized to be the reason that
bioluminescence evolved from previous species, very little is known about the
true evolutionary path of bioluminescence. There are theories of how the
different uses evolved, to adapt with camouflage, repulsion, illumination,
communication and protection. Others adapted to their changing environment,
supplying them with access to the elements required to produce the chemical
reaction of bioluminescence.
The best example of adaptation that resulted in bioluminescence is the
change from environments that had no oxygen, to ones that did. The
bioluminescent reaction was originally used to remove oxygen toxic to primitive
types of bacteria that developed when oxygen was absent or very rare in Earths
atmosphere. The metabolic reaction that combines the oxygen with a reducing
substance (luciferin) liberates sufficient energy to excite a molecule in the
organism to emit visible radiation. Most of those luminous primitive organisms
subsequently developed systems of using oxygen, but they have retained the
luminescent capability as parts of related metabolic pathways or for some
survival value that luminescence may confer on the organism.
("bioluminescence | chemical reaction", 2016)

As there is no definitive evolution trail that has been proved to be the

cause for bioluminescences existence, it is assumed that the answer lies in the
randomized genetic mutations that occur in nature, as well as natural

selection. Organisms that could survive, reproduce or feed more efficiently

because of their new found mutations, would then survive to pass on the
mutated genes. This explains how only certain geneses of a species can be
bioluminescent, rather than the whole species itself. For example, only certain
beetles can be bioluminescent, not all.
There are many wonders of bioluminescence. Even though the
phenomenon has been studied for centuries; there is much that is still
unknown about bioluminescence. Bioluminescence serves numerous purposes
in the environment, and now, it is being used in modern science to make
advances in science. The main inquiry question that was addressed in this
paper, in what way is bioluminescence a useful trait, was answered with the
three sub questions; how is bioluminescence a useful trait, which chemical
processes cause bioluminescence and which adaptations caused
bioluminescence to evolve? Bioluminescence seems to defy the laws of nature,
therefore it seems only natural to want to learn more about such a wonderful
phenomenon that can bring light to even the darkest of places.

Bibliography:
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Haddock, S., McDougall, C., & Case, J. (2016). The Bioluminescence Web Page.
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from http://photobiology.info/LeeBasicBiolum.html
Meighen, E. (2016). Bioluminescence - Chemistry Encyclopedia - reaction, molecule,
Beetles/Fireflies, Dinoflagellates, Bacteria. Chemistryexplained.com. Retrieved
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