Bioluminescence: Living Light

Written by Francesca Aguirre-Wong BIOL3090-1

I. ABSTRACT

Bioluminescence is an emission of light by a living organism. It is a phenomenon that has

captivated people for thousands of years, from Darwin all the way back to Aristotle.

Bioluminescence of an organism can occur via a highly complicated symbiosis between an

organism and a bacterial bioluminescence, or in single cells that contains all the tools needed for

a constant glow. There are also many that are intermediates of these two groups. The female

anglerfish attracts prey by dangling and twitching a glowing orb in front of its mouth. This lure is

actually a modified dorsal fin spine containing dense packet of bioluminescent bacteria which

holds a symbiotic relationship with the fish. On the other hand, single-celled dinoflagellates are

marine plankton that responds to mechanical stimulation when the water is disturbed by emitting

a brief bright flash. Bioluminescence is important because of the symbiotic relationship that

some species have with larger organisms, tourism, and the proteins that can be extracted from

organisms for use in the medical field (Haygood et al 2000).

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 II. WHY THIS TOPIC WAS CHOSEN

In December 2009, off the coast of the Adang Archipelago in Southern Thailand, I went on

my first night dive to explore the nocturnal behavior of organisms in coral reefs. We began from

the beach and snorkeled with apprehension, not knowing what sort of sea monsters lurked in the

places our flashlights couldn’t span. After our teacher signaled to turn our flashlights off, I saw

each light go out one by one until mine was the last to put us in darkness.

What happened next was the most psychedelic display of fireworks I had ever seen. The

ominous black sea lit up with blue sparkles after every movement that we made, painting a

milky-way scene for our perplexed eyes. In the four months I spent in Thailand, this 30 minute

night dive with the phosphorescence was one of the highlights of my stay. It was a spiritual

experience that sparked my curiosity in, as quoted by Aristotle 2,500 years ago, “things which

are neither fire nor forms of fire [that] seem to produce light by nature” (Pieribone 2005). It is

these bioluminescent creatures of the sea that inspired me to choose the topic for this paper.

III. INTRODUCTION

Bioluminescence is an emission of light by a living organism that is visible to other

organisms. This “glowing light” is derived from an enzyme-catalyzed chemiluminescence—a

highly exergonic reaction in which chemical energy is transformed into light energy (Hastings

2004). All bioluminescent reactions involve the oxidation of luciferin by an enzyme (luciferase)

to produce an electronically excited state and an inactive oxyluciferin. More luciferin is required

to be brought into the system, through diet or internal synthesis, to repeat this oxidation-

reduction reaction.

Luminescence of an organism can occur in diverse forms of morphology with various

different mechanisms of light emission. Some organisms have highly complicated light organs,

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such as eyes, in which light emission is controlled by their nerve systems and luminescence is

emitted as needed or by stimulation (ex. luminous fish and squids). In some others, single cells

contain all the needed tools (luciferin and luciferase) for light emission and light is emitted

continuously (luminous bacteria and fungi). There are also many intermediates of these two

groups (Shimomura 2006). It is amazing that animals and plants have developed their functional

abilities of bioluminescence which integrate chemistry, physics, morphology, and physiology.

Bioluminescence is usually a rare trait in land dwellers, but in the deep dark ocean more than

90 percent of animal species are capable of generating light (Smith et al 1987). Since most of the

ocean’s volume lies below the reach of the sun’s rays, marine animals that can produce their own

personal flashlights have an advantage. Bioluminescence can aid in the search for prey by

attracting food with bioluminescent lures or use the light to scan the darkness. Bioluminescence

can also deter an enemy with a blinding or distracting flash, or creatures can use species-specific

light shows to attract mates.

The focus of this paper will be on marine bioluminescent organisms, specifically,

dinoflagellates and bioluminescence due to the presence of symbiotic luminous bacteria. These

microorganism’s morphology, physiology, genetics, and relationship with their environment will

be discussed using references that are given in alphabetical order at the end of the paper.

IV. DISCUSSION

a. Dinoflagellates

Ocean “phosphorescence,” commonly seen at night when the water is disturbed, is due in

large part to the bioluminescence of dinoflagellates. These organisms occur everywhere in the

oceans as planktonic forms and respond to mechanical stimulation when the water is disturbed,

such as by waves or fish swimming, by emitting brief bright flashes. Luminescent dinoflagellates

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occur primarily in surface waters because many are photosynthetic. The so-called red tides are

temporal blooms of individual dinoflagellate species that can emanate an unpleasant odor as well

as toxins which can kill fishes. An example of a non-toxic bloom that occurs in New Zealand is

Noctiluca scintillans.

Luminescence in dinoflagellates is emitted from many small (~0.5μm) cortical organelle

flashing units called scintillons (Hastings 2004). They occur as out-pocketings of the cytoplasm

into the cell vacuole, like a balloon, with the neck staying connected. Scintillons contain only

dinoflagellate luciferase, luciferin, and a luciferin binging protein that keeps it from reacting with

luciferase in between flashes. Other cytoplasmic components are excluded from scintillons,

which can be identified ultrastructurally by immunolabling with antibodies raised against the

luminescence proteins. Dinoflagellate luciferin is a highly reduced novel tetrapyrrole related to

chlorophyll and in extracts remains tightly bound to a ~75-kDa specialized protein at

cytoplasmic pH 8. The luciferase, a large single polypeptide chain of about 136kDa, is also

inactive at pH 8. Activity may be obtained by shifting the pH of the environment from pH 8 to

pH 6. At pH 6 the luciferin is released from its binding protein and the luciferase assumes an

active conformation. The pK for both luciferase and luciferin is at pH 6.7. The flashing of

dinoflagellates is initiated by mechanical shear or cell stimulation which results in the generation

of conducted action potential in the vacuolar membrane. As the action potential traverses the

vacuolar membrance it sweeps over the scintillons, opening voltage-gated ion channels, thus

allowing protons from the acidic vacuole to enter, causing a transient pH change in the

scintillons and thus a flash (Fogel et al 1972).

Since dinoflagellates are stimulated to emit light when predators (ex. crustaceans) are active,

predators might thereby be alerted to feed on crustaceans, resulting in a reduced predation on

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dinoflagellates in general. Dinoflagellates also avoid predation by directly using their flash to

startles or divert a predator. Unlike bacteria, luminous dinoflagellates are not known to have

symbiotic relationships based on their light emission. However, dinoflagellates are important

symbionts for contributing photosynthesis and carbon fixation in certain animals.

b. Luminous bacteria

Luminous bacteria are widely distributed in the marine environment and are generally in

the genera Photobacterium, Beneckea, and Vibrio. They are often found in some kind of

symbiotic association with higher organisms (ex. fish or squid), in which the light emission is of

functional importance to the host. The female deep sea angler fish (Melanocetus johnsoni)

attracts prey by dangling and twitching a glowing orb in front of its mouth. The lure is actually a

modified dorsal fin spine containing a dense packet of bioluminescent bacteria (Vibrio). These

bacteria are located between irregular in-foldings of the plasma membrane of the glandular cells

inside the esca (luminescent bait). Studies have shown that the absence of bacteria membranes

suggests some measure of obligate parasitism between the bacteria and angler fish’s glandular

cells (Hulet et al 1968). While the production of light by these luminescent bacteria results from

a chemiluminescence reaction, it is uncertain how the angler fish exerts biochemical control on

light emission (Herring et al 1994).

The symbiosis between the deep sea angler fish and the luminescent bacteria Vibrio arose

when free-living Vibrios developed a stable association with their host. The present obligate

symbiosis had developed from a facultative one. The obligate bacteria are very specialized for

the symbiotic habitat and lost the traits necessary to adapt to varying conditions (Nishiguchi et al

1998). Now the angler fish can use the luminescent bacteria to reel in prey, and the bacteria are

given a hospitable environment to live inside the esca.

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 V. UNANSWERED QUESTIONS

While writing this paper, I found it difficult to focus on one luminescent organism since there

are so many intriguing bioluminescent systems that are not related to each other evolutionarily.

For example, both deep sea angler fish and pyrosomes both have symbiotic relationships with

bioluminescent bacteria, though they have very different morphologies and physiological

relationships with their symbiont. The diversity in the chemical reactions, enzymes, mechanisms,

and microbe-environment interactions of the various luminescent organisms was somewhat

overwhelming and difficult to summarize in a short page paper. I also found that not all

bioluminescent mechanisms and symbiotic relationships are clearly understood by scientists,

which made it difficult to explain the symbiosis between luminescent bacteria and deep sea

angler fish. After writing this paper, I realized I have only begun to understand the

bioluminescent world.

I would still like to research more about the color emission by bioluminescent organisms. It

would be interesting to see why the color blue is emitted by dinoflagellates and not green, pink,

or some other color. Uses for bioluminescence in the field of science and conservation would

also be an interesting topic to explore. Aside from the Green Fluorescent Protein (GFP) which

has been extracted from jellyfish to create fluorescent micrographs that aid in treating

Alzheimer’s disease, I am sure that bioluminescent organisms can further aid in the field of

medicine.

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VI. LITERATURE CITED

Fogel, M. and J.W. Hastings 1972. "Bioluminescence: Mechanism and Mode of Control of

Scintillon Activity." Proc. Nat. Acad. Sci. USA 69.3: 690-93.

Herring, P.J. and O. Munk 1994. “The Escal Light Gland of the Deep-Sea Anglerfish

Haplophryne mollis (Pisces: Ceratioidei) with Observations on Luminescence Control.”

Journal of the Marine Biological Association of the United Kingdom 74:747-763.

Hastings, J. W. 2004. "Ch. 14 Bioluminescence, Microbial." The Desk Encyclopedia of

Microbiology. By Moselio Schaechter and Joshua Lederberg. Amsterdam; Boston:

Elsevier/Academic. 180-88.

Haygood, M. G., and S. Allen 2000. "Ch.19 Luminous Bacteria." Journey to Diverse Microbial

Worlds: Adaptation to Exotic Environments. Ed. Joseph Seckbach. Vol. 1. Dordrecht:

Kluwer Academic Publ. 269-85.

Hulet, W. H., and G. Musil 1968. "Intracellular Bacteria in the Light Organ of the Deep Sea

Angler Fish, Melanocetus Murrayi." Copeia 1968.3: 506-12.

Nishiguchi, M.K., E.G. Ruby, and M.J McFall-Ngai 1998. "Competitive Dominance among

Strains of Luminous Bacteria Provides an Unusual Form of Evidence for Parallel

Evolution in Sepiolid Squid-Vibrio Symbioses." Applied and Environmental

Microbiology 64.9: 3209-213.

Pieribone, V., and D. F. Gruber 2005. Aglow in the Dark: the Revolutionary Science of

Biofluorescence. Cambridge, Mass.: Belknap of Harvard UP.

Shimomura, O. 2006. Bioluminescence: Chemical Principles and Methods. Hackensack, N.J.:

World Scientific.

Smith, D.C., and A.E. Douglas 1987. The Biology of Symbiosis. London: Edward Arnold. 224.