Reported Bioluminescent Organisms on Land and in the Sea

 

M Jyothi, Suneetha V

VIT University, Vellore, Tamilnadu ,India -632014.

 

ABSTRACT:

Some living organisms produce light, thus production of cold light by living organisms is called bioluminescence. Bioluminescence is emission of light as a result of a chemical reaction inside the living organism during the process chemical energy gets converted into light energy. It is a form of chemiluminescence.   Bioluminescent organisms are found in marine and terrestrial environments.  But bioluminescence is most prevalent in marine organisms. Marine bioluminescence is produced by incredible range of organisms. Bioluminescence producing living organisms produce light and utilize it for vital functions ranging from camouflage, illumination, mimicry, attracting prey, attracting mates, for protection and it may be most common form of communication on the planet. bioluminescence responsible genes are not related in bacteria, algae, fishes, beetles and others because it has evolved independently many times. Bioluminescent organisms can be harnessed to produce natural light in the lamps that could light up our planet one day. In addition to that bioluminescent applications can be used to test extra-terrestrial life on other planets so bioluminescence has scope for many futuristic applications. While, bioluminescence research has offered many potential societal benefits in fields ranging from genetic engineering to medicine. By helping to reveal the evolutionary origins of bioluminescence, we can better know how other complex features arise in nature. So there is greater need study bioluminescence and the organisms producing bioluminescence. In this process scientists continue to discover more bioluminescent species.

 

 

 

INTRODUCTION:

Evolutionary origins:

Oxidation of luciferin by luciferase produces bioluminescence. Nonsymbiotic organisms which produce bioluminescence contain genes for luciferase or photoprotein. Bioluminescence occurred independently many times¹. In bacteria it may have evolved once but the organisms which harbour these bacteria developed different ways to produce light. Studies show that bioluminescence evolved more than 50 times in organisms.

This confirms that bioluminescence is important to organisms and its evolution is smooth. This could be credited to readily available prey. Antioxidant molecules in organisms also helps in the evolution of bioluminescence ². After Cambrian period dietary linkages of bioluminescence was established since luciferins should be evolved in prey and then arise in predators³. Ostrocod crustaceans halocyprida and myodocopida utilize two different luciferins diverged 400mya⁴. In many marine organisms’ bioluminescence is primary visual stimulus.

 

Bioluminescence chemistry:

Bioluminescence is the result of oxidation of light emitting luciferins by luciferases or photoproteins. Luciferins responsible for light production are conserved in living organisms. But luciferases and photoproteins are not conserved and have different evolutionary origins. Different organisms use different luciferins for light production some use coelenterazine as luciferin. Coelenterazine is either synthesized or obtained from diet. In bacteria FMNH2 is oxidised to produce light. Luciferin in dianoflagellates is tetrapyrrole and the most extensively used luciferin is coelenterazine.

 

Bioluminescent organisms:

Bioluminescence has been reported across numerous phyla. In majority of these cases, luminescence is not produced by bacterial symbionts   but by the organisms themselves. Few living organisms which are not known to produce bioluminescence are flowering plants, and land vertebrates like amphibians, birds and mammals. Luminescence is generally seen in deep-living and planktonic organisms than in shallow species. The distribution of bioluminescence across the various phyla does not appear to follow any phylogenetic or oceanographic constraint.

 

Bioluminescence is typically created by the oxidation of a light-emitting molecule universally called the luciferin with a catalysing enzyme either a luciferase or photoprotein. Light is produced when luciferin reacts with oxygen. Organisms produce light by ways other than luciferase reactions. Like Chaetopteru has another photoprotein, aequorin, instead of luciferase. In the presence of calcium ions, the aequorin's catalysis produces a brief flash unlike the prolonged glow produced by luciferase. Nonsymbiotic luminous organisms possess the gene for either luciferase or photoprotein.

 

Marine bioluminescence:

Most of the reported bioluminescent organisms are from marine environment. Here bioluminescence is exhibited both by invertebrates and vertebrates. Parasites are the only non-bioluminescent group with hyperiid amphipods as the exception.

 

Bioluminescent Bacteria:

Bioluminescent bacteria are commonly found in the oceans, especially in temperate and warmer waters⁵.

 

Luminous bacteria reach sufficiently high concentrations to initiate quorum sensing, and once generated, they glow continuously rather than producing distinct flashes. These properties are unique to bacteria, which makes them suitable as photogenic symbionts and can lead to magnificent marine development such as bioluminescent seas⁶ milky seas are continuous glows, which can be visible from satellites orbiting the Earth. In bacteria, the Lux operon controls the expression of genes related to bioluminescence.

 

The extensively studied bacterial symbionts are in genus vibrio Certain species of family Vibrionaceae showing bioluminescence are (Vibrio fischeri, Vibrio harveyi) vibrio fischeri lives in symbiotic relation with loliginid squid and monocentrid fish. Photobacterium phosphoreum is a symbiotic bacteria living on marine animals and it is gram negative. photobacterium leiognathi live in symbiotic relation with apogonid and morid fishes⁷.

 

some species of the family Shewanellaceae (Shewanella hanedai, Shewanella woodyi) also show bioluminescence.

 

Bioluminescence in Dinoflagellates:

After fireflies, dinoflagellates are the most commonly seen bioluminescent organism. Moreover, they produce luciferin on their own. Bioluminescence in dinoflagellates is in a bluish-green colour. Bioluminescent dinoflagellates are a type of photosynthetic plankton that can sometimes make the surface of the ocean sparkle at night producing the “bioluminescent bays”⁸.

 

Certain Dinoflagellates (Noctiluca scintillans, Pyrodinium bahamense, Pyrocystis) showing bioluminescence. Dinoflagellates played important role in environmental monitoring, dinoflagellates which are bioluminescent decrease their light emissions in the presence of toxic compounds and heavy metals and they have invaluable in toxicity assays.

 

Bioluminescence in Radiolarians:

Amoeboid protist are called radiolarians, primarily the Polycystinea and Phaeodarea. Within the polycystines, the only order, is known to be bioluminescent is collodaria which includes members Collozoum and Thalassico. Here light emitting substrate is photoproteins attached to coelenterazine.

 

Bioluminescence in Ctenophores:

Planktonic Comb jellies mostly are bioluminescent with the exception of sea gooseberry, pleurobrachia and some benthic species. Ctenophores (comb jellies) like Tortugas red and Mertensia ovum exhibit bioluminescence⁹.

 

Some species like Eurhamphaea vexilligera, Euplokamis stationis, Mertensia ovum also emit glowing particles as part of an escape response. Ctenophores use coelenterazine and some calcium activated proteins as their light emitting substrate.

 

Bioluminescence in Cnidarians:

Bioluminescence is common phenomenon in many deep-sea animals and is wide spread among the cnidarians. Both planktonic and benthic cnidarians exhibit bioluminescence. Bioluminescent cnidarians use coelenterazine as their substrate for emission of light.  Bioluminescent groups include scyphozoa (sea jellies), Hydrozoa (hydroida, siphonophores), Anthozoa (sea pens and sea pansies).

 

Ex: Aatolla wyvillei, Atolla vanhoffeni, Atolla parva, Nausithoe rubra, Errena sp, Periphyllopsis braueri, Periphylla, Paraphyllina intermedia.

 

Sea pens like Ptilosarcus gurneyi, Renilla amethystine are colonial cnidarians which produce bioluminescence. Cyphastrea microphthalma, Moon coral are corals that exhibit bioluminescence.

 

Bioluminescence occurs in Aequorea Victoria also known as crystal jelly¹⁰. The Green Fluorescent Protein (GFP), first found in the Crystal sea jelly. It is the source for noble prize winning GFP¹¹. The GFP has been used to track how cancer cells metastasize, how infections spread and even which male ends up fertilising a female fruit fly’s egg. Cnidarians use bioluminescence for attracting prey and for defensive purposes.

 

Bioluminescence in Annelid:

Marine polychaetes have different bioluminescent lineages and luminescence have many independent origins. In Odontosyllis females produce luminous secretions during spawning. In Eusyllis, fragments continue to luminescing for weeks even after head is detached. The tube-dwelling chaetopterid Chaetopterus and the terebellid Polycirrus both produce light^12. Glowing particles are released from their tubes when the worms are disturbed. Most of them produce internal bioluminescence when disturbed¹³.

 

In the planktonic polychaetes species of Tomopteris produces yellow light. other planktonic members like Poeobius meseres and Flota vitjasi, carries bioluminescent bombs, it drops when it is disturbed.

 

Bioluminescence in Chaetognatha:

bioluminescent organs in Caecosagitta marcocephala and Eukrohnia fowleri are present on their fins. Chaetognatha make use of luciferase and coelenterazine to shed light during escaping response¹⁴.

 

Bioluminescence in Molluscs:

Bioluminescence has evolved several times in this phylum alone. Certain Clams, Nudibranchs, exhibit bioluminescence. Luminous marine molluscs include a few atypical gastropods like the whelk Planaxis and the amazing nudibranch Phylliroe. One of the longest-known and best studied luminous molluscs is the bivalve pholas.  Pholasin was further studied and characterized¹⁵.

Cephalopods are the most important bioluminescent marine molluscs. In the group squids alone, there are at least 70 luminous genera. Bacterial symbionts produce luminescence in some genera in the family’s Sepiolidae and Loliginidae. Sea snail Hinea brasiliana shows bioluminescence. Only Land snail Quantula striata and only fresh water snail Latia neritoides show bioluminescence. Certain octopuses from order Bolitaenidae exhibit bioluminescence Bioluminescence is seen in Vampire squids and Bobtail squids of order Sepeolida. Bioluminescence also occurs in squids like colossal squid, sparkling enope squid and whip-lash squid of family mastigoteuthidae

 

Bioluminescence in Crustaceans:

Ostracods, Copepods, Krill exhibit bioluminescence. Krill use physical mechanisms to control light production¹⁶. Dramatic displays of luminescence at dusk are often attributable to ostracods.

 

Copepods, are one of the most abundant marine invertebrates, in addition to that they are the most abundant bioluminescent groups in the ocean, while the widespread genus Calanus is not luminescent. Common luminous species include Euaugaptilus magnus, pleuromamma and Gaussia.

 

Bioluminescence in Echinoderms:

Major groups of echinoderms like Ophiuroid, Asteroid (Brisingidae and paxillosida) olothuroid (pannychia, scotoanassa) and crinoid (Annacrinus and thaumatocrinus) show bioluminescence¹⁷. In which it has an anti-predator effect. Ex: Amphipholis squamata. Echinoderms use unknown photoproteins and coelenterazine luciferase as light emitting substrates.

 

Bioluminescence in Tunicates:

Bioluminescence is seen colonial salp pyrosoma. Further studies showed bacteria was involved in light production. Some of the other species showing luminescence are Paradoliopsis harbisoni, Clavelina miniata, Pseudusa bostigrinus. Planktonic larvaceans (Appendicularia) also reported to show bioluminescence.

 

Bioluminescence in Fish:

Bioluminescence is found in at least 42 families of bony fishes¹⁸, in addition to one family of sharks. In contrast to invertebrate taxa, several of these fishes use bacterial symbionts for light production, including the well-known anglerfishes¹⁹.

Lantern fish (Symbolophorus barnardi)

Gulper eel (Eurypharynx pelecanoides)

Viperfish (Chauliodus sloani)

Black dragonfish (Grammatostomias flagellibarba)

Flashlight fish (Photoblepharon palpebratum)

Midshipman fish (Porichthys notatus)

Anglerfish (Centrophryne spinulosa)

Catshark (Scyliorhinus meadi)

Pinecone fish (Monocentris japonica)

Cookiecutter shark (Isistius brasiliensis)

Marine hatchetfish (Argyropelecus gigas)

 

In the order Chondrichthyes, lantern sharks of Squalidae family use bioluminescence for both defensive and offensive purposes.

 

Bioluminescence in Terrestrial animals:

In the terrestrial environment, bioluminescence is sparse than in the sea.  Terrestrial bioluminescence is observed in various organisms which include bacteria, fungi, mollusc’s, arthropods and annelids. It is probable that more luminescent species are hidden in the layers of soil, where the darkness is as constant as in the deep oceans. unlike marine species, which live in constant darkness, terrestrial bioluminescent species are active during night. Luminescence is used for a variety of functions like for attracting mates, for luring prey, for defense and illumination.

 

In the marine environment the predominant bioluminescence colour is blue, whereas in the terrestrial environment, green is the main bioluminescence colour.

 

Bacteria showing bioluminescence:

Only few bioluminescent bacteria are reported from terrestrial environment. Photorhabdus luminescens is one of the luminous bacteria that lives in the gut entomopathogenic nematode ²⁰.

 

Fungi showing bioluminescence:

We recognize numerous species of bioluminescent fungi belonging to Omphalotus, Armillaria and mycenoid lineages. Here luminescence plays an important role in the propagation and dispersal of species²¹.

Armillaria species,

Mycena species,

Dictyopanus follicollus,

Favolaschia manipularis,

Filoboletus species,

Gerronema viridilucens,

Nothopanus species,

Omphalotus species,

Panellus species,

Pleurotus decipiens,

Roridomyces species,

Xylaria hypoxylon.

 

Bioluminescence in Molluscs:

Bioluminescence is seen in land snail Quantula striata. The limpet, Latia neritoides, is the only reported fresh-water bioluminescent organism. This small snail lives in streams in New Zealand.

Bioluminescence in Annelids:

some annelids show bioluminescence including earthworms. Three families out of 16 families in Oligochaeta have bioluminescent species. A number of species are found in the soil around the world. Diplocardia longa is a bioluminescent worm, which has been extensively studied. The luminescence is produced by the coelomic fluid. Luminescence colour differs from blue to orange depending on the organism. A member of Emlea (Enchytraeidae) also displays luminescence from the coelomic fluid.

 

Bioluminescence in Arthropods:

Arthropods has the largest number of luminescent species, most of them found in insects. Today, thousands of luminescent species are found in the orders Collembola, Diptera and mainly Coleoptera. species of luminescent cockroaches (Orthoptera) were recently discovered in the Amazon forest. In Diptera, luminescence is found in Mycetophilidae²². bioluminescent species are found in the Australasian Arachnocampa spp, Keroplatus spp and Orfelia fultonii. The order of Coleoptera comprise most of the bioluminescent species, mainly in the family Elateroidea, which includes fireflies (Lampyridae), click beetles (Elateridae) and railroadworms (Phengodidae).

 

Centipedes like Geophilus carpophagus and millipedes like Motyxia exhibit bioluminescence. Motyxia bistipita was discovered recently in California. Another millipede Luminodesmus sequoia lives in the mountain forests of California, and emit greenish light through the body.

 

CONCLUSIONS:

Bioluminescent organisms are selected for many areas of research. Bioluminescence applications are widely used in the field of genetic engineering²³. Bioluminescence imaging has emerged as powerful tool in analysing infectious diseases in biomedical research.  Assays using bacterial bioluminescence for the detection and classification of microbes is rapid, sensitive, accurate and specific. while exploring potential new application such as creating plants that luminesce when they need water, the light producing organs in bioluminescent organisms are being explored by industrial designers. integrated bioluminescence could perhaps one day be used to reduce the need for street lighting, or decorative lighting²⁴. The gene that makes fireflies glow has been added to mustard plants. The plants glow faintly for an hour when touched. Some research is going on to use of genetically engineered bioluminescent E. coli bacteria, as bioluminescent bacteria in a light bulb. The Glowing Plant project raised nearly $500,000 on the crowd funding site Kickstarter to create a bio luminescent plant.

 

The following are the few topics of bioluminescence for future research

 

The process of luciferin synthesis is almost entirely unknown. This will be a promising area for future research.

Better access to live animals in their habitat will give opportunities to understand natural functions of luminescence. It may also provide answers to why so many animals are bioluminescent.

To study the chemistry of luminescence for many organisms which remained completely unknown. Promising taxa include echinoderms, polychaetes, and tunicates.

Improvements in remote and automated methods of detecting oceanic bioluminescence will make us understand marine ecosystem dynamics.

 

On the other hand, bioluminescence must provide important ecological roles, given its overall distribution in the ocean. Many interesting questions remain to be answered²⁵.

 

REFERENCES:

1.     Rees JF, De Wergifosse B, Noiset O, Dubuisson M, Janssens B, Thompson EM. 1998. The origins of marine bioluminescence: Turning oxygen defence mechanisms into deep-sea communication tools. J. Exp. Biol. 201:1211–21

2.     Labas YA, Matz MV, Zakhartchenko VA. 2001. on the origin of bioluminescent systems. In Bioluminescence and Chemiluminescence 2000, ed. JF Case, PJ Herring, BH Robison, SHD Haddock, LJ Kricka, PE Stanley, pp. 91–94.

3.     Yamaguchi S, Endo K. 2003. Molecular phylogeny of Ostracoda (Crustacea) inferred from 18S ribosomal DNA sequences: implication for its origin and diversification. Mar. Biol. 143:23–38

4.     Tinn O, Oakley T. 2008. Erratic rates of molecular evolution and incongruence of fossil and molecular divergence time estimates in Ostracoda (Crustacea). Mol. Phylogenet. Evol. 48:157–67

5.     Dunlap PV, Kita-Tsukamoto K. 2006. Luminous bacteria. Prokaryotes 2:863–92

6.     Nealson KH, Hastings JW. 2006. Quorum sensing on a global scale Bioluminescent bacteria make milky seas. Appl. Environ. Microbiol. 72:2295–97

7.     Kaeding AJ, Ast JC, Pearce MM, Urbanczyk H, Kimura , et al. 2007. Phylogenetic diversity and cosymbiosis in the bioluminescent symbioses of “Photobacterium mandapamensis.”Appl. Envir. Microbiol. 73:3173–82

8.     Latz MI, Jeong HJ. 1996. Effect of red tide dinoflagellate diet and cannibalism on the bioluminescence of the heterotrophic dinoflagellates Protoperidinium spp. Mar. Ecol. Prog. Ser. 132:275–85

9.     Steven H.D. Haddock, Mark A. Moline, and James F. Case Bioluminescence in the Sea, Annual Review of Marine Science, Vol. 2: 443-493 (Volume publication date January 2010)

10.   Prasher DC, Eckenrode VK, Ward WW, Prendergast FG, Cormier. 1992. Primary structure of the Aequorea Victoria green fluorescent protein. Gene 111:229–33

11.   Shimomura O. 2005. The discovery of aequorin and green fluorescent protein. J. Microsc. 217:3–15 Bradner H, Bartlett M, Blackinton G, Clem J, Karl D, et al. 1987. Bioluminescence profile in the deep Pacific Ocean. Deep-Sea Res.34:1831–40

12.   Fischer A, Fischer U. 1995. On the life-style and life-cycle of the luminescent polychaete Odontosyllis enopla (Annelida: Polychaeta). Invert. Biol. 114:236–47

13.   Deheyn DD, Latz MI. 2009. bioluminescence of the marine polychaete Odontosyllis phosphorea (Syllidae). Invertebrate Biol. 128:31–45

14.   Case JF, Haddock SHD, Harper RD. 1994. The ecology of bioluminescence. In Bioluminescence and Chemiluminescence. Fundamentals and Applied Aspects, ed. AK Campbell, LJ Kricka, PE Stanley, pp. 115–22. New York: Wiley

15.   Dunstan SL, Sala-Newby, Fajardo, Taylor, Campbell AK. 2000. Cloning and expression of the bioluminescent photo protein pholasin from the bivalve mollusc Pholas dactylus. J. Biol. Chem. 275:9403–9409

16.   .Krönström J, Karlsson W, Johansson BR, Holmgren S. 2009. Involvement of contractile elements in control of bioluminescence in Northern krill, Meganyctiphanes norvegica (M. Sars). Cell Tissue Res. 336:299–308

17.   Herring PJ, Cope C. 2005. Red bioluminescence in fishes: on the suborbital photophores of Malacosteus, Pachystomias and Aristostomias. Mar. Biol. 148:383–94

18.   Haygood MG, Prince RC. 1993. Light organ symbioses in fishes.Crit. Rev. Microbiol. 19:191–216

19.   Suntsov AV, Widder EA, Sutton TT. 2008. Bioluminescence in larval fishes. In Fish Larval Physiology, ed. RN Finn, BG Kapoor, pp. 51–88. Bergen, Norway

20.   Wilson T. and Hastings J. W. (1998) Bioluminescence. Annu. Rev. Cell Dev. Biol. 14:197-230.

21.   Desjardin, D., Oliveira A.G., Stevani C.V. Fungi Bioluminescence revisited. Photochem. Photobiol. Sci 7: 170.

22.   Viviani, V. R. (2002) The origin, diversity and structure function relationships of insect luciferases. CMLS59:1833-1850 

23.   Steven H.D. Haddock, Mark A. Moline, and James F. Case Bioluminescence in the Sea, Annual Review of Marine Science, Vol. 2: 443-493 (Volume publication date January 2010)

24.   Steven D Miller, Steven H. D. Haddock, Christopher D. Elvidge and Thomas F Lee Detection of Bioluminescent Milky sea from space, Proceedings of the National Academy of Sciences of the United states of America.

25.   Herring PJ. 1987. Systematic distribution of bioluminescence in living organisms. J. Biolum. Chemilum. 1:147–63

 

 

 

 

Accepted on 29.06.2017                  © RJPT All right reserved