Glowing In the Woods – what is it?

1. Glowing In the Woods – what is it? Biolumenescence -- terrestrial organisms

2. Bioluminescence: (Bio = life; luminescence =chemical light energy) Many different organisms are endowed with the ability to emit light . These luminescent organisms have been observed for ages. The first scientific attempts to understand the origin in luminescence began in the 1600’s. The first studies determined the “light” was dependent on the air (oxygen).1

3. Luciferin • Identified as the light emitting molecule • Is oxidized in the presence of the enzyme luciferase* • Utilizes energy; ATP (adenosine triphosphate) in the reaction • Utilizes oxygen in the reaction *Luciferase is an oxygenase optimized for light emission that catalyzes the oxidation of luciferin - to obtain oxyluciferin in the singlet-excited state.2

4. Chemical Reaction for luminescence Firefly luminescence Luciferin consists of a benzothiazole moiety attached to a thiozole carboxycilic acid 2

5. Terrestrial organisms identified as being bioluminescent include: Bacteria Fireflies other Insects/larvae Fungi

6. USES OF LUMINESCENCE • Communication • Finding or Attracting Prey • Defense Against Preditors

7. FIREFLIES Use bioluminescenc for: COMMUNICATION Males and females exchange flashes for the purpose of attraction and mating. The female sits on the ground to flash to a certain selected male only. She chooses the male based on his flash pattern.

8. Firefly signal regulation The female firefly signals her response to the males that have the longer and faster flashes. This female choice favors the most conspicuous flashes.3 Scientists question why there is not more of an evolutionary adaptation towards a longer and quicker flash. Why don’t we see the majority of the males with this trait.3 The answer to this question may be a survival mechanism.

9. The predator- Photuris The firefly signaling not only attracts potential mates, But also attracts the bigger beetle-Photuris Photuris mimics signals of the firefly and essentially attracts her next meal.

10. Bacteria Bacteria identified as being luminescent belong to the following genera: • Photobacterium • Vibrio • Photorhabdus

11. Bacteria use luminescence for: Symbiosis- requiring unique nutritional supplements, the bacteria grow on the host ; the host gains the bioluminescent properties of the bacteria4. The host organism uses this new found property for communication within its own species interactions.

12. Bacterial Luciferase and the Lux Gene For continuous light production in bacteria includes not only bacterial luciferase, but also the enzymes that supply and regenerate the substrates of bacterial luciferase.4 The DNA sequences coding the proteins in the luminescent system are termed the lux genes. Only 5 genes in the lux operon (luxCDABE) are needed to produce light emission.4

13. The substrates of bacterial luciferase are reduced flavin mononucleotide (FMNH2 ), molecular oxygen, and long chain fatty aldehydes.4 http://www.modares.ac.ir/sci/saman_h/Pages/bacterial.htm http://www.modares.ac.ir/sci/saman_h/Pages/bacterial.htm

14. The Luminous glow of bacterial colonies is explained: These bacteria utilize FMNH2 , O2 , and long fatty aldehyde as substrates for the bioluminescence reaction catatlyzed by luciferase (LuxAB) with the fatty acid resiductase complex (LuxCDE) synthesizing the long chain aldehyde substrate of tetradecanal. A NAD(P)H-dependent FMN reductase found in most, if not all, bacteria generates the FMNH2 substrate.4

15. Fungus use bioluminesce for: Finding and attracting insects; not as prey but as a means of dispersing fungal spores.5 Like the firefly, the use of bioluminescence not only attracts the wanted insects, but the predatory ones as well. The fungus is subject to organisms who can destroy their very essence.

16. Fungal Families Exhibiting Bioluminesce: • Myxomycetes (slime molds) • Schizomycetes (bacteria) • Phycomycetes (molds) • Ascomycetes (yeasts, sac fungi and some molds) • Basidiomycetes (smuts, rusts, and mushrooms) http://news.nationalgeographic.com/news/2006/10/photogalleries/glowing-fungi/photo2.html

17. Fungi The “glowing in the woods” or Foxfire, has been discussed as far back as Aristotle in 382 B.C.5 Fungal bioluminescent studies have been conducted in the sixteenth, seventeenth, and eighteeth centuries.5 Studies have identified many fungal species, attached to rotting and decaying wood, as having bioluminescent properties.

18. Other Insects/Larvae with Bioluminesce • Phengodidae beetles • Luminodesmus millipedes • Keroplatus fungus gnats

19. Phengodidae: Beetles use bioluminescence as a defensive mechanism. The adult female remains in larvaeform.6 The light is also used as a sexual attractant from the female to male of the species. MALE BEETLE FEMALE BEETLE

20. Lampyridae larvae (glowworm) use luminescence as a signal for unpalability to the nocturnal, visually guided predators. Example: Wild toads (Bufo Bufo) are reluctant to attack luminescent glow worm larvae.6

21. Luminodesmus: The millipede is blind and therefore can not utilize bioluminescence for visual attraction of the species. Its luminance is used as a warning signal known as Aposematism, and it will additionally release HCN which produces a pungent odor. It is noted, that this type of bioluminescence is of no voluntary control, and the light glow is continual.

22. Keroplatus: The Japanese fungus gnat spins a web festooned with poisons and adhesives and block predatory or parasitic enemies.7 Another example of luminescence used for an Aposematism. Defense from predators.

23. SUMMARY: Terrestrial bioluminescence: “Glowing in the Woods” is demonstrated by many species of organisms. Bioluminescence used for communication, attraction, or defense, has been observed for many centuries. Currently, bioluminescence is being utilized in the research laboratories for biochemical toxicity studies, and chemiluminescence for medical imaging techniques. The “ glowing” phenomena exhibits wonder for all.

24. REFERENCES: 1. Biron, K., Fireflies, Dead Fish and a Glowing Bunny: a Primer on bioluminescence. BioTeach Journal. April 2003; Vol. 1: pgs. 19-26. Internet: retrieved October 11, 2007 from www.bioteach.ubc.ca. 2. Roda, A. [Guardigli, M., Michelini, E., Mirasoli, M., Pasini, P.] Analytical Bioluminescence and Chemiluminescence. Analytical Chemistry. American Chemical Society. November 2003; pgs. 463-470. Internet: retrieved October 11, 2007 from http://avogadro.chem.iastate.edu/CHEM513/1103feature _roda%20chemiluminescence.pdf. 3. Tufts University. Biologists Expose Hidden Costs of Firefly Flashes: Risky Balalnce Bewteen Sex and Death. Science Daily. September 2007; Internet: retrieved October 11, 2007 from http://www.sciencedaily.com/releases/2007/09 /070919164811.htm. • Yen-Cheng Lin, L. [Meighen, E.] The Biochemistry and Molecular Biology of Bacterial Bioluminescence. Tarbiat Modares University. January 2004; Internet: retrieved October 11, 2007 from http://www.modares.ac.ir/sci/saman_h / Pages/bacterial.htm.

25. REFERENCES (CONT.) 5. Sivinski, J. Phototrophism, Bioluminescence, and Diptera. U.S. Department of Agriculture, Center for Medical, Agricultural, and Veterinary Entomology. Florida Entomologist. September 1998; vol. 81: n.3 pg. 282. Internet: retrieved October 11, 2007 from http://www.fcla.edu/FlaEnt/fe81p282.html. 6. De Cock, R. [Matthysen, E.] Glow-worm larvae bioluminescence (Coleoptera: Lampyridae) operates as an aposematic signal upon toads (Bufo Bufo). Behavioral Ecology. 2003; vol 14: n. 1 pgs. 103-108. Internet: retrieved October 11, 2007 from http://beheco.oxfordjournals.org/cgi/content/abstract/14/1/103. 7. Davenport, D., [Wootton, D.,Cushing, J.]. The Biology of the Sierra Luminous Millipede. Luminodesmu sequoiae, Loomis and Davenport. University of California. No year given. pgs.100-111. Internet: retrieved October 11, 2007 from http://www.biolbull.org/cgi/reprint/102/2/100.pdf