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 luciferases are often distinguished by including the species or group, i.e. Firefly 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 animal groups, especially in marine environments where dinoflagellates cause 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 plant has been created.
E. N. Harvey (1932) was among the first to propose how bioluminescence could have evolved. 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). 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. 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. 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. 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. 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.