Electric organ (biology)
In biology, the electric organ is an organ common to all electric fish used for the purposes of creating an electric field. The electric organ is derived from modified nerve or muscle tissue. The electric discharge from this organ is used for navigation, communication, defense and also sometimes for the incapacitation of prey.
In the 1770s the electric organs of the torpedo and electric eel were the subject of Royal Society papers by Hunter, Walsh and Williamson. They appear to have influenced the thinking of Luigi Galvani and Alessandro Volta - the founders of electrophysiology and electrochemistry.
In the 19th century, Charles Darwin discussed the electric organ in his Origin of Species as a likely example of convergent evolution: "But if the electric organs had been inherited from one ancient progenitor thus provided, we might have expected that all electric fishes would have been specially related to each other…I am inclined to believe that in nearly the same way as two men have sometimes independently hit on the very same invention, so natural selection, working for the good of each being and taking advantage of analogous variations, has sometimes modified in very nearly the same manner two parts in two organic beings".
Electrocytes, electroplaques or electroplaxes are cells used by electric eels, rays, and other fish for electrogenesis. They are flat disk-like cells. Electric eels have several thousand of these cells stacked, each producing 0.15 V. The cells function by pumping positive sodium and potassium ions out of the cell via transport proteins powered by adenosine triphosphate (ATP). Postsynaptically, electrocytes work much like muscle cells.[clarification needed] They have nicotinic acetylcholine receptors.
These cells are used in research because of their resemblance to nerve-muscle junctions.
The stack of electrocytes has long been compared to a voltaic pile, and may even have inspired the invention of the battery, since the analogy was already noted by Alessandro Volta. While the electric organ is structurally similar to a battery, its cycle of operation is more like a Marx generator, in that the individual elements are slowly charged in parallel, then suddenly and nearly simultaneously discharged in series to produce a high voltage pulse.
To discharge the electrocytes at the correct time, the electric eel uses its pacemaker nucleus, a nucleus of pacemaker neurons. When an electric eel spots its prey, the pacemaker neurons fire and acetylcholine is subsequently released from electromotor neurons to the electrocytes, resulting in an electric organ discharge.
In most fishes, electric organs are oriented to fire along the length of the body, usually lying along the length of the tail and within the fish's musculature, with smaller accessory electric organs in the head. However, there are some exceptions; in stargazers and in rays the electric organs are oriented along the dorso-ventral (up-down) axis. In the electric torpedo ray, the organ is near the pectoral muscles and the gills (see the image). The stargazer's electric organs lie between the mouth and the eye. In the electric catfish, the organs are located just below the skin and encase most of the body like a sheath.
Electric organ discharge
Electric organ discharge (EOD) is the electric field generated by the organs of animals including electric fish. In some cases the electric discharge is strong and is used for protection from predators; in other cases it is weak and it is used for navigation and communication. Communicating through EODs occurs when a fish uses its own electroreceptors to sense the electric signals of a nearby fish. Electric fish navigate by detecting distortions in their electrical field by using their cutaneous electroreceptors 
- Review Article on the molecular evolution of the electric organ.
- Phylogeny of weakly electric fishes.
- Comprehensive overview of electric organ function and evolution in weakly electric fishes.
- Succinct Nature article describing differences in gene expression between sarcomeres and electrocytes of electric organs.
- Kramer, Bernd (1996). "Electroreception and communication in fishes" (PDF). Progress in Zoology. 42.
- Castello, M. E., A. Rodriguez-Cattaneo, P. A. Aguilera, L. Iribarne, A. C. Pereira, and A. A. Caputi (2009). "Waveform generation in the weakly electric fish Gymnotus coropinae (Hoedeman): the electric organ and the electric organ discharge". Journal of Experimental Biology. 212 (9): 1351–1364. PMID 19376956. doi:10.1242/jeb.022566.
- Alexander Mauro, "The role of the voltaic pile in the Galvani-Volta controversy concerning animal vs. metallic electricity" in Journal of the History of Medicine and Allied Sciences, volume XXIV, number 2, April, 1969 available online at jhmas.oxfordjournals.org/cgi/reprint/XXIV/2/140.pdf
- Darwin, C. (1859). On the origin of species by means of natural selection, or the preservation of favoured races in the struggle for life. London: John Murray. ISBN 1-4353-9386-4.
- Lissmann, H. W. (1951). "CONTINUOUS ELECTRICAL SIGNALS FROM THE TAIL OF A FISH, GYMNARCHUS-NILOTICUS CUV". Nature. 167 (4240): 201–202. PMID 14806425. doi:10.1038/167201a0.
- Lissmann, H. W. (1958). "ON THE FUNCTION AND EVOLUTION OF ELECTRIC ORGANS IN FISH". Journal of Experimental Biology. 35: 156–&.
- Zakon, H. H., D. J. Zwickl, Y. Lu, and D. M. Hillis (2008). "Molecular evolution of communication signals in electric fish". Journal of Experimental Biology. 211 (11): 1814–1818. PMID 18490397. doi:10.1242/jeb.015982.
- Caputi, A. A., B. A. Carlson, and O. Macadar. 2005. Electric organs and their control. Pages 410-451 in T. H. Bullock, C. D. Hopkins, A. N. Popper, and R. R. Fay, eds. Electroreception. New York.
- Crampton W. G. R., Davis J. K., Lovejoy N. R., Pensky M. (2008). "Multivariate classification of animal communication signals: A simulation-based comparison of alternative signal processing procedures using electric fishes". Journal of Physiology-Paris. 102: 304–321. doi:10.1016/j.jphysparis.2008.10.001.
- Bastian J. 1986. Electrolocation: behavior, anatomy, and physiology. Pages 577-612 in T. H. Bullock and W. Heiligenberg, eds. Electroreception. New York.
- Aguilera P. A., Caputi A. A. (2003). "Electroreception in G. carapo: detection of changes in waveform of the electrosensory signals". Journal of Experimental Biology. 206: 989–998. doi:10.1242/jeb.00198.
- Pereira, A. C., and A. A. Caputi. 2010. Imaging in Electrosensory Systems. Interdisciplinary Sciences-Computational Life Sciences 2:291-307.
- Lavoue, S., R. Bigorne, G. Lecointre, and J. F. Agnese (2000). "Phylogenetic relationships of mormyrid electric fishes (Mormyridae; Teleostei) inferred from cytochrome b sequences". Molecular Phylogenetics and Evolution. 14 (1): 1–10. PMID 10631038. doi:10.1006/mpev.1999.0687.
- Lavoué S., Miya M., Arnegard M. E., Sullivan J. P., Hopkins C. D., Nishida M. (2012). "Comparable ages for the independent origins of electrogenesis in African and South American weakly electric fishes". PLoS ONE. 7: e36287. doi:10.1371/journal.pone.0036287.
- Kawasaki, M. (2009). "Evolution of Time-Coding Systems in Weakly Electric Fishes". Zoological Science. 26 (9): 587–599. PMID 19799509. doi:10.2108/zsj.26.587.
- Gallant, J. R., L. L. Traeger, J. D. Volkening, H. Moffett, P. H. Chen, C. D. Novina, G. N. Phillips, R. Anand, G. B. Wells, M. Pinch, R. Guth, G. A. Unguez, J. S. Albert, H. H. Zakon, M. P. Samanta, and M. R. Sussman (2008). "Genomic basis for the convergent evolution of electric organs". Science. 344: 1522–1525. PMID 24970089. doi:10.1126/science.1254432.