On the Function and Evolution of Electric Organs in Fish

Abstract

1. The electric discharges of Gymnarchus niloticus and of representative species of seven genera of the Mormyridae have been examined in their natural habitat in Africa and in the laboratory. 2. Comparable investigations of the South American Gymnotidae have shown the existence of two discharge types in both these unrelated fish families. 3. The first type of electric discharge consists of very regular sequences of continuously emitted, monophasic pulses, varying from species to species in frequency, and within narrower limits from individual to individual. 4. Fish emitting this first type of pulses include Gymnarchus, Hypopomus and Eigenmannia. The frequency range for these fish lies between 60 and 400 discharges/sec. 5. The frequency does not alter with the state of excitation of the fish. The duration of individual pulses is relatively long, i.e. 2-10 msec. 6. The second type of discharge is less regular in frequency, the pulse duration much shorter and the pulse shape more complex. The individual discharge from the whole electric organ lasts about 0.2 msec, in Petrocephalus. 7. This type of discharge is found in all the examined species of the Mormyridae and in such forms as Gymnotus carapo and Staetogenes elegans. 8. The basic discharge rate of a resting mormyrid is somewhat variable and not strictly rhythmical. It usually lies between 1 and 6 pulses/sec. 9. Stimuli which excite the mormyrids cause an increase in the discharge frequency. The recorded maximum is about 130 pulses/sec. 10. Suitable stimuli can inhibit the discharges of the Mormyridae for prolonged periods. 11. In Gymnotus carapo and Staetogenes elegans the basic discharge rate is higher and of regular rhythmicity. Depending on temperature the frequencies lie between 30 and 87 pulses/sec. When these fish are excited the frequencies are increased up to 200 pulses/sec, for a short time. 12. The shape of the electric field, which is set up with each pulse around the fish, has been examined. 13. A theory has been proposed which suggests that these fish, by means of their electric pulses, can locate objects if their electrical conductivity differs from that of water. 14. These fish have shown themselves extremely sensitive to influences affecting the electric field. This has been studied by applying artificial electric stimuli, by studying the effects of conductors and non-conductors introduced into the field, and the reactions towards magnetic fields and electrostatic charges. 15. Conditioned reflex experiments with Gymnarchus niloticus and Gymnotus carapo have shown that these fish can detect the presence of a stationary magnet, and that they can discriminate between conductors and non-conductors. 16. The prey of these fish does not appear to be affected by the discharges. Inter alia, the electric pulses have a social significance. 17. This locating mechanism may be considered as an adaptation to life in turbid water. 18. Gymnotidae and Mormyridae (taken to include Gymnarchus) show striking features of convergent evolution. 19. Unusual locomotory adaptations such as swimming by means of the dorsal fin (Gymnarchus), the anal fin (Gymnotidae) and ‘Gemminger's bones’ (Mormyridae) may be considered as a means which tends to make the axis of symmetry of the fish and of its electric field coincide during active movements. 20. A new theory for the evolution of electric organs has been suggested. A major prerequisite appears to be a receptor sensitive to electrical stimulation. 21. It is suggested that special sensory and nervous differentiations of the lateralis system (‘mormyromasts’, valvulae cerebelli) are concerned with the perception and integration of electric stimuli. 22. Muscular action potentials have been recorded in the water at some distance from non-electric fish. 23. The easiest explanation for the evolution of strong electric organs would appear to start from such muscular action potentials, and proceed via weak electric organs used for orientation, to the powerful offensive and defensive electric organs.