1. Previous experiments have demonstrated that (a) the shark Scyliorhinus canicula and the ray Raja clavata are extremely sensitive to weak electric fields; (b) their electrical sensitivity is due to the ampullae of Lorenzini; (c) the sharks and rays can be stimulated by the bioelectric fields emanating from the flatfish Pleuronectes platessa.
2. When hungry, Scyliorhinus and Raja perform well-aimed feeding responses to flatfish, even if the prey have covered themselves with sand. The object of the present study was to determine whether the sharks and rays use the bioelectric fields of the flatfish to detect the position of their prey.
3. To analyse the feeding responses of the sharks and rays, a flatfish was put into an agar chamber. The predators responded to the so screened prey from the same distance, and tried to feed on it in the same way as if there were no agar at all. As the flatfish in the agar chamber was completely hidden from view, the sharks and rays were thus shown not to need visual contact to locate the prey.
4. If the agar chamber was filled with cut-up pieces of whiting, the sharks and rays did not respond to the food, although the odour of whiting juice normally attracts them strongly. Therefore, the sharks and rays did not detect the position of the agarscreened flatfish by smell.
5. The feeding responses to the flatfish could be entirely abolished by covering the agar chamber with a very thin sheet of plastic. The mechanical attenuation offered by the plastic film was too weak to explain its dramatic inhibitory effect, and, thus, a purely mechanical detection of the agar-screened flatfish without plastic film was also ruled out.
6. As the responses to the agar-screened flatfish were not merely due to visual, chemical, or mechanical stimuli, it was tentatively concluded that the sharks and rays perceived the prey electrically. This conclusion was fully in agreement with the results of the experiments, for the agar chamber did not appreciably distort the bioelectric fields of the flatfish, and the electrical impedance of the plastic film was extremely high.
7. Further, the bioelectric field of a flatfish was simulated with a pair of electrodes, buried in the sand. Now, the sharks and rays displayed exactly the same feeding responses to the electrodes as they did previously to the real prey. This crucial experiment confirmed the electrical hypothesis in a very direct way.
8. The experiments described demonstrate clearly that the shark Scyliorhinus canicula and the ray Raja clavata make a biologically significant use of their electrical sensitivity. Therefore, we now are justified in accrediting the animals with an electric sense and in designating the ampullae of Lorenzini as electroreceptors.
9. When the sharks and rays were offered a piece of whiting in the vicinity of two electrodes simulating a flatfish, they were attracted by the odour of the food but usually performed their well-aimed responses to the electrodes. Thus, at short range, the electric fields act as a much stronger directive force than do the visual and chemical stimuli. Only direct mechanical contact dominates over the electrical stimuli.
10. Theoretically, the sharks and rays can detect the electric fields resulting from ceanic and tidal currents. Whether they make use of the available information for orientation in the open sea is not yet known. Furthermore, the observations and measurements described indicate that, in studying shark attacks, the electric fields of the prey and the electric sense of the predators should be taken into account.
Present address: Department of Neurosciences, School of Medicine, University of California, San DiegoLa Jolla, California 92037.