6. SOUND PRODUCTION AND DETECTION
171
Kleerekoper and Roggenkamp, 1959). Upper frequency limits for these
fishes were given as over 7000 Hz (Farkas, 1936; Kleerekoper and
Chagnon, 1954; von Boutteville, 1935; von Frisch, 1938b).
The first attempt to determine a complete audiogram for a fish was
done with the bullhead, Zctalurus nebulosus, by Poggendorf ( 1952) (Fig.
21). In recent years, audiograms for two additional ostariophysine species
have been reported: the goldfish, Carassius uurutus, by Enger (1966,
1967b), Jacobs and Tavolga ( 1967), and Weiss ( 1967, 1969); and the
Mexican blind characin, Astyanux mexicanus, by Popper. ( 1970).
A comparison of the three audiograms obtained for the goldfish shows
some interesting results of different techniques. Enger (1967b) used a
positive reward conditioning ( Fig. 22) , while Jacobs and Tavolga ( 1967)
(Fig. 21) and Weiss (1967) (Fig. 23) used different forms of avoidance
conditioning. Enger used a loudspeaker in the water and also in air;
Jacobs and Tavolga used a loudspeaker in air, but in an enclosed chamber such that the backwave of the speaker was damped out; Weiss used
a tank with two large speaker surfaces at the ends, with the speakers
operating 180" out of phase. The quality of the stimulus in each of these
experiments was somewhat different. The Enger underwater-loudspeaker
+40 r
t
8 + I 0
5 0
6 -301
-40 t
i
I
0
Holocentrus
i
Haemulon f
1' fCarassius
- 50
I
1
I
I
1
I I I I I I
1
I I I L I
50
100
m
500
lo00 2o00
5000
Frequency (in Hz)
Fig. 21. Comparison of audiograms for four species of teleost fishes, Goldfish,
Carassius auratus, from Jacobs and Tavolga ( 1967); brown bullhead, lctalurus
nebulosus, from Poggendorf ( 1952 ) ; blue-striped grunt, Haemulon sciurus, from
Tavolga and Wodinsky ( 1965) ; and squirrelfish, Holocentrus ascensionis, from
Tavolga and Wodinsky (1963).