Sonar gain control and echo detection thresholds in the echolocating bat, Eptesicus fuscus

The echolocating bat, Eptesicus fuscus, detects sonar echoes with a sensitivity that changes according to the time elapsed between broadcasting of each sonar signal and reception of echoes. When tested in an electronic target simulator on a two-choice echo-detection task, the bat's threshold improved by 11.5 dB as echo delay changed from 2.3 to 4.6 ms (target ranges of 40 and 80 cm). Earlier experiments measured the change in detection threshold for delays from 1 to 6.4 ms (target ranges from about 17 to 110 cm) and obtained about 11 dB of improvement per doubling of delay. The new experiments used electronic delay lines to simulate echo delay, thus avoiding movement of loudspeakers to different distances and the possible creation of delay-dependent backward masking between stimulus echoes and cluttering echoes from the loudspeaker surfaces. The slope of the threshold shift defines an echo gain control that keeps echoes from point targets at a fixed sensation level--reducing sensitivity by 11 to 12 dB as echo amplitude increases by 12 dB per halving of range during the bat's approach to the target. A recent experiment using loudness discrimination of echoes at 70 to 80 dB SPL (roughly 50 dB above threshold) found a slope of about 6 dB per halving of range, so the gain-control effect may be level dependent. The observed effect is operationally equivalent to forward masking of echoes by the transmission, but any events correlated with vocalization which impair hearing sensitivity for a short interval following transmissions could cause a decline in sensitivity to echoes. Contractions of the bat's middle-ear muscles synchronized to transmissions may account for the observed threshold shift, at least for a span of echo delays associated with the most critical portion of the approach stage of pursuit. Forward masking by the sonar transmissions may contribute to the threshold shift, too, but middle-ear muscle contractions do occur and must be a significant part of the cause.