Automatic gain control in the bat's sonar receiver and the neuroethology of echolocation

Abstract

The sensitivity of the echolocating bat, Eptesicus fuscus, to sonar echoes at different time delays after sonar emissions was measured in a two-choice echo detection experiment. Since echo delay is perceptually equivalent to target range, the experiment effectively measured sensitivity to targets at different ranges. The bat's threshold for detecting sonar echoes at a short delay of only 1.0 msec after emissions (corresponding to a range of 17 cm) was 36 dB SPL (peak to peak), but the threshold decreased to 8 dB SPL at a longer delay of 6.4 msec (a range of 1.1 m). Prior research has shown that, at even longer delays (corresponding to ranges of 3 to 5 m), the bat's threshold is in the region of 0 dB SPL. Contractions of the bat's middle ear muscles synchronized with the production of echolocation sounds cause a transient loss in hearing sensitivity which appears to account for the observed echo detection threshold shifts. The bat's echo detection thresholds increase by approximately 11 dB for each reduction in target range by a factor of 2 over the span from 17 cm to 1.1 m. As range shortens, the amplitude of echoes from small targets also increases, by 12 dB for each 2-fold reduction in range. Thus, when approaching a target, the bat compensates for changes in echo strength as target range shortens by changing its hearing threshold. Since this compensation appears to occur in the middle ear, the bat regulates echoes reaching the cochlea to a stable amplitude during its approach to a target such as a flying insect. In addition to this automatic gain control linked to target range, the bat aims its head to track a target's position during approach, thus stabilizing echo amplitude even if the target's direction changes. We hypothesize that the bat's directional emissions, directional hearing, middle ear muscle contractions, and head aim response collectively create a three-dimensional spatial tracking filter which the bat locks onto targets to stabilize echo amplitudes during interception of prey. We further hypothesize that this regulation, which cancels echo amplitude changes caused by the target's changing spatial position, leaves the bat free to observe echo amplitude changes caused by the target's own actions, such as insect wing beats. Elimination of spatially dependent echo amplitude changes removes the cause of potentially troublesome changes in neural response latency and keeps stimulation from echoes in the "tip" region of auditory nerve fiber tuning curves.(ABSTRACT TRUNCATED AT 400 WORDS)

Tempo and mode of antibat ultrasound production and sonar jamming in the diverse hawkmoth radiation

“…We examined sonar emissions and found that bats, in both experiments, attacked all silent control and sound-producing moths with comparable echolocation patterns (Fig. 2C), similarly progressing through the approach, track, and terminal phases of attack (34).…”

Steering by Hearing: A Bat’s Acoustic Gaze Is Linked to Its Flight Motor Output by a Delayed, Adaptive Linear Law

“…Griffin first reported the dramatic increase in PPR by echolocating bats during the final attack of insect prey and termed it the buzz (Griffin, 1958;Griffin et al, 1960). Subsequent studies in echolocating bats have used changes in PPR to infer changes in the bat's behavioral state (Kick and Simmons, 1984;Schnitzler et al, 1987;Kalko, 1995). Foraging flight has been divided into four stages, according to the bat's vocal behavior: searching, approaching, tracking, and attacking (Kick and Simmons, 1984).…”

“…Subsequent studies in echolocating bats have used changes in PPR to infer changes in the bat's behavioral state (Kick and Simmons, 1984;Schnitzler et al, 1987;Kalko, 1995). Foraging flight has been divided into four stages, according to the bat's vocal behavior: searching, approaching, tracking, and attacking (Kick and Simmons, 1984). The tracking and attack phases correspond to the terminal I and terminal II stages described by some authors (Kalko, 1995).…”

“…After the bat detects the insect, the bat moves into the approach stage and the pulse rate rises (20 -50 Hz). It then transitions to the tracking stage (50 Hz) (Kick and Simmons, 1984) and finally to the attacking stage (up to 200 Hz). These PPR values are estimates from field studies and vary with species of bat (for review, see Schnitzler and Kalko, 2001).…”

Sensory feedback control of mammalian vocalizations

“…Call intensity has a profound impact on ranging capabilities of the echolocation system, but very loud calls also carry substantial energy requirements, thus bats must efficiently regulate call intensity to match behapvioral contexts. It is well known that many bats decrease the intensity of their echolocation calls as they approach obstacles or prey [42,59,65,151]. This echo intensity compensation behavior is believed to optimize echo intensities for signal analysis within the auditory system [21,46,69], particularly during critical phases such as prey capture and navigating through cluttered spaces.…”