Effects of echo intensity on Doppler-shift compensation behavior in horseshoe bats

Abstract

Echolocating horseshoe bats respond to flight-speed induced shifts in echo frequency by adjusting the frequency of subsequent calls. Under natural conditions, Doppler effects may force the frequency of a returning echo several kilohertz above the original emission frequency. By lowering subsequent call frequencies, the bat can return echo frequencies to within a narrow spectral bandwidth to which its highly specialized auditory system is most sensitive. While Doppler-shift compensation (DSC) behavior specifically refers to frequency compensation, other parameters of the returning echo, such as delay, duration, and interaural time and intensity differences have been shown to influence DSC performance. Understanding the nature of these influences has already led to a better appreciation of the neurophysiology of DSC. Here we provide a quantitative analysis of the effects of a prominent feature of the returning echo, its intensity, on DSC performance in horseshoe bats. Although DSC performance generally tolerates echo attenuation up to approximately 40 dB relative to the outgoing emission intensity, a systematic decline in DSC performance can be observed over this range. Generally, the effects of echo attenuation are characterized by a reduction in 1) the overall amount of compensation relative to the size of the shift in echo frequency and 2) the rate at which the bat responds to perceived echo shifts. These effects appear to be the consequence of a systematic shift in the range of echo frequencies capable of inducing DSC behavior. In particular, the reference frequency (the minimum shift in echo frequency that will elicit DSC behavior) appears to be highly sensitive to echo intensity. Every 10-dB reduction in echo intensity shifts the reference upward nearly 250 Hz. Our results indicate that, even at the highest intensity levels, relatively minor changes in echo intensity critically influence frequency compensation during normal DSC. We conclude with a discussion of how these results might impact echolocation behavior of horseshoe bats under natural and experimental conditions.