Response to frequency shifted artificial echoes in the batRhinolophus ferrumequinum
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In 5 roosting bats the resting frequency, that is the mean frequency of the cf-portion of consecutive sounds, is kept constant with a standard deviation which varies between 30–120 Hz in different bats and at different days. In 15 bats the emitted sounds were electronically shifted in frequency and played back as artificial echoes. Upward frequency shifts were responded by a decrease of the emission frequency. This frequency compensation occurred at frequency shifts of up to 4400 Hz in all bats and up to 6000 Hz in a few bats. The frequency decrease in different bats over the whole compensation range was 50–300 Hz smaller than the frequency shifts in the echoes. The echoes, therefore, returned at a frequency, called the reference frequency, which was by this compensation offset higher than the resting frequency. The standard deviations of the emission frequency in compensating bats were only slightly larger than in roosting bats and the same in the whole compensation range. All bats started to compensate frequency shifts when they were slightly larger than the compensation offset. Downward frequency shifts were not responded by a change of the emission frequency, but the accuracy with which the emission frequency was kept decreased somewhat. From these results it is concluded that the Doppler shift compensation system of the Horseshoe bats compares the echo frequency with the reference frequency and compensates deviations of upward frequency shifts.