Role of Broadcast Harmonics in Echo Delay Perception by Big Brown Bats

Big brown bats (Eptesicus fuscus) emit frequency-modulated (FM) echolocation sounds containing two principal down-sweeping harmonics (FM(1) approximately 55-25 kHz, FM(2) approximately 105-50 kHz). To determine whether each harmonic contributes to perception of echo delay, bats were trained to discriminate between "split-harmonic" echoes that differed in delay. The bat's broadcasts were picked up with microphones, and FM(1) and FM(2) were separated with highpass and lowpass filters at about 55 kHz, where they overlap in frequency. Both harmonics then were delivered from loudspeakers as positive stimuli in a 2-choice delay discrimination procedure with FM(1) delayed 3.16 ms and FM(2) delayed 3.46 ms (300 mus delay split). Negative stimuli contained FM(1) and FM(2) with the same filtering but no delay separation. These were presented at different overall delays from 11 down to 3 ms to measure the bat's delay discrimination acuity for each harmonic in the split harmonic echoes. The bats determined the delays of both FM(1) and FM(2), but performance was overlaid by a broad pedestal of poor performance that extended for 800 micros. Splitting the harmonics by 300 micros appears to defocus the bat's representation of delay, revealing the existence of a process for recognizing the normally simultaneous occurrence of the harmonics.