1. In Eptesicus the auditory cortex, as defined by electrical activity recorded from microelectrodes in response to tone bursts, FM sweeps, and combinations of FM sweeps, encompasses an average cortical surface area of 5.7 mm2. This area is large with respect to the total cortical surface area and reflects the importance of auditory processing to this species of bat. 2. The predominant pattern of organization in response to tone bursts observed in each cortex is tonotopic, with three discernible divisions revealed by our data. However, although cortical best-frequency (BF) maps from most of the individual bats are similar, no two maps are identical. The largest division contains an average of 84% of the auditory cortical surface area, with BF tonotopically mapped from high to low along the anteroposterior axis and is part of the primary auditory cortex. The medium division encompasses an average of 13% of the auditory cortical surface area, with highly variable BF organization across bats. The third region is the smallest, with an average of only 3% of auditory cortical surface area and is located at the anterolateral edge of the cortex. This region is marked by a reversal of the tonotopic axis and a restriction in the range of BFs as compared with the larger, tonotopically organized division. 3. A population of cortical neurons was found (n = 39) in which each neuron exhibited two BF threshold minima (BF1 and BF2) in response to tone bursts. These neurons thus have multipeaked frequency threshold tuning curves. In Eptesicus the majority of multipeaked frequency-tuned neurons (n = 27) have threshold minima at frequencies that correspond to a harmonic ratio of three-to-one. In contrast, the majority of multipeaked neurons in cats have threshold minima at frequencies in a ratio of three-to-two. A three-to-one harmonic ratio corresponds to the "spectral notches" produced by interference between overlapping echoes from multiple reflective surfaces in complex sonar targets. Behavioral experiments have demonstrated the ability of Eptesicus to use spectral interference notches for perceiving target shape, and this subpopulation of multipeaked frequency-tuned neurons may be involved in coding of spectral notches. 4. The auditory cortex contains delay-tuned neurons that encode target range (n = 99). Most delay-tuned neurons respond poorly to tones or individual FM sweeps and require combinations of FM sweeps. They are combination sensitive and delay tuned.(ABSTRACT TRUNCATED AT 400 WORDS)
To measure the directionality of the external ear of the echolocating bat, Eptesicus fuscus, the left or right eardrum of a dead bat was replaced by a microphone which recorded signals received from a sound source that was moved around the stationary head. The test signal was a 0.5-ms FM sweep from 100 kHz to 10 kHz (covering all frequencies in the bat's biosonar sounds). Notches and peaks in transfer functions for 7 tested ears varied systematically with changes in elevation. For the most prominent notch, center frequency decreased from about 50 kHz for elevations at or near the horizontal to 30-40 kHz for elevations 30 degrees-40 degrees below the horizontal. A second notch shifted from about 85 kHz to 70 kHz over these same elevations. Above the horizontal, a peak that flanks these notches changed in amplitude by 15 dB with changes in elevation. Removal of the tragus from the external ear disrupted the systematic movement of notch frequencies with elevation but did not disrupt changes in the peak's amplitude. Smaller changes in notch frequency also occurred with changes in azimuth, so monaural notch information alone cannot determine the position of sound sources away from the median plane. However, because bats routinely keep the head pointed at the target's azimuth, median-plane localization occurs with monaural cues delivered to the two ears. Corresponding changes with elevation occurred in the impulse-response, which consists of a series of 3-6 peaks spaced 10-20 microseconds apart. The time separation of two prominent impulse peaks systematically increased from 22-26 microseconds above the horizontal to about 36-40 microseconds below the horizontal, and removal of the tragus disrupted this time shift below the horizontal.
Neurons in the inferior colliculus (IC) of the awake big brown bat, Eptesicus fuscus, were examined for joint frequency and latency response properties which could register the timing of the bat's frequency-modulated (FM) biosonar echoes. Best frequencies (BFs) range from 10 kHz to 100 kHz with 50% tuning widths mostly from 1 kHz to 8 kHz. Neurons respond with one discharge per 2-ms tone burst or FM stimulus at a characteristic latency in the range of 3-45 ms, with latency variability (SD) of 50 microseconds to 4-6 ms or more. BF distribution is related to biosonar signal structure. As observed previously, on a linear frequency scale BFs appear biased to lower frequencies, with 20-40 kHz overrepresented. However, on a hyperbolic frequency (linear period) scale BFs appear more uniformly distributed, with little overrepresentation. The cumulative proportion of BFs in FM1 and FM2 bands reconstructs a scaled version of the spectrogram of FM broadcasts. Correcting FM latencies for absolute BF latencies and BF time-in-sweep reveals a subset of IC cells which respond dynamically to the timing of their BFs in FM sweeps. Behaviorally, Eptesicus perceives echo delay and phase with microsecond or even submicrosecond accuracy and resolution, but even with use of phase-locked FM and tone-burst stimuli the cell-by-cell precision of IC time-frequency registration seems inadequate by itself to account for the temporal acuity exhibited by the bat.
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