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4 sounds, respectively. The initial frequencies of these FM sounds were approximately 30, 60, 90, and 120kHz, respectively. The duration and rise-decay time of the FM sounds were 3.0 and 0.5 ms, respectively. Acoustic stimuli were delivered from a condenser loudspeaker placed 72.4 cm in front of the bat's ears in an echo-attenuated soundproof chamber. The parameters of individual FM sounds and the time interval between paired sounds (hereafter called the echo delay) were manually varied to initially determine the best FM frequencies and best amplitudes of the FM sounds and the best delay (BD) to excite a neuron (15, 16). Then, the paired FM stimuli were fixed at the best FM frequencies and best amplitudes of the neuron, and echo delay was varied with a computer program.
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For electrical stimulation, a tungsten wire electrode (negative pole) was placed at a depth of 600 to 700 μm from the cortical surface and another (positive pole) was placed on the cortical surface just above the negative pole. The thickness of the FM-FM area was ∼900 μm. The BD of multiple neurons was again measured. Then, an electrical stimulus (100 nA, 0.2 ms) was delivered through these electrodes synchronously with each pulse stimulus in a delay scan; that is, the electrical stimulus was repeated every 150 ms, 11 times in each delay scan
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Two injections of 41.6 nl of 1.0% lidocaine solution were made at a depth of 600 to 700 μm with a mechanical microinjection unit (Picospritzer II, General Value Corp., Fairfield, NJ). Each injection lasted 700 ms and was separated by 1 s.
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The change in auditory response during the 6.7-min period of electrical stimulation was small and was often obscured by the stimulus artifact. Therefore, all the data described in this paper were obtained after the termination of the electrical stimulus.
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A 50% delay width is the width of a delay tuning curve at 50% of the maximum response (27).
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This positive feedback mediated by cortical auditory neurons is similar to that mediated by cortical visual neurons. However, synchronized oscillatory discharges mediated by the corticofugal fibers appeared to be very rare and weak in the auditory system, because cortical FM-FM neurons show almost no oscillatory discharges [J. Horikawa et al., Hear. Res. 76, 45 (1994)].
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We thank the Natural Resource Conservation Authority and the Ministry of Agriculture of Jamaica for permitting us to collect and export the mustached bats used in our research. We also thank S. Kuwada, J. F. Olsen, W. E. O'Neill, and A. Kadir for their comments on the manuscript. This work was supported by research grants from the National Institute on Deafness and Other Communication Disorders (DC 00175) and the Office of Naval Research (N00014-90-J-1068). The protocol of our research was approved by the Animal Studies Committee of Washington University (approval number 92279).
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