This is the first paper of a series dealing with sound-power collection by the auditory periphery of the gerbil. The purpose of the series is to quantify the physiological action of the gerbil's relatively large tympanic membrane and middle-ear air cavities. To this end the middle-ear input impedance ZT was measured at frequencies between 10 Hz and 18 kHz before and after manipulations of the middle-ear cavity. The frequency dependence of ZT is consistent with that of the middle-ear transfer function computed from extant data. Comparison of the impedance and transfer function suggests a middle-ear transformer ratio of 50 at frequencies below 1 kHz, substantially smaller than the anatomical value of 90 [Lay, J. Morph. 138, 41-120 (1972)]. Below 1 kHz the data suggest a low-frequency acoustic stiffness KT for the middle ear of 970 Pa/mm3 and a stiffness of the middle-ear cavity of 720 Pa/mm3 (middle-ear volume V MEC of 195 mm3); thus the middle-ear air spaces contribute about 70% of the acoustic stiffness of the auditory periphery. Manipulations of a middle-ear model suggest that decreases in V MEC lead to proportionate increases in KT but that further increases in middle-ear cavity volume produce only limited decreases in middle-ear stiffness. The data and the model point out that the real part of the middle-ear impedance at frequencies below 100 Hz is determined primarily by losses within the middle-ear cavity. The measured impedance is comparable in magnitude and frequency dependence to the impedance in several larger mammalian species commonly used in auditory research. A comparison of low-frequency stiffness and anatomical dimensions among several species suggests that the large middle-ear cavities in gerbil act to reduce the middle-ear stiffness at low frequencies. A description of sound-power collection by the gerbil ear requires a description of the function of the external ear.
1. The electrophysiological responses of single units in the dorsal cochlear nucleus of unanesthetized decerebrate Mongolian gerbil (Meriones unguiculatus) were recorded. Units were classified according to the response map scheme of Evans and Nelson as modified by Young and Brownell, Young and Voigt, and Shofner and Young. Type II units have a V-shaped excitatory response map similar to typical auditory nerve tuning curves but little or no spontaneous activity (SpAc < 2.5 spikes/s) and little or no response to noise. Type I/III units also have a V-shaped excitatory map and SpAc < 2.5 spikes/s, but have an excitatory response to noise. Type III units have a V-shaped excitatory map with inhibitory sidebands, SpAc > 2.5 spikes/s, and an excitatory response to noise. Type IV-T units typically also have a V-shaped excitatory map with inhibitory sidebands, but have a highly nonmonotonic rate versus level response to best frequency (BF) tones like type IV units, SpAc > 2.5 spikes/s, and an excitatory response to noise. Type IV units have a predominantly inhibitory response map above an island of excitation of BF, SpAc > 2.5 spikes/s, and an excitatory response to noise. We present results for 133 units recorded with glass micropipette electrodes. The purpose of this study was to establish a normative response map data base in this species for ongoing structure/function and correlation studies. 2. The major types of units (type II, type I/III, type III, type IV-T, and type IV) found in decerebrate cat are found in decerebrate gerbil. However, the percentage of type II (7.5%) and type IV (11.3%) units encountered are smaller and the percentage of type III (62.4%) units is larger in decerebrate gerbil than in decerebrate cat. In comparison, Shofner and Young found 18.5% type II units, 30.6% type IV units, and 23.1% type III units using metal electrodes. 3. Two new unit subtypes are described in gerbil: type III-i and type IV-i units. Type III-i units are similar to type III units except that type III-i units are inhibited by low levels of noise and excited by high levels of noise whereas type III units have strictly excitatory responses to noise. Type IV-i units are similar to type IV units except that type IV-i units are excited by low levels of noise and become inhibited by high levels of noise whereas type IV units have strictly excitatory responses to noise. Type III-i units are approximately 30% of the type III population and type IV-i units are approximately 50% of the type IV population. 4. On the basis of the paucity of classic type II units and the reciprocal responses to broadband noise of type III-i and type IV-i units, we postulate that some gerbil type III-i units are the same cell type and have similar synaptic connections as cat type II units. 5. Type II and type I/III units are distinguished from one another on the basis of both their relative noise response, rho, and the normalized slope of the BF tone rate versus level functions beyond the first maximum. Previously, type II units were defined to be those ...
Responses of auditory nerve fibers to steady-state vowels presented alone and in the presence of background noise were obtained from anesthetized cats. Representation of vowels based on average discharge rate and representation based primarily on phase-locked properties of responses are considered. Profiles of average discharge rate versus characteristic frequency (CF) ("rate-place" representation) can show peaks of discharge rate in the vicinity of formant frequencies when vowels are presented alone. These profiles change drastically in the presence of background noise, however. At moderate vowel and noise levels and signal/noise ratios of +9 dB, there are not peaks of rate near the second and third formant frequencies. In fact, because of two-tone suppression, rate to vowels plus noise is less than rate to noise alone for fibers with CFs above the first formant. Rate profiles measured over 5-ms intervals near stimulus onset show clear formant-related peaks at higher sound levels than do profiles measured over intervals later in the stimulus (i.e., in the steady state). However, in background noise, rate profiles at onset are similar to those in the steady state. Specifically, for fibers with CFs above the first formant, response rates to the noise are suppressed by the addition of the vowel at both vowel onset and steady state. When rate profiles are plotted for low spontaneous rate fibers, formant-related peaks appear at stimulus levels higher than those at which peaks disappear for high spontaneous fibers. In the presence of background noise, however, the low spontaneous fibers do not preserve formant peaks better than do the high spontaneous fibers. In fact, the suppression of noise-evoked rate mentioned above is greater for the low spontaneous fibers than for high. Representations that reflect phase-locked properties as well as discharge rate ("temporal-place" representations) are much less affected by background noise. We have used synchronized discharge rate averaged over fibers with CFs near (+/- 0.25 octave) a stimulus component as a measure of the population temporal response to that component. Plots of this average localized synchronized rate (ALSR) versus frequency show clear first and second formant peaks at all vowel and noise levels used. Except at the highest level (vowel at 85 dB sound pressure level (SPL), signal/noise = +9 dB), there is also a clear third formant peak. At signal-to-noise ratios where there are no second formant peaks in rate profiles, human observers are able to discriminate second formant shifts of less than 112 Hz. ALSR plots show clear second formant peaks at these signal/noise ratios.
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