Directional sensitivity of the auditory midbrain in the mustached bat to free-field tones

1986 ◽  
Vol 24 (1) ◽  
pp. 73-88 ◽  
Author(s):  
James C. Makous ◽  
William E. O'Neill
1983 ◽  
Vol 50 (5) ◽  
pp. 1182-1196 ◽  
Author(s):  
A. Asanuma ◽  
D. Wong ◽  
N. Suga

The orientation sound emitted by the Panamanian mustached bat, Pteronotus parnellii rubiginosus, consists of four harmonics. The third harmonic is 6-12 dB weaker than the predominant second harmonic and consists of a long constant-frequency component (CF3) at about 92 kHz and a short frequency-modulated component (FM3) sweeping from about 92 to 74 kHz. Our primary aim is to examine how CF3 and FM3 are represented in a region of the primary auditory cortex anterior to the Doppler-shifted constant-frequency (DSCF) area. Extracellular recordings of neuronal responses from the unanesthetized animal were obtained during free-field stimulation of the ears with pure tones. FM sounds, and signals simulating their orientation sounds and echoes. Response properties of neurons and tonotopic and amplitopic representations were examined in the primary and the anteroventral nonprimary auditory cortex. In the anterior primary auditory cortex, neurons responded strongly to single pure tones but showed no facilitative responses to paired stimuli. Neurons with best frequencies from 110 to 90 kHz were tonotopically organized rostrocaudally, with higher frequencies located more rostrally. Neurons tuned to 92-94 kHz were overpresented, whereas neurons tuned to sound between 64 and 91 kHz were rarely found. Consequently a striking discontinuity in frequency representation from 91 to 64 kHz was found across the anterior DSCF border. Most neurons exhibited monotonic impulse-count functions and responded maximally to sound pressure level (SPL). There were also neurons that responded best to weak sounds but unlike the DSCF area, amplitopic representation was not found. Thus, the DSCF area is quite unique not only in its extensive representation of frequencies in the second harmonic CF component but also in its amplitopic representation. The anteroventral nonprimary auditory cortex consisted of neurons broadly tuned to pure tones between 88 and 99 kHz. Neither tonotopic nor amplitopic representation was observed. Caudal to this area and near the anteroventral border of the DSCF area, a small cluster of FM-FM neurons sensitive to particular echo delays was identified. The responses of these neurons fluctuated significantly during repetitive stimulation.


1980 ◽  
Vol 86 (1) ◽  
pp. 153-170
Author(s):  
R. B. COLES ◽  
D. B. LEWIS ◽  
K. G. HILL ◽  
M. E. HUTCHINGS ◽  
D. M. GOWER

The directional sensitivity of cochlear microphonics (CM) was studied inthe quail by rotating a free-field sound source (pure tones, 160-10 kHz)through 360° in the horizontal plane, under anechoic conditions. Sound diffraction by the head was monitored simultaneously by a microphone at the entrance to the ipsilateral (recorded) ear canal. Pressure-field fluctuations measured by the microphone were non-directional (≤ 4 dB) up to 4 kHz; the maximum head shadow was 8 dB at 6.3 kHz. In comparison, the CM sensitivity under went directional fluctuations ranging up to 25 dB for certain low, mid and high frequency band widths. There was noticeable variation between quail for frequencies producing maximum directional effects, although consistently poor directionality was seen near 820 Hz andto a lesser extent near 3.5 kHz. Well-defined CM directivity patterns reflected the presence of nulls (insensitive regions) at critical positions around the head and the number of nulls increased with frequency. Five major types of directivity patterns were defined using polar co-ordinates: cardioid, supercardioid, figure-of-eight, tripartite and multilobed. Such patterns were largely unrelated to head shadow effects. Blocking the ear canal contralateral to there corded ear was shown to effectively abolish CM directionality, largely by eliminating regions of insensitivity to sound. It is inferred that the quail ear functions as an asym metrical pressure gradient receiver, the pressure gradient function being mediated by the interauralcavity. It is proposed that the central auditory system codes directional information by a null detecting method and computes an unambiguous (i.e.intensity independent) directional cue. This spatial cue is achieved by the difference between the directional sensitivities of the two ears, defined as the Directional Index (DI). The spatial distribution of DI values (difference pattern) demonstrated ranges and peaks which closely reflected the extent and position of nulls determined from monaural directivity functions. Large directional cues (up to 25 dB) extended throughout most of the audible spectrum of the quail and the sharpness of difference patterns increased with frequency. Primary ‘best’ directions, estimated from peaks in difference patterns, tended to move towards the front of the head at higher frequencies; rearward secondary peaks also occurred. From the properties of directional cues it is suggested that the ability of birds to localize sound need not necessarily depend on frequency; however, spatial acuity may be both frequency and direction dependent, and include the possibility of front-torearerrors. The directional properties of bird vocalizations may need to bere assessed on the basis of the proposed mechanism for directional hearing.


1988 ◽  
Vol 60 (4) ◽  
pp. 1384-1404 ◽  
Author(s):  
J. J. Wenstrup ◽  
Z. M. Fuzessery ◽  
G. D. Pollak

1. The responses to closed-field and free-field sound stimuli were obtained from EI neurons in the 60-kHz isofrequency representation of the mustache bat's inferior colliculus. The goals of the study were 1) to examine how binaural interactions and the directional properties of the ear shape the spatial response of 60-kHz EI neurons and 2) to examine features of the spatial responses of these isofrequency neurons which may encode the location of a sound. 2. The free-field response of monaural neurons was used to describe directional effects of the head and external ears at 60 kHz. Monaural neurons were most sensitive to sounds originating 26 degrees into the contralateral field, with sharply reduced sensitivity in the ipsilateral field and the lateral extreme of the contralateral field. The interaural intensity difference (IID), computed by assuming that the ears had mirror-image directional sensitivities, was greatest for sounds between 26 and 52 degrees off the vertical midline, where IIDs typically ranged from 20 to 30 dB. These values agree well with previous measures of the directional sensitivity of the ear obtained from cochlear microphonic recordings. 3. The response of a monaural unit to closed-field stimulation was used to predict its response to free-field stimulation. This prediction agreed closely with the observed free-field response, indicating that the monaural unit's free-field response could be explained by the directional sensitivity of the contralateral ear and the unit's response as a function of contralateral sound intensity. 4. EI neurons differed in basic features of their binaural response properties, as described in a companion paper. To examine how these features of binaural inhibition shaped spatial responses, we obtained responses to free-field sounds among neurons whose binaural properties had been thoroughly characterized. We then compared the spatial response to a prediction based upon the directional properties of the ear and the sensitivity of the EI neurons to IIDs and intensity. The close agreement between the observed and predicted spatial responses in 11 of 12 EI units indicated that the spatial responses were determined largely by sensitivity to IIDs and intensity and by the directional properties of the ear. 5. EI neurons in our sample were most sensitive, and responded best, to sounds originating 26-39 degrees into the contralateral field. These spatial response features were independent of the binaural properties of EI neurons. Instead, they depended upon the directional properties of the ears at 60 kHz.(ABSTRACT TRUNCATED AT 400 WORDS)


1978 ◽  
Vol 41 (4) ◽  
pp. 870-884 ◽  
Author(s):  
E. I. Knudsen ◽  
M. Konishi

1. The influence of sound location and sound frequency on the responses of single units in the midbrain auditory area (MLD) of the owl (Tyto alba) were studied using a movable sound source under free-field conditions. With this technique, two functionally distinct regions in MLD have been identified: a tonotopic region and a space-mapped region. 2. MLD units were classified according to their receptive-field properties: 1) limited-field units responded only to sound from a small, discrete area of space; 2) complex-field units exhibited two to four different excitatory areas separated by areas of reduced response or inhibition: 3) space-preferring units responded best to a certain area of space, but their fields expanded considerably with increasing sound intensities; 4) Space-independent units responded similarly to a sound stimulus regardless of its location in space. 3. Limited-field units were located exclusively along the lateral and anterior borders of MLD. These units were tuned to sound frequencies at the high end of the owl's audible range (5-8.7 kHz). They usually responded only at the onset of a tonal stimulus; but most importantly, the units were systematically arranged in this region according to the azimuths and elevations of their receptive fields, thus creating a physiological map of auditory space. Because of this latter, dominant aspect of its functional organization, this region is named the space-mapped region of MLD. 4. The receptive fields of units in the larger, medial portion of MLD were of the space-independent, space-preferring, or complex-field types. These units tended to respond in a sustained fashion to tone and noise bursts, and these units were arranged in a strict frequency-dependent order. Based on this last property, this region is named the tonotopic region of MLD. 5. Because of the salient differences in the response properties of their constituent units, it is argued that the space-mapped region and the tonotopic region are involved in different aspects of sound analysis.


1984 ◽  
Vol 222 (1227) ◽  
pp. 231-242 ◽  

Field potentials have been recorded in the torus semicircularis of the toad, Bufo marinus, in response to brief tones presented in the free field. The amplitude of the potentials varied with the frequency of the stimulus and location of the electrode along the rostro-caudal axis of the torus. All frequencies in the auditory range evoked largest potentials when the stimulus was located in the contralateral auditory field. Potentials evoked by low to mid frequencies were largest when the stimulus was located near the line orthogonal to the long axis of the animal. For progressively higher frequencies, the optimal stimulus position was progressively more anterior in the contralateral field. In animals in which one eighth nerve had been sectioned, field potentials evoked by tones of low to mid frequency were less sensitive to changes in stimulus direction than in normal animals. However, the directional sensitivity of field potentials evoked by mid to high frequencies was similar in monaural and normal animals. These observations suggest that binaural neural integration is important in determining the directional sensitivity of field potentials in the torus evoked by low to mid frequencies but not for potentials evoked by mid to high frequencies.


Author(s):  
T. R. Letowski ◽  
G. L. Ricard ◽  
J. E. Greives

We measured listeners' directional sensitivity for narrow bands of noise and for speech recognition while they were wearing the XM45 gas mask and hood and compared it to normal free-field listening. Bekesy tracking was used to measure absolute hearing thresholds for 1/3 octave bands of noise centered at 0.125, 0.25, 0.5, 1, 2, 3, 4, 6, and 8 kHz. and masked intelligibility thresholds for speech were measured using the Modified Rhyme Test. Both the noise and speech signals were presented from loudspeakers located at azimuths of 0°, 45°, 90°, 135°, and 180°. The mask and hood significantly elevated noise-band thresholds for those bands centered at 1.0 kHz and above. Sensitivity to masked speech was increased by over 12 dB by lateral loudspeaker placement, and wearing a gas mask and hood reduced this gain by 80 percent. These results document a loss of auditory spatial awareness that has not been apparent from previous studies of speech intelligibility while wearing chemical protective equipment.


1990 ◽  
Vol 64 (1) ◽  
pp. 225-235 ◽  
Author(s):  
N. Suga ◽  
M. Kawasaki ◽  
R. F. Burkard

1. The mustached bat, Pteronotus parnellii parnellii, emits bisonar pulses each consisting of eight components: CF1-4 and FM1-4. In the auditory cortex of the bat there are arrays of FM-FM neurons that are tuned to particular delays of echo FMn (n = 2, 3, or 4) from pulse FM1. They are specialized for the processing of target-range information. The FM signal is suited for ranging and also for target localization. Therefore we studied the directional sensitivity of FM-FM neurons with pulse FM1 and echo FMn. One of the FM1-FMn pair was moved around the bat's head while the other was fixed in front of the bat. 2. FM-FM neurons are sharply tuned in echo delay and are broadly tuned in echo amplitude. That is, they are tuned to a target that has a particular cross-sectional area and that is located at a particular distance from the bat. Their best amplitudes for echoes range between 8 and 73 dB sound pressure level (SPL). The best amplitude is approximately 30 dB higher than minimum threshold in the majority of neurons. 3. The higher the best amplitude is relative to minimum threshold, the larger the receptive field is at the best amplitude. The receptive field of FM-FM neurons at 30 dB above minimum threshold is always so large that it covers the entire contralateral auditory field or the entire contralateral field and the medial half of the ipsilateral auditory field. The large size of the receptive field and the uniform distribution of response magnitudes within the receptive field indicate that FM-FM neurons are not suited for sound localization. Directional information is probably processed in parallel by a separate population of neurons other than FM-FM neurons. 4. The receptive field of FM-FM neurons at 10 dB above minimum threshold is much smaller than that at 30 dB above minimum threshold, but it is still large. The mean azimuthal and elevational widths for echo FMn are greater than 70 degrees in all directions. There is no sign that FM-FM neurons are more directional than peripheral neurons. Furthermore, there is neither an azimuthal nor an elevational axis within the FM-FM area. 5. Mean best azimuths of FM-FM neurons are different for each echo FM harmonic: lateral 35 degrees for FM2 and lateral 19 degrees for FM3 and FM4.(ABSTRACT TRUNCATED AT 400 WORDS)


2019 ◽  
Vol 62 (3) ◽  
pp. 745-757 ◽  
Author(s):  
Jessica M. Wess ◽  
Joshua G. W. Bernstein

PurposeFor listeners with single-sided deafness, a cochlear implant (CI) can improve speech understanding by giving the listener access to the ear with the better target-to-masker ratio (TMR; head shadow) or by providing interaural difference cues to facilitate the perceptual separation of concurrent talkers (squelch). CI simulations presented to listeners with normal hearing examined how these benefits could be affected by interaural differences in loudness growth in a speech-on-speech masking task.MethodExperiment 1 examined a target–masker spatial configuration where the vocoded ear had a poorer TMR than the nonvocoded ear. Experiment 2 examined the reverse configuration. Generic head-related transfer functions simulated free-field listening. Compression or expansion was applied independently to each vocoder channel (power-law exponents: 0.25, 0.5, 1, 1.5, or 2).ResultsCompression reduced the benefit provided by the vocoder ear in both experiments. There was some evidence that expansion increased squelch in Experiment 1 but reduced the benefit in Experiment 2 where the vocoder ear provided a combination of head-shadow and squelch benefits.ConclusionsThe effects of compression and expansion are interpreted in terms of envelope distortion and changes in the vocoded-ear TMR (for head shadow) or changes in perceived target–masker spatial separation (for squelch). The compression parameter is a candidate for clinical optimization to improve single-sided deafness CI outcomes.


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