Temporal Damping in Response to Broadband Noise. II. Auditory Nerve

2008 ◽  
Vol 99 (4) ◽  
pp. 1942-1952 ◽  
Author(s):  
Philip X. Joris ◽  
Dries H. Louage ◽  
Marcel van der Heijden
Keyword(s):  
Inferior Colliculus ◽  
Auditory Nerve ◽  
Low Frequency ◽  
Auditory Pathway ◽  
Broadband Noise ◽  
Linear Summation ◽  
Interaural Time Differences ◽  
Oscillatory Response ◽  
Original Report ◽  
Composite Curve

II. Auditory nerve. Low-frequency neurons in the inferior colliculus (IC) show a damped oscillatory response as a function of interaural time differences (ITDs) of broadband noise. It was previously shown that several features of such noise-delay functions are well predicted by the composite curve, generated by the linear summation of responses to tones with varying ITD. This indicates a surprising degree of linearity at the midbrain level of the auditory pathway. A similar comparison between responses to tones and to noise has not been made at a more peripheral, monaural level and it is therefore unclear to what extent this linearity reflects peripheral physiology. Here, we compare cat auditory nerve responses to broadband noise and to isolevel tones. We constructed shuffled autocorrelograms for responses to tones and summed across frequencies to obtain a monaural composite curve. We then compare this composite curve to the shuffled autocorrelogram of responses to broadband noise and find that the match between tonal and noise responses is poorer at the level of the auditory nerve than at the level of the IC. The apparent linearity of responses in the IC is thus even more surprising than was apparent from its original report because it results from mechanisms interposed between the auditory nerve and the IC.

Comparison of Bandwidths in the Inferior Colliculus and the Auditory Nerve. II: Measurement Using a Temporally Manipulated Stimulus

2008 ◽  
Vol 100 (4) ◽  
pp. 2312-2327 ◽  
Author(s):  
Myles Mc Laughlin ◽  
Joelle Nsimire Chabwine ◽  
Marcel van der Heijden ◽  
Philip X. Joris
Keyword(s):  
Correlation Function ◽  
Inferior Colliculus ◽  
Auditory Nerve ◽  
Low Frequency ◽  
Population Level ◽  
Broadband Noise ◽  
Center Frequency ◽  
Future Studies ◽  
Two Populations

To localize low-frequency sounds, humans rely on an interaural comparison of the temporally encoded sound waveform after peripheral filtering. This process can be compared with cross-correlation. For a broadband stimulus, after filtering, the correlation function has a damped oscillatory shape where the periodicity reflects the filter's center frequency and the damping reflects the bandwidth (BW). The physiological equivalent of the correlation function is the noise delay (ND) function, which is obtained from binaural cells by measuring response rate to broadband noise with varying interaural time delays (ITDs). For monaural neurons, delay functions are obtained by counting coincidences for varying delays across spike trains obtained to the same stimulus. Previously, we showed that BWs in monaural and binaural neurons were similar. However, earlier work showed that the damping of delay functions differs significantly between these two populations. Here, we address this paradox by looking at the role of sensitivity to changes in interaural correlation. We measured delay and correlation functions in the cat inferior colliculus (IC) and auditory nerve (AN). We find that, at a population level, AN and IC neurons with similar characteristic frequencies (CF) and BWs can have different responses to changes in correlation. Notably, binaural neurons often show compression, which is not found in the AN and which makes the shape of delay functions more invariant with CF at the level of the IC than at the AN. We conclude that binaural sensitivity is more dependent on correlation sensitivity than has hitherto been appreciated and that the mechanisms underlying correlation sensitivity should be addressed in future studies.

Effects of interaural time delays of noise stimuli on low-frequency cells in the cat's inferior colliculus. II. Responses to band-pass filtered noises

1987 ◽  
Vol 58 (3) ◽  
pp. 543-561 ◽  
Author(s):  
J. C. Chan ◽  
T. C. Yin ◽  
A. D. Musicant
Keyword(s):  
Inferior Colliculus ◽  
Time Delays ◽  
Discharge Rate ◽  
Low Frequency ◽  
Central Peak ◽  
Broadband Noise ◽  
Central Nucleus ◽  
Rate Curve ◽  
Median Frequency ◽  
Band Pass

1. We studied cells in the central nucleus of the inferior colliculus of the cat that were sensitive to interaural time delays (ITDs) in order to evaluate the influence of the stimulus spectrum of noise signals. Stimuli were sharply filtered low-, high-, and band-pass noise signals whose cutoff frequencies and bandwidths were systematically varied. The responses to ITDs of these noise signals were compared with responses obtained to ITDs of broadband noise and pure tones. 2. The discharge rate in response to band-pass noise as a function of ITD was usually a cyclic function with decreasing peak amplitudes at longer ITDs. The reciprocal of the mean interval between adjacent peaks indicated how rapidly the response rate varied with ITD and was termed the response frequency (RF). This RF was approximately equal to the median frequency of the stimulus spectrum filtered by the cell's sync-rate curve, which was the product of the synchronization to interaural phase and the discharge rate plotted against frequency. This suggests that the RF was determined by all the spectral components in the stimulus that fell within the frequency range in which the cell's response was synchronized. The contribution of each component was proportional to the sync-rate for that frequency. 3. The central peak of the ITD function usually fell within the physiological range of ITDs (+/- 400 microseconds). The location of this peak did not vary significantly with changes in stimulus spectrum by comparison with responses to tones of different frequency. Its shape also remained constant, except for a decrease in width when high-frequency components within the range of the sync-rate curve were added to the stimulus. A few cells responded with a minimal discharge instead of a maximal near-zero ITD, and this central minimum had similar properties as the central peak. The amplitude of the secondary peaks of the ITD function decreased as the stimulus bandwidth that overlapped the sync-rate curve broadened. 4. The sum of the ITD functions to two band-pass signals was similar to that of a broadband signal whose spectrum was composed of the sum of the band-pass spectra. 5. From these binaural responses we could make inferences about the response characteristics of the monaural inputs to binaural neurons. We then verified these predictions by studying responses of low-frequency trapezoid body fibers to band-pass noises.

Temporal coding of envelopes and their interaural delays in the inferior colliculus of the unanesthetized rabbit

1989 ◽  
Vol 61 (2) ◽  
pp. 257-268 ◽  
Author(s):  
R. Batra ◽  
S. Kuwada ◽  
T. R. Stanford
Keyword(s):  
Inferior Colliculus ◽  
Phase Locking ◽  
Acoustic Stimulus ◽  
Low Frequency ◽  
Auditory Pathway ◽  
Temporal Coding ◽  
Time Of Arrival ◽  
Unanesthetized Rabbit ◽  
The Difference ◽  
Ipsilateral Stimulation

1. The difference in the time of arrival of a sound at the two ears can be used to locate its source along the azimuth. Traditionally, it has been thought that only the on-going interaural temporal disparities (ITDs) produced by sounds of lower frequency (approximately less than 2 kHz) could be used for this purpose. However, ongoing ITDs of low frequency are also produced by envelopes of amplitude-modulated (AM) tones. These ITDs can be detected and used to lateralize complex high-frequency sounds (1, 8, 12, 15, 22, 24, 26). Auditory neurons synchronize to the modulation envelope, but do so at progressively lower modulation frequencies at higher levels of the auditory pathway. Some neurons of the cochlear nucleus synchronize best to frequencies as high as 700 Hz, but those of the inferior colliculus (IC) exhibit their best synchrony below 200 Hz. Even though synchrony to higher modulation frequencies is reduced at higher levels of the auditory pathway, is information about ITDs retained? 2. We answered this question by extracellularly recording the responses of neurons in the IC of the unanesthetized rabbit. We used an unanesthetized preparation because anesthesia alters the responses of neurons in the IC to both monaurally presented tones and ITDs. The unanesthetized rabbit is ideal for auditory research. Recordings can be maintained for long periods, and the acoustic stimulus to each ear can be independently controlled. 3. We studied the responses of 89 units to sinusoidally AM tones presented to the contralateral ear. For each unit, we recorded the response at several modulation frequencies. The degree of phase locking to the envelope at each frequency was measured using the synchronization coefficient. Two measures were used to assess the range of modulation frequencies over which phase locking occurred. The "best AM frequency" was the frequency at which we observed the greatest phase locking. The "highest AM frequency" was the highest frequency at which significant phase locking (0.001 level) was observed. We could not assess synchrony to ipsilateral AM tones directly, because most units did not respond to ipsilateral stimulation. 4. We studied the sensitivity of 63 units to ITDs produced by the envelopes of AM tones. Sensitivity to ITDs was tested by presenting AM tones to the two ears that had the same carrier frequency, but modulation frequencies that differed by 1 Hz. Units that were sensitive to ITDs responded to this stimulus by varying their response rate cyclically at the difference frequency, i.e., 1 Hz.(ABSTRACT TRUNCATED AT 400 WORDS)

Role of GABAergic Inhibition in the Coding of Interaural Time Differences of Low-Frequency Sounds in the Inferior Colliculus

2005 ◽  
Vol 93 (6) ◽  
pp. 3390-3400 ◽  
Author(s):  
W. R. D’Angelo ◽  
S. J. Sterbing ◽  
E.-M. Ostapoff ◽  
S. Kuwada
Keyword(s):  
Inferior Colliculus ◽  
Signal To Noise Ratio ◽  
Interaural Time Difference ◽  
Low Frequency ◽  
Gabaergic Inhibition ◽  
Interaural Time Differences ◽  
Gaba Antagonists ◽  
Inhibitory Transmitter ◽  
Almost All

A major cue for the localization of sound in space is the interaural time difference (ITD). We examined the role of inhibition in the shaping of ITD responses in the inferior colliculus (IC) by iontophoretically ejecting γ-aminobutyric acid (GABA) antagonists and GABA itself using a multibarrel pipette. The GABA antagonists block inhibition, whereas the applied GABA provides a constant level of inhibition. The effects on ITD responses were evaluated before, during and after the application of the drugs. If GABA-mediated inhibition is involved in shaping ITD tuning in IC neurons, then applying additional amounts of this inhibitory transmitter should alter ITD tuning. Indeed, for almost all neurons tested, applying GABA reduced the firing rate and consequently sharpened ITD tuning. Conversely, blocking GABA-mediated inhibition increased the activity of IC neurons, often reduced the signal-to-noise ratio and often broadened ITD tuning. Blocking GABA could also alter the shape of the ITD function and shift its peak suggesting that the role of inhibition is multifaceted. These effects indicate that GABAergic inhibition at the level of the IC is important for ITD coding.

scholarly journals Temporal Features of Spectral Integration in the Inferior Colliculus: Effects of Stimulus Duration and Rise Time

2009 ◽  
Vol 102 (1) ◽  
pp. 167-180 ◽  
Author(s):  
Donald Gans ◽  
Kianoush Sheykholeslami ◽  
Diana Coomes Peterson ◽  
Jeffrey Wenstrup
Keyword(s):  
Inferior Colliculus ◽  
Rise Time ◽  
Low Frequency ◽  
Auditory Pathway ◽  
Spectral Integration ◽  
Auditory Responses ◽  
Temporal Features ◽  
Vocal Signals ◽  
Inhibitory Interactions ◽  
Different Levels

This report examines temporal features of facilitation and suppression that underlie spectrally integrative responses to complex vocal signals. Auditory responses were recorded from 160 neurons in the inferior colliculus (IC) of awake mustached bats. Sixty-two neurons showed combination-sensitive facilitation: responses to best frequency (BF) signals were facilitated by well-timed signals at least an octave lower in frequency, in the range 16–31 kHz. Temporal features and strength of facilitation were generally unaffected by changes in duration of facilitating signals from 4 to 31 ms. Changes in stimulus rise time from 0.5 to 5.0 ms had little effect on facilitatory strength. These results suggest that low frequency facilitating inputs to high BF neurons have phasic-on temporal patterns and are responsive to stimulus rise times over the tested range. We also recorded from 98 neurons showing low-frequency (11–32 kHz) suppression of higher BF responses. Effects of changing duration were related to the frequency of suppressive signals. Signals <23 kHz usually evoked suppression sustained throughout signal duration. This and other features of such suppression are consistent with a cochlear origin that results in masking of responses to higher, near-BF signal frequencies. Signals in the 23- to 30-kHz range—frequencies in the first sonar harmonic—generally evoked phasic suppression of BF responses. This may result from neural inhibitory interactions within and below IC. In many neurons, we observed two or more forms of the spectral interactions described here. Thus IC neurons display temporally and spectrally complex responses to sound that result from multiple spectral interactions at different levels of the ascending auditory pathway.

Tuning to Interaural Time Difference and Frequency Differs Between the Auditory Arcopallium and the External Nucleus of the Inferior Colliculus

2009 ◽  
Vol 101 (5) ◽  
pp. 2348-2361 ◽  
Author(s):  
Katrin Vonderschen ◽  
Hermann Wagner
Keyword(s):  
Inferior Colliculus ◽  
Tuning Curve ◽  
Interaural Time Difference ◽  
Low Frequency ◽  
Steep Slope ◽  
Time Difference ◽  
Interaural Time Differences ◽  
Tuning Curves ◽  
Barn Owls ◽  
Zero Time

Barn owls process sound-localization information in two parallel pathways, the midbrain and the forebrain pathway. Exctracellular recordings of neural responses to auditory stimuli from far advanced stations of these pathways, the auditory arcopallium in the forebrain and the external nucleus of the inferior colliculus in the midbrain, demonstrated that the representations of interaural time difference and frequency in the forebrain pathway differ from those in the midbrain pathway. Specifically, low-frequency representation was conserved in the forebrain pathway, while it was lost in the midbrain pathway. Variation of interaural time difference yielded symmetrical tuning curves in the midbrain pathway. By contrast, the typical forebrain-tuning curve was asymmetric with a steep slope crossing zero time difference and a less-steep slope toward larger contralateral time disparities. Low sound frequencies contributed sensitivity to contralateral leading sounds underlying these asymmetries, whereas high frequencies enhanced the steepness of slopes at small interaural time differences. Furthermore, the peaks of time-disparity tuning curves were wider in the forebrain than in the midbrain. The distribution of the steepest slopes of best interaural time differences in the auditory arcopallium, but not in the external nucleus of the inferior colliculus, was centered at zero time difference. The distribution observed in the auditory arocpallium is reminiscent of the situation observed in small mammals. We speculate that the forebrain representation may serve as a population code supporting fine discrimination of central interaural time differences and coarse indication of laterality of a stimulus for large interaural time differences.

scholarly journals Neural Correlates and Mechanisms of Spatial Release From Masking: Single-Unit and Population Responses in the Inferior Colliculus

2005 ◽  
Vol 94 (2) ◽  
pp. 1180-1198 ◽  
Author(s):  
Courtney C. Lane ◽  
Bertrand Delgutte
Keyword(s):  
Inferior Colliculus ◽  
Single Unit ◽  
Signal To Noise Ratio ◽  
Low Frequency ◽  
Broadband Noise ◽  
Auditory Signal ◽  
Spatial Release ◽  
Additional Processing ◽  
Release From Masking ◽  
Correlator Model

Spatial release from masking (SRM), a factor in listening in noisy environments, is the improvement in auditory signal detection obtained when a signal is separated in space from a masker. To study the neural mechanisms of SRM, we recorded from single units in the inferior colliculus (IC) of barbiturate-anesthetized cats, focusing on low-frequency neurons sensitive to interaural time differences. The stimulus was a broadband chirp train with a 40-Hz repetition rate in continuous broadband noise, and the unit responses were measured for several signal and masker (virtual) locations. Masked thresholds (the lowest signal-to-noise ratio, SNR, for which the signal could be detected for 75% of the stimulus presentations) changed systematically with signal and masker location. Single-unit thresholds did not necessarily improve with signal and masker separation; instead, they tended to reflect the units' azimuth preference. Both how the signal was detected (through a rate increase or decrease) and how the noise masked the signal response (suppressive or excitatory masking) changed with signal and masker azimuth, consistent with a cross-correlator model of binaural processing. However, additional processing, perhaps related to the signal's amplitude modulation rate, appeared to influence the units' responses. The population masked thresholds (the most sensitive unit's threshold at each signal and masker location) did improve with signal and masker separation as a result of the variety of azimuth preferences in our unit sample. The population thresholds were similar to human behavioral thresholds in both SNR value and shape, indicating that these units may provide a neural substrate for low-frequency SRM.

Comparison of Bandwidths in the Inferior Colliculus and the Auditory Nerve. I. Measurement Using a Spectrally Manipulated Stimulus

2007 ◽  
Vol 98 (5) ◽  
pp. 2566-2579 ◽  
Author(s):  
Myles Mc Laughlin ◽  
Bram Van de Sande ◽  
Marcel van der Heijden ◽  
Philip X. Joris
Keyword(s):  
Inferior Colliculus ◽  
Auditory Nerve ◽  
Spike Timing ◽  
Temporal Coding ◽  
Broadband Noise ◽  
Auditory Periphery ◽  
Interaural Phase ◽  
Interaural Phase Difference ◽  
Frequency Components ◽  
Psychophysical Studies

A defining feature of auditory systems across animal divisions is the ability to sort different frequency components of a sound into separate neural frequency channels. Narrowband filtering in the auditory periphery is of obvious advantage for the representation of sound spectrum and manifests itself pervasively in human psychophysical studies as the critical band. Peripheral filtering also alters coding of the temporal waveform, so that temporal responses in the auditory periphery reflect both the stimulus waveform and peripheral filtering. Temporal coding is essential for the measurement of the time delay between waveforms at the two ears—a critical component of sound localization. A number of human psychophysical studies have shown a wider effective critical bandwidth with binaural stimuli than with monaural stimuli, although other studies found no difference. Here we directly compare binaural and monaural bandwidths (BWs) in the anesthetized cat. We measure monaural BW in the auditory nerve (AN) and binaural BW in the inferior colliculus (IC) using spectrally manipulated broadband noise and response metrics that reflect spike timing. The stimulus was a pair of noise tokens that were interaurally in phase for all frequencies below a certain flip frequency (fflip) and that had an interaural phase difference of π above fflip. The response was measured as a function of fflip and, using a separate stimulus protocol, as a function of interaural correlation. We find that both AN and IC filter BW depend on characteristic frequency, but that there is no difference in mean BW between the AN and IC.

Effects of interaural time delays of noise stimuli on low-frequency cells in the cat's inferior colliculus. III. Evidence for cross-correlation

1987 ◽  
Vol 58 (3) ◽  
pp. 562-583 ◽  
Author(s):  
T. C. Yin ◽  
J. C. Chan ◽  
L. H. Carney
Keyword(s):  
Inferior Colliculus ◽  
Acoustic Signal ◽  
Cross Correlation ◽  
Low Frequency ◽  
Nerve Fibers ◽  
Broadband Noise ◽  
The Other ◽  
Central Nucleus ◽  
Interaural Phase ◽  
Phase Relationships

1. We tested the coincidence, or cross-correlation, model of Jeffress, which proposes a neuronal mechanism for sensitivity to interaural time differences (ITDs) in low-frequency cells in the central nucleus of the inferior colliculus (ICC) of the cat. Different tokens of Gaussian noise stimuli were delivered to the two ears. We studied the neural responses to changes in ITDs of these stimuli and examined the manner in which the binaural cells responded to them. All of our results support the idea that the central binaural neurons perform an operation very similar to cross-correlation on the inputs arriving from each side. These inputs are transformed from the actual acoustic signal by the peripheral auditory system, and these transformations are reflected in the properties of the cross-correlations. 2. The responses to ITDs of identical broadband noise stimuli to the two ears varies cyclically as a function of ITD at a frequency close to the best frequency of the neuron. This cyclic response is a consequence of the narrowband filtering of the wideband acoustic signal by the auditory nerve fibers. To examine the effects of using stimuli to the two ears that were correlated to each other to different degrees, we generated pairs of noises. Each pair consisted of one standard noise, which was delivered to one ear, and a linear sum of two standard uncorrelated noises, which was delivered to the other ear. The responses of 34 neurons in the ICC to ITDs of noises with variable interaural coherence were examined. When partially correlated noises were delivered, there was a positive and approximately linear relationship between the degree of modulation of the response as a function of ITD and interaural coherence. The degree of modulation was measured by the synchronization coefficient, or vector strength, over one period of the ITD curve. 3. We examined the effects of altering the interaural phase relationships of the input noise stimuli. The phase of the noise stimuli was changed by digitally filtering the standard noise so that only a phase delay was imposed. The responses to ITDs with differing interaural phase relationships were then studied by delivering a phase-shifted noise to one ear and the standard noise to the other. The ITD curves in response to phase-shifted noise were shifted by about the same amount as the shift of the stimulus; the shift of the response was measured with respect to the case with identical noises to the two ears.(ABSTRACT TRUNCATED AT 400 WORDS)

scholarly journals Neural coding of time-varying interaural time differences and time-varying amplitude in the inferior colliculus

2017 ◽  
Vol 118 (1) ◽  
pp. 544-563 ◽  
Author(s):  
Nathaniel Zuk ◽  
Bertrand Delgutte
Keyword(s):  
Firing Rate ◽  
Inferior Colliculus ◽  
Amplitude Modulation ◽  
Temporal Coding ◽  
Broadband Noise ◽  
Neural Mechanisms ◽  
Time Varying ◽  
Neural Correlate ◽  
Interaural Time Differences ◽  
Binaural Cues

Binaural cues occurring in natural environments are frequently time varying, either from the motion of a sound source or through interactions between the cues produced by multiple sources. Yet, a broad understanding of how the auditory system processes dynamic binaural cues is still lacking. In the current study, we directly compared neural responses in the inferior colliculus (IC) of unanesthetized rabbits to broadband noise with time-varying interaural time differences (ITD) with responses to noise with sinusoidal amplitude modulation (SAM) over a wide range of modulation frequencies. On the basis of prior research, we hypothesized that the IC, one of the first stages to exhibit tuning of firing rate to modulation frequency, might use a common mechanism to encode time-varying information in general. Instead, we found weaker temporal coding for dynamic ITD compared with amplitude modulation and stronger effects of adaptation for amplitude modulation. The differences in temporal coding of dynamic ITD compared with SAM at the single-neuron level could be a neural correlate of “binaural sluggishness,” the inability to perceive fluctuations in time-varying binaural cues at high modulation frequencies, for which a physiological explanation has so far remained elusive. At ITD-variation frequencies of 64 Hz and above, where a temporal code was less effective, noise with a dynamic ITD could still be distinguished from noise with a constant ITD through differences in average firing rate in many neurons, suggesting a frequency-dependent tradeoff between rate and temporal coding of time-varying binaural information. NEW & NOTEWORTHY Humans use time-varying binaural cues to parse auditory scenes comprising multiple sound sources and reverberation. However, the neural mechanisms for doing so are poorly understood. Our results demonstrate a potential neural correlate for the reduced detectability of fluctuations in time-varying binaural information at high speeds, as occurs in reverberation. The results also suggest that the neural mechanisms for processing time-varying binaural and monaural cues are largely distinct.

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