scholarly journals Neural Sensitivity to Periodicity in the Inferior Colliculus: Evidence for the Role of Cochlear Distortions

2004 ◽  
Vol 92 (3) ◽  
pp. 1295-1311 ◽  
Author(s):  
David McAlpine
Keyword(s):  
Inferior Colliculus ◽  
Basilar Membrane ◽  
Low Frequency ◽  
Neural Response ◽  
Auditory Midbrain ◽  
Discharge Rates ◽  
Interaural Phase ◽  
Neural Input ◽  
Interaural Phase Difference ◽  
Modulation Rate

Responses of low characteristic-frequency (CF) neurons in the inferior colliculus were obtained to amplitude-modulated (AM) high-frequency tones in which the modulation rate was equal to the neuron's CF. Despite all spectral components lying outside the pure tone–evoked response areas, discharge rates were modulated by the AM signals. Introducing a low-frequency tone (CF − 1 Hz) to the same ear as the AM tones produced a 1-Hz beat in the neural response. Introducing a tone (CF − 1 Hz) to the opposite ear to the AM tone also produced a beat in the neural response, with the beat at the period of the interaural phase difference between the CF − 1 Hz tone in one ear, and the AM rate in the other ear. The monaural and interaural interactions of the AM signals with introduced pure tones suggest that AM tones generate combination tones, (inter-modulation distortion) on the basilar membrane. These interact with low-frequency tones presented to the same ear to produce monaural beats on the basilar membrane, modulating the responses of inferior colliculus (IC) neurons on the 1-Hz period of the monaural beats or interacting binaurally with neural input generated in response to stimulation of the opposite ear. The auditory midbrain appears to show a robust representation of cochlear distortions generated by amplitude-modulated sounds.

Responses of Neurons in the Inferior Colliculus to Dynamic Interaural Phase Cues: Evidence for a Mechanism of Binaural Adaptation

2000 ◽  
Vol 83 (3) ◽  
pp. 1356-1365 ◽  
Author(s):  
David McAlpine ◽  
Dan Jiang ◽  
Trevor M. Shackleton ◽  
Alan R. Palmer
Keyword(s):  
Inferior Colliculus ◽  
The Other ◽  
Auditory Midbrain ◽  
Adaptation Mechanism ◽  
Motion Cues ◽  
Depth Direction ◽  
Discharge Rates ◽  
Interaural Phase ◽  
History Of ◽  
Different Depths

Responses to sound stimuli that humans perceive as moving were obtained for 89 neurons in the inferior colliculus (IC) of urethan-anesthetized guinea pigs. Triangular and sinusoidal interaural phase modulation (IPM), which produced dynamically varying interaural phase disparities (IPDs), was used to present stimuli with different depths, directions, centers, and rates of apparent motion. Many neurons appeared sensitive to dynamic IPDs, with responses at any given IPD depending strongly on the IPDs the stimulus had just passed through. However, it was the temporal pattern of the response, rather than the motion cues in the IPM, that determined sensitivity to features such as motion depth, direction, and center locus. IPM restricted only to the center of the IPD responsive area, evoked lower discharge rates than when the stimulus either moved through the IPD responsive area from outside, or up and down its flanks. When the stimulus was moved through the response area first in one direction and then back in the other, and the same IPDs evoked different responses, the response to the motion away from the center of the IPD responsive area was always lower than the response to the motion toward the center. When the IPD was closer at which the direction of motion reversed was to the center, the response to the following motion was lower. In no case did we find any evidence for neurons that under all conditions preferred one direction of motion to the other. We conclude that responses of IC neurons to IPM stimuli depend not on the history of stimulation, per se, but on the history of their response to stimulation, irrespective of the specific motion cues that evoke those responses. These data are consistent with the involvement of an adaptation mechanism that resides at or above the level of binaural integration. We conclude that our data provide no evidence for specialized motion detection involving dynamic IPD cues in the auditory midbrain of the mammal.

scholarly journals First Spike Latency Code for Interaural Phase Difference Discrimination in the Guinea Pig Inferior Colliculus

2011 ◽  
Vol 31 (25) ◽  
pp. 9192-9204 ◽  
Author(s):  
O. Zohar ◽  
T. M. Shackleton ◽  
I. Nelken ◽  
A. R. Palmer ◽  
M. Shamir
Keyword(s):  
Guinea Pig ◽  
Inferior Colliculus ◽  
Phase Difference ◽  
Interaural Phase ◽  
Interaural Phase Difference ◽  
Spike Latency

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)

Binaural interaction in low-frequency neurons in inferior colliculus of the cat. I. Effects of long interaural delays, intensity, and repetition rate on interaural delay function

1983 ◽  
Vol 50 (4) ◽  
pp. 981-999 ◽  
Author(s):  
S. Kuwada ◽  
T. C. Yin
Keyword(s):  
Inferior Colliculus ◽  
Repetition Rate ◽  
Single Cells ◽  
Continuous Distribution ◽  
Low Frequency ◽  
Intensity Difference ◽  
Onset Delay ◽  
Interaural Intensity Difference ◽  
Interaural Phase ◽  
Stimulus Offset

Detailed, quantitative studies were made of the interaural phase sensitivity of 197 neurons with low best frequency in the inferior colliculus (IC) of the barbiturate-anesthetized cat. We analyzed the responses of single cells to interaural delays in which tone bursts were delivered to the two ears via sealed earphones and the onset of the tone to one ear with respect to the other was varied. For most (80%) cells the discharge rate is a cyclic function of interaural delay at a period corresponding to that of the stimulating frequency. The cyclic nature of the interaural delay curve indicates that these cells are sensitive to the interaural phase difference. These cells are distributed throughout the low-frequency zone of the IC, but they are less numerous in the medial and caudal zones. Cells with a wide variety of response patterns will exhibit interaural phase sensitivities at stimulating frequencies up to 3,100 Hz, although above 2,500 Hz the number of such cells decrease markedly. Using dichotic stimuli we could study the cell's sensitivity to the onset delay and interaural phase independently. The large majority of IC cells respond only to changes in interaural phase, with no sensitivity to the onset delay. However, a small number (7%) of cells exhibit a sensitivity to the onset delay as well as to the interaural phase disparity, and most of these cells show an onset response. The effects of changing the stimulus intensity equally to both ears or of changing the interaural intensity difference on the mean interaural phase were studied. While some neurons are not affected by level changes, others exhibit systematic phase shifts for both average and interaural intensity variations, and there is a continuous distribution of sensitivities between these extremes. A few cells also showed systematic changes in the shape of the interaural delay curves as a function of interaural intensity difference, especially at very long delays. These shifts can be interpreted as a form of time-intensity trading. A few cells demonstrated orderly changes in the interaural delay curve as the repetition rate of the stimulus was varied. Some of these changes are consonant with an inhibitory effect that occurs at stimulus offset. The responses of the neurons show a strong bias for stimuli that would originate from he contralateral sound field; 77% of the responses display mean interaural phase angles that are less than 0.5 of a cycle, which are delays to the ipsilateral tone.(ABSTRACT TRUNCATED AT 400 WORDS)

Effects of interaural time delays of noise stimuli on low-frequency cells in the cat's inferior colliculus. I. Responses to wideband noise

1986 ◽  
Vol 55 (2) ◽  
pp. 280-300 ◽  
Author(s):  
T. C. Yin ◽  
J. C. Chan ◽  
D. R. Irvine
Keyword(s):  
Inferior Colliculus ◽  
Discharge Rate ◽  
Low Frequency ◽  
Central Peak ◽  
Stimulus Level ◽  
Central Nucleus ◽  
Interaural Phase ◽  
Wideband Noise ◽  
Pure Tones ◽  
Phase Differences

We examined the responses of low-frequency neurons in the central nucleus of the inferior colliculus (ICC) of the cat to interaurally delayed, wideband noise stimuli. The stimuli were pseudorandom noise signals that were generated digitally with a nominal bandwidth of 60-4,000 Hz. We also compared the responses to noise with those obtained from interaural phase differences of pure tones. We studied 144 neurons with characteristic frequencies below 2.5 kHz. Eighty-five percent (85%) of these were sensitive to changes in both interaural time differences (ITDs) of noise and interaural phase differences of pure tones, only 2% were sensitive to one stimulus but not the other, and the remainder were insensitive to both stimuli. For most cells the discharge rate was modulated in an approximately cyclic fashion by changes in ITDs of the wideband noise stimuli. The maximal spike counts often occurred near zero ITD, and there was considerable variability in the nature of the cycling, though it usually disappeared for ITDs greater than +/- 4,000 microseconds. The position of the central peak was usually (65%) within the physiologically relevant range of +/- 400 microseconds, and most (80%) occurred at positive ITDs, which corresponded to delays to the ipsilateral stimulus. In general, the shapes of the responses were not affected by changes in stimulus level above threshold. As long as identical noises were delivered to both ears, the responses were not sensitive to the particular noise stimulus used. When uncorrelated noises were delivered to the two ears, there was no sensitivity to ITDs. Composite curves were computed by linear summation of the responses to ITDs of pure tones at frequencies spaced at equal intervals throughout each cell's response area. The shapes of composite curves were similar to the responses of the same cell to ITDs of wideband noise stimuli. The positions of the central peaks of these two functions were highly correlated (r = 0.91, slope = 0.97). The values of characteristic delay and characteristic phase computed from the tonal responses were found to be good indicators of the shapes of the noise delay curves. Characteristic phases (CPs) near zero were associated with noise delay curves symmetric about the central peak, CPs near 0.5 cycles with those symmetric about the trough, while CPs between 0 and 0.5 or between 0.5 and 1.0 had noise delay curves that were asymmetric with a prominent trough to the left or right, respectively, of the central peak.(ABSTRACT TRUNCATED AT 400 WORDS)

Binaural interaction in low-frequency neurons in inferior colliculus of the cat. III. Effects of changing frequency

1983 ◽  
Vol 50 (4) ◽  
pp. 1020-1042 ◽  
Author(s):  
T. C. Yin ◽  
S. Kuwada
Keyword(s):  
Inferior Colliculus ◽  
Visual Inspection ◽  
Regression Line ◽  
Stimulus Frequency ◽  
Low Frequency ◽  
Analysis Procedure ◽  
Statistical Criterion ◽  
Phase Sensitivity ◽  
Interaural Phase ◽  
The Mean

The effects of changing stimulus frequency on the interaural phase sensitivity of neurons in the inferior colliculus (IC) were studied in barbiturate-anesthetized cats in order to reexamine the issue of characteristic delay (CD). Since the results obtained with the interaural delay and binaural beat stimuli are similar, we used the averaged interaural delay curves and binaural beat period histograms as comparable expressions of a neuron's interaural phase sensitivity. When the averaged interaural delay curves at different frequencies are plotted on a common time axis, for some cells the resulting superimposed delay curves show peaks or troughs that coincide at some CD. For most cells, though, this method of detecting a CD by visual inspection yields ambiguous and uncertain results. Composite curves, computed from the average of all the normalized superimposed delay curves, are also not helpful for showing CD. In order to provide a more objective means of analyzing the data, we plotted the mean interaural phase versus the stimulating frequency and computed the linear regression line, using the mean square error as a measure of linearity. The slope of the regression line is the CD for the neuron, and the phase intercept is referred to as the characteristic phase (CP). Cells that display a CD at the peak discharge have a CP = 0.0 cycles, while those that show a CD at the minimum discharge have a CP = 0.5. Cells that exhibit a CP at any value other than 0.0, 0.5, or 1.0 will have a CD at some relative amplitude other than the peak or trough. For cells that exhibit a CD at the peak or trough, results of the analysis procedure using the phase-frequency plot correspond to those obtained from visual inspection. For cells that do not show a common peak or trough, the analysis procedure not only specifies the location of the CD but also provides a statistical criterion of the linearity. From this analysis about 60% of the runs were identified as satisfying the criteria for CD at the P less than 0.005 level and 71% of these CDs are between +/- 300 micros. Most CD cells do not have the CD at the peak or trough of the response. Our results differ from those found in previous studies but they are in essential agreement with the original concept put forth by Rose et al. (31). Some cels exhibit little change in the CD or CP with variations in intensity, while others display marked systematic shifts in both CD and CP. In general, the peaks and troughs of the composite curves show less variability with intensity than the CD.(ABSTRACT TRUNCATED AT 400 WORDS)

The neural coding of auditory space

1989 ◽  
Vol 146 (1) ◽  
pp. 307-322 ◽  
Author(s):  
T. T. Takahashi
Keyword(s):  
Inferior Colliculus ◽  
Auditory System ◽  
Phase Difference ◽  
Neural Coding ◽  
Interaural Time Difference ◽  
Limited Range ◽  
Directional Sensitivity ◽  
Amplitude Difference ◽  
Interaural Phase ◽  
Interaural Phase Difference

The barn owl's auditory system computes interaural differences in time and amplitude and derives from them the horizontal and vertical coordinates of the sound source, respectively. Within the external nucleus of its inferior colliculus are auditory neurones, called ‘space-specific neurones’, that have spatial receptive fields. To activate a space-specific neurone, a sound must originate from a circumscribed region of space, or, if the sounds are delivered to each ear separately, using earphones, the stimuli must have the combination of interaural time and amplitude difference that simulates a sound broadcast from their receptive field. The sound-localization cues are processed in parallel, non-overlapping pathways extending from the cochlear nuclei to the subdivision of the inferior colliculus that innervates the space-specific neurones. Processing in the time pathway involves the coding of monaural phase angle, the derivation of sensitivity for interaural phase difference, and the calculation of interaural time difference (ITD) from interaural phase difference. The last process involves groups of neurones in the inferior colliculus whose collective firing signals a unique ITD, even though the activity of each constituent neurone signals multiple ITDs. The projections of these ensembles to the space-specific neurone endow the latter with a selectivity for ITD. Processing in the amplitude channel, about which less is known, initially involves an inhibitory process that sharpens the directional sensitivity of neurones in a lateral lemniscal nucleus. The inhibition is mediated by a commissural projection from the same lemniscal nucleus of the opposite side. At higher levels of the auditory system, neurones that are tuned to a limited range of interaural amplitude differences are found. It is proposed that at these higher stages, interaural amplitude difference, like ITD, is coded amidst an ensemble of neurones.

scholarly journals Temporal characteristics of neural sensitivities to the interaural phase difference in the inferior colliculus

2002 ◽  
Vol 23 (5) ◽  
pp. 286-286
Author(s):  
Shigeto Furukawa ◽  
Katuhiro Maki ◽  
Makio Kashino ◽  
Hiroshi Riquimaroux ◽  
Tatsuya Hirahara
Keyword(s):  
Inferior Colliculus ◽  
Phase Difference ◽  
Temporal Characteristics ◽  
Interaural Phase ◽  
Interaural Phase Difference

Desynchronizing Responses to Correlated Noise: A Mechanism for Binaural Masking Level Differences at the Inferior Colliculus

1999 ◽  
Vol 81 (2) ◽  
pp. 722-734 ◽  
Author(s):  
Alan R. Palmer ◽  
Dan Jiang ◽  
David McAlpine
Keyword(s):  
Inferior Colliculus ◽  
Low Frequency ◽  
Central Peak ◽  
Correlated Noise ◽  
Interaural Correlation ◽  
Correlation Models ◽  
Interaural Phase ◽  
Level Of Activity ◽  
Masking Level Difference ◽  
The Mean

Desynchronizing responses to correlated noise: a mechanism for binaural masking level differences at the inferior colliculus. We examined the adequacy of decorrelation of the responses to dichotic noise as an explanation for the binaural masking level difference (BMLD). The responses of 48 low-frequency neurons in the inferior colliculus of anesthetized guinea pigs were recorded to binaurally presented noise with various degrees of interaural correlation and to interaurally correlated noise in the presence of 500-Hz tones in either zero or π interaural phase. In response to fully correlated noise, neurons’ responses were modulated with interaural delay, showing quasiperiodic noise delay functions (NDFs) with a central peak and side peaks, separated by intervals roughly equivalent to the period of the neuron’s best frequency. For noise with zero interaural correlation (independent noises presented to each ear), neurons were insensitive to the interaural delay. Their NDFs were unmodulated, with the majority showing a level of activity approximately equal to the mean of the peaks and troughs of the NDF obtained with fully correlated noise. Partial decorrelation of the noise resulted in NDFs that were, in general, intermediate between the fully correlated and fully decorrelated noise. Presenting 500-Hz tones simultaneously with fully correlated noise also had the effect of demodulating the NDFs. In the case of tones with zero interaural phase, this demodulation appeared to be a saturation process, raising the discharge at all noise delays to that at the largest peak in the NDF. In the majority of neurons, presenting the tones in π phase had a similar effect on the NDFs to decorrelating the noise; the response was demodulated toward the mean of the peaks and troughs of the NDF. Thus the effect of added tones on the responses of delay-sensitive inferior colliculus neurons to noise could be accounted for by a desynchronizing effect. This result is entirely consistent with cross-correlation models of the BMLD. However, in some neurons, the effects of an added tone on the NDF appeared more extreme than the effect of decorrelating the noise, suggesting the possibility of additional inhibitory influences.

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