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

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.

2008 ◽  
Vol 99 (4) ◽  
pp. 1942-1952 ◽  
Author(s):  
Philip X. Joris ◽  
Dries H. Louage ◽  
Marcel van der Heijden

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.


1987 ◽  
Vol 58 (3) ◽  
pp. 543-561 ◽  
Author(s):  
J. C. Chan ◽  
T. C. Yin ◽  
A. D. Musicant

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.


2005 ◽  
Vol 93 (6) ◽  
pp. 3390-3400 ◽  
Author(s):  
W. R. D’Angelo ◽  
S. J. Sterbing ◽  
E.-M. Ostapoff ◽  
S. Kuwada

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.


2005 ◽  
Vol 94 (2) ◽  
pp. 1180-1198 ◽  
Author(s):  
Courtney C. Lane ◽  
Bertrand Delgutte

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.


2007 ◽  
Vol 98 (5) ◽  
pp. 2566-2579 ◽  
Author(s):  
Myles Mc Laughlin ◽  
Bram Van de Sande ◽  
Marcel van der Heijden ◽  
Philip X. Joris

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.


1987 ◽  
Vol 58 (3) ◽  
pp. 562-583 ◽  
Author(s):  
T. C. Yin ◽  
J. C. Chan ◽  
L. H. Carney

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)


1997 ◽  
Vol 77 (6) ◽  
pp. 3085-3106 ◽  
Author(s):  
Dan Jiang ◽  
David McAlpine ◽  
Alan R. Palmer

Jiang, Dan, David McAlpine, and Alan R. Palmer. Responses of neurons in the inferior colliculus to binaural masking level difference stimuli measured by rate-versus-level functions. J. Neurophysiol. 77: 3085–3106, 1997. The psychophysical detection threshold of a low-frequency tone masked by broadband noise is reduced by ≤15 dB by inversion of the tone in one ear (called the binaural masking level difference: BMLD). The contribution of 120 low-frequency neurons (best frequencies 168–2,090 Hz) in the inferior colliculus (ICC) of the guinea pig to binaural unmasking of 500-Hz tones masked by broadband noise was examined. We measured rate-level functions of the responses to identical signals (So) and noise (No) at the two ears (NoSo) and to identical noise but with the signal inverted at one ear (NoSπ): the noise was 7–15 dB suprathreshold. The masked threshold was estimated by the standard separation, “D”. The neural BMLD was estimated as the difference between the masked thresholds for NoSo and NoSπ. The presence of So and Sπ tones was indicated by discharge rate increases in 55.3% of neurons. In 36.4% of neurons, the presence of So tones was indicated by an increase in discharge rate and Sπ tones by a decrease. In 6.8% of neurons, both So and Sπ tones caused a decrease in discharge rate. In only 1.5% of neurons was So indicated by a decrease and Sπ by an increase in discharge rate. Responses to the binaural configurations were consistent with the neuron's interaural delay sensitivities; 34.4% of neurons showing increases in discharge rate to both So and Sπ tones gave positive BMLDs ≥3 dB (Sπ tones were detected at lower levels than So), whereas 37.3% gave negative BMLDs ≥3 dB. For neurons in which So signals caused an increase in the discharge rate and Sπ a decrease, 72.7% gave positive BMLDs ≥3 dB and only 4.5% gave negative BMLDs ≥3 dB. The results suggest that the responses of single ICC neurons are consistent with the psychophysical BMLDs for NoSo versus NoSπ at 500 Hz, and with current binaural interaction models based on coincidence detection. The neurons likely to contribute to the psychophysical BMLD are those with BFs near 500 Hz, but detection of So and Sπ tones may depend on different populations of neurons.


2018 ◽  
Vol 30 (7) ◽  
pp. 1801-1829
Author(s):  
Irina Higgins ◽  
Simon Stringer ◽  
Jan Schnupp

It is well known that auditory nerve (AN) fibers overcome bandwidth limitations through the volley principle, a form of multiplexing. What is less well known is that the volley principle introduces a degree of unpredictability into AN neural firing patterns that may be affecting even simple stimulus categorization learning. We use a physiologically grounded, unsupervised spiking neural network model of the auditory brain with spike time dependent plasticity learning to demonstrate that plastic auditory cortex is unable to learn even simple auditory object categories when exposed to the raw AN firing input without subcortical preprocessing. We then demonstrate the importance of nonplastic subcortical preprocessing within the cochlear nucleus and the inferior colliculus for stabilizing and denoising AN responses. Such preprocessing enables the plastic auditory cortex to learn efficient robust representations of the auditory object categories. The biological realism of our model makes it suitable for generating neurophysiologically testable hypotheses.


2016 ◽  
Author(s):  
Irina Higgins ◽  
Simon Stringer ◽  
Jan Schnupp

AbstractIt is well known that auditory nerve (AN) fibers overcome bandwidth limitations through the “volley principle”, a form of multiplexing. What is less well known is that the volley principle introduces a degree of unpredictability into AN neural firing patterns which makes even simple stimulus categorization tasks difficult. We use a physiologically grounded, unsupervised spiking neural network model of the auditory brain with STDP learning to demonstrate that plastic auditory cortex is unable to learn even simple auditory object categories when exposed to the raw AN firing input without subcortical preprocessing. We then demonstrate the importance of non-plastic subcortical preprocessing within the cochlear nucleus (CN) and the inferior colliculus (IC) for stabilising and denoising AN responses. Such preprocessing enables the plastic auditory cortex to learn efficient robust representations of the auditory object categories. The biological realism of our model makes it suitable for generating neurophysiologically testable hypotheses.


2005 ◽  
Vol 93 (4) ◽  
pp. 1857-1870 ◽  
Author(s):  
Philip X. Joris ◽  
Bram Van De Sande ◽  
Marcel van der Heijden

Many cells in the inferior colliculus (IC) are sensitive to interaural time differences (ITDs), in the form of an oscillatory dependency of average firing rate on ITD. We studied the degree of damping in such binaural responses, recording from neurons in the inferior colliculus of pentobarbital-anesthetized cats to binaural broadband noise and tones. Noise-delay functions and composite curves were characterized by computing the difference between responses to correlated and anticorrelated stimuli. We use a new metric, based on the envelope of this difference, to quantify damping. There is a clear relationship between damping and characteristic frequency (CF), but even neurons of the same CF can differ in their damping. For individual cells, damping can be stronger to tones or to noise; at the population level the two are positively correlated and are scarcely affected by SPL. The frequencies that dominate ITD sensitivity are near the CF in response to noise, but are often below CF in response to tones. These findings qualify conclusions from earlier reports but overall they support the conclusion that, at a population level, basic aspects of binaural responses to wideband noise are consistent with summed responses to pure tones.


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