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

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)

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.


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)


1992 ◽  
Vol 67 (6) ◽  
pp. 1428-1436 ◽  
Author(s):  
A. Moiseff ◽  
T. Haresign

1. We studied the response of single units in the central nucleus of the inferior colliculus (ICc) of the barn owl (Tyto alba) to continuously varying interaural phase differences (IPDs) and static IPDs. Interaural phase was varied in two ways: continuously, by delivering tones to each ear that varied by a few hertz (binaural beat, Fig. 1), and discretely, by delaying in fixed steps the phase of sound delivered to one ear relative to the other (static phase). Static presentations were repeated at several IPDs to characterize interaural phase sensitivity. 2. Units sensitive to IPDs responded to the binaural beat stimulus over a broad range of delta f(Fig. 4). We selected a 3-Hz delta f for most of our comparative measurements on the basis of constraints imposed by our stimulus generation system and because it allowed us to reduce the influence of responses to stimulus onset and offset (Fig. 3A). 3. Characteristic interaural time or phase sensitivity obtained by the use of the binaural beat stimulus were comparable with those obtained by the use of the static technique (Fig. 5; r2 = 0.93, Fig. 6). 4. The binaural beat stimulus facilitated the measurement of characteristic delay (CD) and characteristic phase (CP) of auditory units. We demonstrated that units in the owl's inferior colliculus (IC) include those that are maximally excited by specific IPDs (CP = 0 or 1.0) as well as those that are maximally suppressed by specific IPDs (CP = 0.5; Figs. 7 and 8). 5. The selectivity of units sensitive to IPD or interaural time difference (ITD) were weakly influenced by interaural intensity difference (IID).(ABSTRACT TRUNCATED AT 250 WORDS)


1999 ◽  
Vol 81 (2) ◽  
pp. 722-734 ◽  
Author(s):  
Alan R. Palmer ◽  
Dan Jiang ◽  
David McAlpine

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.


2006 ◽  
Vol 95 (3) ◽  
pp. 1926-1935 ◽  
Author(s):  
Liang-Fa Liu ◽  
Alan R. Palmer ◽  
Mark N. Wallace

In the auditory system, some ascending pathways preserve the precise timing information present in a temporal code of frequency. This can be measured by studying responses that are phase-locked to the stimulus waveform. At each stage along a pathway, there is a reduction in the upper frequency limit of the phase-locking and an increase in the steady-state latency. In the guinea pig, phase-locked responses to pure tones have been described at various levels from auditory nerve to neocortex but not in the inferior colliculus (IC). Therefore we made recordings from 161 single units in guinea pig IC. Of these single units, 68% (110/161) showed phase-locked responses. Cells that phase-locked were mainly located in the central nucleus but also occurred in the dorsal cortex and external nucleus. The upper limiting frequency of phase-locking varied greatly between units (80−1,034 Hz) and between anatomical divisions. The upper limits in the three divisions were central nucleus, >1,000 Hz; dorsal cortex, 700 Hz; external nucleus, 320 Hz. The mean latencies also varied and were central nucleus, 8.2 ± 2.8 (SD) ms; dorsal cortex, 17.2 ms; external nucleus, 13.3 ms. We conclude that many cells in the central nucleus receive direct inputs from the brain stem, whereas cells in the external and dorsal divisions receive input from other structures that may include the forebrain.


1987 ◽  
Vol 57 (4) ◽  
pp. 1130-1147 ◽  
Author(s):  
M. N. Semple ◽  
L. M. Kitzes

The central auditory system could encode information about the location of a high-frequency sound source by comparing the sound pressure levels at the ears. Two potential computations are the interaural intensity difference (IID) and the average binaural intensity (ABI). In this study of the central nucleus of the inferior colliculus (ICC) of the anesthetized gerbil, we demonstrate that responses of 85% of the 97 single units in our sample were jointly influenced by IID and ABI. For a given ABI, discharge rate of most units is a sigmoidal function of IID, and peak rates occur at IIDs favoring the contralateral ear. Most commonly, successive increments of ABI cause successive shifts of the IID functions toward IIDs favoring the ipsilateral ear. Neurons displaying this behavior include many that would conventionally be classified EI (receiving predominantly excitatory input arising from one ear and inhibitory input from the other), many that would be classified EE (receiving predominantly excitatory input arising from each ear), and all that are responsive only to contralateral stimulation. The IID sensitivity of a very few EI neurons is unaffected by ABI, except near threshold. Such units could provide directional information that is independent of source intensity. A few EE neurons are very sensitive to ABI, but are minimally sensitive to IID. Nevertheless, our data indicate that responses of most EE units in ICC are strongly dominated by excitation of contralateral origin. For some units, discharge rate is nonmonotonically related to IID and is maximal when the stimuli at the two ears are of comparable sound pressure. This preference for zero IID is common for all binaural levels. Many EI neurons respond nonmonotonically to ABI. Discharge rates are greater for IIDs representative of contralateral space and are maximal at a single best ABI. For a subset of these neurons, the influence arising from the ipsilateral ear is comprised of a mixture of excitation and inhibition. As a consequence, discharge rates are nonmonotonically related not only to ABI but also to IID. This dual nonmonotonicity creates a clear focus of peak response at a particular ABI/IID combination. Because of their mixed monaural influences, such units would be ascribed to different classes of the conventional (EE/EI) binaural classification scheme depending on the binaural level presented. Several response classes were identified in this study, and each might contribute differently to the encoding of spatial information.(ABSTRACT TRUNCATED AT 400 WORDS)


1993 ◽  
Vol 70 (1) ◽  
pp. 64-80 ◽  
Author(s):  
R. Batra ◽  
S. Kuwada ◽  
T. R. Stanford

1. Localization of sounds has traditionally been considered to be performed by a duplex mechanism utilizing interaural temporal differences (ITDs) at low frequencies and interaural intensity differences at higher frequencies. More recently, it has been found that listeners can detect ITDs at high frequencies if the amplitude of the sound varies and an ITD is present in the envelope. Here we report the responses of neurons in the inferior colliculi of unanesthetized rabbits to ITDs of the envelopes of sinusoidally amplitude-modulated (SAM) tones. 2. Neurons were studied extracellularly with glass-coated Pt-Ir or Pt-W microelectrodes. Their sensitivity to ITDs in the envelopes of high-frequency sounds (> or = 2 kHz) was assessed using SAM tones that were presented binaurally. The tones at the two ears had the same carrier frequency but modulation frequencies that differed by 1 Hz. This caused a cyclic variation in the ITD produced by the envelope. In this "binaural SAM" stimulus, the carriers caused no ITD because they were in phase. In addition to the binaural SAM stimulus, pure tones were used to investigate responses to ipsilateral and contralateral stimulation and the nature of the interaction during binaural stimulation. 3. Neurons tended to display one of two kinds of sensitivity to ITDs. Some neurons discharged maximally at the same ITD at all modulation frequencies > 250 Hz (peak-type neurons), whereas others were maximally suppressed at the same ITD (trough-type neurons). 4. At these higher modulation frequencies (> 250 Hz), the characteristic delays that neurons exhibited tended to lie within the range that a rabbit might normally encounter (+/- 300 microseconds). The peak-type neurons favored ipsilateral delays, which correspond to sounds in the contralateral sound field. The trough-type neurons showed no such preference. 5. The preference of peak-type neurons for a particular delay was sharper than that of trough-type neurons and was comparable to that observed in neurons of the inferior colliculus that are sensitive to delays of low-frequency pure tones. 6. At lower modulation frequencies (< 150 Hz) characteristic delays often lay beyond +/- 300 microseconds. 7. Increasing the ipsilateral intensity tended to shift the preferred delay ipsilaterally at lower (< 250 Hz), but not at higher, modulation frequencies. 8. When tested with pure tones, a substantial number of peak-type neurons were found to be excited by contralateral stimulation but inhibited by ipsilateral stimulation.(ABSTRACT TRUNCATED AT 400 WORDS)


1997 ◽  
Vol 77 (6) ◽  
pp. 2945-2965 ◽  
Author(s):  
Adrian Rees ◽  
Ali Sarbaz ◽  
Manuel S. Malmierca ◽  
Fiona E. N. Le Beau

Rees, Adrian, Ali Sarbaz, Manuel S. Malmierca, and Fiona E. N. Le Beau. Regularity of firing of neurons in the inferior colliculus. J. Neurophysiol. 77: 2945–2965, 1997. The spike discharge regularity of 254 tonically firing units in the inferior colliculus (IC) of the anesthetized guinea pig was studied in response to tones presented at best frequency (BF) to the ear contralateral to the recorded IC. Regularity of firing was measured by calculating the coefficient of variation (CV) as a function of time over the course of a unit's response. Two hundred and fifteen units (56 under urethan and 159 under chloralose anesthesia) in the central nucleus of the IC (CNIC) were studied in detail. In response to tones at 15–25 dB above threshold, 80% of units in the urethan sample fired regularly (CV < 0.5) during their sustained response, and 46% were highly regular (CV ≤ 0.35). For chloralose the values were 68% and 23%, respectively. Units recorded under urethan were significantly more regular than those recorded under chloralose. For units in the sample with a measurable onset CV, 63% were regular and 44% highly regular under urethan, and 73% were regular and 54% highly regular under chloralose. The units' peristimulus time histogram (PSTH) patterns were classified into subdivisions of four categories: choppers [9%: chop-sustained (Cs), chop-onset (Co)]; pausers [42%: pauser-chop-sustained(P/Cs), pauser-chop-onset (P/Co), pauser-no-chop]; on-sustained(43%: primary-type, L-type, h-type); and sustained (6%). The presence of chopping was a reliable predictor of regularity: Cs and P/Cs units were highly regular throughout their response, whereas Co and P/Co units were highly regular at onset and became less regular. Some units in the other PSTH categories were highly regular despite the absence of chopping, and units with virtually identical PSTHs showed very different sustained CVs. Regularity was measured as a function of firing rate in 71 units. In 23%, regularity remained constant when firing rate changed with stimulus level. Forty-six percent fired more regularly as firing rate increased, 8% fired less regularly, and 23% of units showed no consistent relationship between CV and firing rate. Regularity did not correlate with the neurons' frequency response areas or BFs. Regular firing was also found in a smaller sample of units recorded in cortices surrounding the CNIC. We conclude that regular firing is a characteristic feature of most neurons in the IC. Regularity is a specific feature correlated with four PSTH types (Cs, Co, P/Cs, and P/Co). Other PSTH types may or may not exhibit regularity.


1986 ◽  
Vol 55 (3) ◽  
pp. 587-603 ◽  
Author(s):  
M. B. Calford ◽  
D. R. Moore ◽  
M. E. Hutchings

Recordings of response to free-field stimuli at best frequency were made from single units in the central nucleus of the inferior colliculus of anesthetized cats. Stimulus position was varied in azimuth, and the responses of units were compared with variation in the intensity and arrival time of the sound at each ear, derived from cochlear microphonic (CM) recordings. CM recordings were made at each frequency and at every point in space for which single-unit data were collected. Interaural time difference (delay) increased monotonically, but not linearly, as the stimulus was moved away from the midline. However, a given delay did not represent a single azimuth across frequency. Low-frequency interaural intensity differences (IIDs) were monotonic across azimuth and peaked at, or near, the poles. Higher-frequency IIDs were nonmonotonic and peaked relatively close to the midline, decreasing toward the poles. Units that showed little variation in discharge across azimuth formed 28% of the sample and were classified as omnidirectional. For other units, the spike-count intensity function and the variation of the CM with azimuth were combined to form a derived monaural azimuth function. For 29% of those units showing azimuthal sensitivity, the derived monaural azimuth function matched the actual azimuth function. This suggested that these units received input from only one ear. The largest group of azimuthally sensitive units (47%) was formed from those units inferred to be IID sensitive. At higher frequencies these units displayed a peaked azimuth function paralleling the nonmonotonic relation of IID to azimuth. The proportion of inferred IID-sensitive units was close to that found in dichotic studies.


1995 ◽  
Vol 73 (2) ◽  
pp. 449-467 ◽  
Author(s):  
R. Snyder ◽  
P. Leake ◽  
S. Rebscher ◽  
R. Beitel

1. Cochlear implants have been available for > 20 yr to profoundly deaf adults who have lost their hearing after acquiring language. The success of these cochlear prostheses has encouraged the application of implants in prelingually deaf children as young as 2 yr old. To further characterize the consequences of chronic intracochlear electrical stimulation (ICES) on the developing auditory system, the temporal-response properties of single neurons in the inferior colliculus (IC) were recorded in deafened anesthetized cats. 2. The neurons were excited by unilateral ICES with the use of a scala tympani stimulating electrode implanted in the left cochlea. The electrodes were modeled after those used in cochlear implant patients. Responses of 443 units were recorded extracellularly in the contralateral (right) IC with the use of tungsten microelectrodes. Recordings were made in three groups of adult animals: neonatally deafened/chronically stimulated animals (192 units), neonatally deafened/unstimulated animals (80 units), and adult-deafened/prior normal-hearing animals (171 units). The neonatally deafened cats were deafened by multiple intramuscular injections of neomycin sulfate and never developed demonstrable hearing. Most of the deafened, chronically stimulated animals were implanted at 6 wk of age and stimulated at suprathreshold levels for 4 h/day for 3-6 mo. The unstimulated animals were implanted as adults at least 2 wk before the acute physiological experiment and were left unstimulated until the acute experiment was conducted. Prior-normal adults were deafened and implanted at least 2 wk before the acute experiment. 3. IC units were isolated with the use of a search stimulus consisting of three cycles of a 100-Hz sinusoid. Most units responded to sinusoidal stimulation with either an onset response or a sustained response. Onset units were the predominant unit found in the external nucleus, whereas sustained units were found almost exclusively in the central nucleus. The temporal resolution of sustained response units was measured with the use of pulse trains of increasing frequency and calculating the discharges/pulse. 4. The range of electrical pulse frequencies to which IC units responded in a temporally synchronized manner was comparable with that produced by acoustic stimulation. The discharge rate/pulse-versus-pulse frequency transfer functions of IC units were uniformly low-pass, although they varied widely in their cutoff frequencies. This variation in pulse response was partially correlated with the unit's response to sinusoids. Most onset neurons responded only to pulse frequencies below 20 pulses per second (pps). Most sustained units responded best to pulse frequencies < 100 pps, and most ceased to respond to pulse frequencies > 300 pps.(ABSTRACT TRUNCATED AT 400 WORDS)


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