Detection of interaural correlation by neurons in the superior olivary complex, inferior colliculus and auditory cortex of the unanesthetized rabbit

2006 ◽  
Vol 221 (1-2) ◽  
pp. 1-16 ◽  
Author(s):  
Charles S. Coffey ◽  
Charles S. Ebert ◽  
Allen F. Marshall ◽  
John D. Skaggs ◽  
Stephanie E. Falk ◽  
...  
2006 ◽  
Vol 95 (3) ◽  
pp. 1309-1322 ◽  
Author(s):  
Shigeyuki Kuwada ◽  
Douglas C. Fitzpatrick ◽  
Ranjan Batra ◽  
Ernst-Michael Ostapoff

Interaural time differences, a cue for azimuthal sound location, are first encoded in the superior olivary complex (SOC), and this information is then conveyed to the dorsal nucleus of the lateral lemniscus (DNLL) and inferior colliculus (IC). The DNLL provides a strong inhibitory input to the IC and may serve to transform the coding of interaural time differences (ITDs) in the IC. Consistent with the projections from the SOC, the DNLL and IC had similar distributions of peak- and trough-type neurons, characteristic delays, and best ITDs. The ITD tuning widths of DNLL neurons were intermediate between those of the SOC and IC. Further sharpening is seen in the auditory thalamus, indicating that sharpening mechanisms are not restricted to the midbrain. The proportion of neurons that phase-locked to the tones delivered to each ear progressively decreased from the SOC to the auditory thalamus. The degree of phase-locking for a large majority of DNLL neurons was too weak to support their involvement in processing monaural inputs to generate a sensitivity to ITDs. The response rates of DNLL neurons were on average ∼60% greater than in the IC or SOC, indicating that the inhibitory input provided to the IC by the DNLL is robust.


1995 ◽  
Vol 74 (6) ◽  
pp. 2469-2486 ◽  
Author(s):  
D. C. Fitzpatrick ◽  
S. Kuwada ◽  
R. Batra ◽  
C. Trahiotis

1. In most natural environments, sound waves from a single source will reach a listener through both direct and reflected paths. Sound traveling the direct path arrives first, and determines the perceived location of the source despite the presence of reflections from many different locations. This phenomenon is called the "law of the first wavefront" or "precedence effect." The time at which the reflection is first perceived as a separately localizable sound defines the end of the precedence window and is called "echo threshold." The precedence effect represents an important property of the auditory system, the neural basis for which has only recently begun to be examined. Here we report the responses of single neurons in the inferior colliculus (IC) and superior olivary complex (SOC) of the unanesthetized rabbit to a sound and its simulated reflection. 2. Stimuli were pairs of monaural or binaural clicks delivered through earphones. The leading click, or conditioner, simulated a direct sound, and the lagging click, or probe, simulated a reflection. Interaural time differences (ITDs) were introduced in the binaural conditioners and probes to adjust their simulated locations. The probe was always set at the neuron's best ITD, whereas the conditioner was set at the neuron's best ITD or its worst ITD. To measure the time course of the effects of the conditioner on the probe, we examined the response to the probe as a function of the conditioner-probe interval (CPI). 3. When IC neurons were tested with conditioners and probes set at the neuron's best ITD, the response to the probe as a function of CPI had one of two forms: early-low or early-high. In early-low neurons the response to the probe was initially suppressed but recovered monotonically at longer CPIs. Early-high neurons showed a nonmonotonic recovery pattern. In these neurons the maximal suppression did not occur at the shortest CPIs, but rather after a period of less suppression. Beyond this point, recovery was similar to that of early-low neurons. The presence of early-high neurons meant that the overall population was never entirely suppressed, even at short CPIs. Taken as a whole. CPIs for 50% recovery of the response to the probe among neurons ranged from 1 to 64 ms with a median of approximately 6 ms. 4. The above results are consistent with the time course of the precedence effect for the following reasons. 1) The lack of complete suppression at any CPI is compatible with behavioral results that show the presence of a probe can be detected even at short CPIs when it is not separately localizable. 2) At a CPI corresponding to echo threshold for human listeners (approximately 4 ms CPI) there was a considerable response to the probe, consistent with it being heard as a separately localizable sound at this CPI. 3) Full recovery for all neurons required a period much longer than that associated with the precedence effect. This is consistent with the relatively long time required for conditioners and probes to be heard with equal loudness. 5. Conditioners with either the best ITD or worst ITD were used to determine the effect of ITD on the response to the probe. The relative amounts of suppression caused by the two ITDs varied among neurons. Some neurons were suppressed about equally by both types of conditioners, others were suppressed more by a conditioner with the best ITD, and still others by a conditioner with the worst ITD. Because the best ITD and worst ITD presumably activate different pathways, these results suggest that different neurons receive a different balance of inhibition from different sources. 6. The recovery functions of neurons not sensitive to ITDs were similar to those of ITD-sensitive, neurons. This suggests that the time course of suppression may be common among different IC populations. 7. We also studied neurons in the SOC. Although many showed binaural interactions, none were sensitive to ITDs. Thus the response of this population may not be


2020 ◽  
Vol 123 (2) ◽  
pp. 695-706
Author(s):  
Lu Luo ◽  
Na Xu ◽  
Qian Wang ◽  
Liang Li

The central mechanisms underlying binaural unmasking for spectrally overlapping concurrent sounds, which are unresolved in the peripheral auditory system, remain largely unknown. In this study, frequency-following responses (FFRs) to two binaurally presented independent narrowband noises (NBNs) with overlapping spectra were recorded simultaneously in the inferior colliculus (IC) and auditory cortex (AC) in anesthetized rats. The results showed that for both IC FFRs and AC FFRs, introducing an interaural time difference (ITD) disparity between the two concurrent NBNs enhanced the representation fidelity, reflected by the increased coherence between the responses evoked by double-NBN stimulation and the responses evoked by single NBNs. The ITD disparity effect varied across frequency bands, being more marked for higher frequency bands in the IC and lower frequency bands in the AC. Moreover, the coherence between IC responses and AC responses was also enhanced by the ITD disparity, and the enhancement was most prominent for low-frequency bands and the IC and the AC on the same side. These results suggest a critical role of the ITD cue in the neural segregation of spectrotemporally overlapping sounds. NEW & NOTEWORTHY When two spectrally overlapped narrowband noises are presented at the same time with the same sound-pressure level, they mask each other. Introducing a disparity in interaural time difference between these two narrowband noises improves the accuracy of the neural representation of individual sounds in both the inferior colliculus and the auditory cortex. The lower frequency signal transformation from the inferior colliculus to the auditory cortex on the same side is also enhanced, showing the effect of binaural unmasking.


1989 ◽  
Vol 61 (2) ◽  
pp. 257-268 ◽  
Author(s):  
R. Batra ◽  
S. Kuwada ◽  
T. R. Stanford

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)


1990 ◽  
Vol 64 (2) ◽  
pp. 582-597 ◽  
Author(s):  
B. J. Blatchley ◽  
J. F. Brugge

1. Responses of single neurons to monaural or binaural CF tones delivered through a closed and calibrated sound delivery system were studied in the central nucleus of the inferior colliculus (ICC) in ketamine and barbiturate-anesthetized kittens 4-105 days old. 2. Neurons from young kittens had elevated thresholds, some greater than 100 dB in the youngest kittens tested. Average thresholds in the ICC matched those previously measured in the auditory nerve (AN), cochlear nuclei (CN), and auditory cortex (CTX), suggesting that the drop in threshold as a function of age is primarily determined by development at the periphery. 3. Minimal first-spike latencies were relatively long in the youngest kittens, approaching adult values by the end of the third postnatal week. Latencies were distributed between values previously determined for the CN and auditory cortex and can be attributed to the centripetal development of the auditory system. 4. The range of frequencies that were effective in exciting ICC neurons was restricted in young kittens. Neurons having characteristic frequencies (CFs) greater than 7 kHz were not recorded before postnatal day 10. CF distribution matched that obtained in previous experiments from AN, CN, and CTX, reflecting the development of the cochlea. 5. Both monotonic and nonomonotonic spike count-versus-intensity functions were found in the youngest kittens. There was a tendency for monotonic functions from the youngest kittens to be steeper than those from older kittens. No age-related changes in the shapes of non-monotonic functions were found. 6. Sensitivity to interaural intensity difference (IID), tested by holding the intensity to the excitatory ear at a suprathreshold level and increasing the intensity of the stimulus to the inhibitory ear, was exhibited as early as 8 days after birth. The majority of the cells exhibiting sensitivity to IID (89.5%) were classified as EI cells, and almost all IID sensitive cells had CFs between 3 and 25 kHz. Within our sample the shapes of IID functions, as well as the operating range of the IID functions, closely resembled those obtained from the adult cat. Thresholds of excitation and of inhibition were highly correlated, suggesting that the ipsilateral and contralateral inputs to the ICC develop as a matched set. 7. Sensitivity to interaural phase difference (IPD), tested either by shifting the onset phase of a CF tone to one ear relative to the other or by presenting tones of slightly different frequency to the two ears, was present as early as 12 days after birth.(ABSTRACT TRUNCATED AT 400 WORDS)


2010 ◽  
Vol 103 (4) ◽  
pp. 2050-2061 ◽  
Author(s):  
Kyle T. Nakamoto ◽  
Trevor M. Shackleton ◽  
Alan R. Palmer

One of the fundamental questions of auditory research is how sounds are segregated because, in natural environments, multiple sounds tend to occur at the same time. Concurrent sounds, such as two talkers, physically add together and arrive at the ear as a single input sound wave. The auditory system easily segregates this input into a coherent percept of each of the multiple sources. A common feature of speech and communication calls is their harmonic structure and in this report we used two harmonic complexes to study the role of the corticofugal pathway in the processing of concurrent sounds. We demonstrate that, in the inferior colliculus (IC) of the anesthetized guinea pig, deactivation of the auditory cortex altered the temporal and/or the spike response to the concurrent, monaural harmonic complexes. More specifically, deactivating the auditory cortex altered the representation of the relative level of the complexes. This suggests that the auditory cortex modulates the representation of the level of two harmonic complexes in the IC. Since sound level is a cue used in the segregation of auditory input, the corticofugal pathway may play a role in this segregation.


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