Binaural summation of amplitude modulation involves weak interaural suppression.
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ABSTRACT:
SUBMITTER: Baker DH
PROVIDER: S-EPMC7044261 | biostudies-literature |
REPOSITORIES: biostudies-literature
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Project description:The subjective representation of the sounds delivered to the two ears of a human listener is closely associated with the interaural delay and correlation of these two-ear sounds. When the two-ear sounds, e.g., arbitrary noises, arrive simultaneously, the single auditory image of the binaurally identical noises becomes increasingly diffuse, and eventually separates into two auditory images as the interaural correlation decreases. When the interaural delay increases from zero to several milliseconds, the auditory image of the binaurally identical noises also changes from a single image to two distinct images. However, measuring the effect of these two factors on an identical group of participants has not been investigated. This study examined the impacts of interaural correlation and delay on detecting a binaurally uncorrelated fragment (interaural correlation = 0) embedded in the binaurally correlated noises (i.e., binaural gap or break in interaural correlation). We found that the minimum duration of the binaural gap for its detection (i.e., duration threshold) increased exponentially as the interaural delay between the binaurally identical noises increased linearly from 0 to 8 ms. When no interaural delay was introduced, the duration threshold also increased exponentially as the interaural correlation of the binaurally correlated noises decreased linearly from 1 to 0.4. A linear relationship between the effect of interaural delay and that of interaural correlation was described for listeners participating in this study: a 1 ms increase in interaural delay appeared to correspond to a 0.07 decrease in interaural correlation specific to raising the duration threshold. Our results imply that a tradeoff may exist between the impacts of interaural correlation and interaural delay on the subjective representation of sounds delivered to two human ears.
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Project description:Understanding binaural perception requires detailed analyses of the neural circuitry responsible for the computation of interaural time differences (ITDs). In the avian brainstem, this circuit consists of internal axonal delay lines innervating an array of coincidence detector neurons that encode external ITDs. Nucleus magnocellularis (NM) neurons project to the dorsal dendritic field of the ipsilateral nucleus laminaris (NL) and to the ventral field of the contralateral NL. Contralateral-projecting axons form a delay line system along a band of NL neurons. Binaural acoustic signals in the form of phase-locked action potentials from NM cells arrive at NL and establish a topographic map of sound source location along the azimuth. These pathways are assumed to represent a circuit similar to the Jeffress model of sound localization, establishing a place code along an isofrequency contour of NL. Three-dimensional measurements of axon lengths reveal major discrepancies with the current model; the temporal offset based on conduction length alone makes encoding of physiological ITDs impossible. However, axon diameter and distances between Nodes of Ranvier also influence signal propagation times along an axon. Our measurements of these parameters reveal that diameter and internode distance can compensate for the temporal offset inferred from axon lengths alone. Together with other recent studies, these unexpected results should inspire new thinking on the cellular biology, evolution, and plasticity of the circuitry underlying low-frequency sound localization in both birds and mammals.
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Project description:Binaural integration in the central nucleus of inferior colliculus (ICC) plays a critical role in sound localization. However, its arithmetic nature and underlying synaptic mechanisms remain unclear. Here, we showed in mouse ICC neurons that the contralateral dominance is created by a "push-pull"-like mechanism, with contralaterally dominant excitation and more bilaterally balanced inhibition. Importantly, binaural spiking response is generated apparently from an ipsilaterally mediated scaling of contralateral response, leaving frequency tuning unchanged. This scaling effect is attributed to a divisive attenuation of contralaterally evoked synaptic excitation onto ICC neurons with their inhibition largely unaffected. Thus, a gain control mediates the linear transformation from monaural to binaural spike responses. The gain value is modulated by interaural level difference (ILD) primarily through scaling excitation to different levels. The ILD-dependent synaptic scaling and gain adjustment allow ICC neurons to dynamically encode interaural sound localization cues while maintaining an invariant representation of other independent sound attributes.
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Project description:Naturally occurring sounds are routinely periodic. The ability to phase-lock to such periodicity facilitates pitch perception and interaural time differences (ITDs) determination in binaural localization. We examined whether deficient pitch processing in individuals with congenital amusia (tone deafness) is accompanied by impaired ability to lateralize musical pitch at auditory periphery and memorize the location of pitch at the working memory level. If common mechanisms subserve processing of temporal-fine-structure based pitch and ITDs, then deficient processing of one feature should impair performance on the other. Thus, we measured ITD discrimination thresholds using an adaptive-tracking procedure for lateralizing musical tone pairs separated by different semitone intervals. Amusic individuals exhibited normal ITD thresholds comparable to those of matched controls, which were not affected by concurrent pitch changes. For working memory tasks, the amusic group performed significantly worse than matched controls in probed pitch recall, irrespective of the complexity level of concurrent variations along the ITD dimension of the melodic sequence. Interestingly, despite normal peripheral ITD thresholds, amusic individuals performed worse than controls in recalling probed locations of tones within a sequence of musical notes originating from different ITD-simulated locations. Findings suggest that individuals with congenital amusia are unimpaired in temporal fine-structure encoding to determine the location of musical pitch based on binaural ITD information at the auditory periphery. However, working memory for a sequence of sounds' ITD-dependent spatial location is here shown to be impaired and dissociated from the pitch feature of sounds at the working memory level.
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Project description:Interaural time difference (ITD) arises whenever a sound outside of the median plane arrives at the two ears. There is evidence that ITD in the rapidly varying fine structure of a sound is most important for sound localization and for understanding speech in noise. Cochlear implants (CIs), neural prosthetic devices that restore hearing in the profoundly deaf, are increasingly implanted to both ears to provide implantees with the advantages of binaural hearing. CI listeners have been shown to be sensitive to fine structure ITD at low pulse rates, but their sensitivity declines at higher pulse rates that are required for speech coding. We hypothesize that this limitation in electric stimulation is at least partially due to binaural adaptation associated with periodic stimulation. Here, we show that introducing binaurally synchronized jitter in the stimulation timing causes large improvements in ITD sensitivity at higher pulse rates. Our experimental results demonstrate that a purely temporal trigger can cause recovery from binaural adaptation. Thus, binaurally jittered stimulation may improve several aspects of binaural hearing in bilateral recipients of neural auditory prostheses.
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Project description:Neurons in the medial superior olive (MSO) enable sound localization by their remarkable sensitivity to submillisecond interaural time differences (ITDs). Each MSO neuron has its own "best ITD" to which it responds optimally. A difference in physical path length of the excitatory inputs from both ears cannot fully account for the ITD tuning of MSO neurons. As a result, it is still debated how these inputs interact and whether the segregation of inputs to opposite dendrites, well-timed synaptic inhibition, or asymmetries in synaptic potentials or cellular morphology further optimize coincidence detection or ITD tuning. Using in vivo whole-cell and juxtacellular recordings, we show here that ITD tuning of MSO neurons is determined by the timing of their excitatory inputs. The inputs from both ears sum linearly, whereas spike probability depends nonlinearly on the size of synaptic inputs. This simple coincidence detection scheme thus makes accurate sound localization possible.
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Project description:When different components of a stimulus carry different binaural information, processing of binaural information in a target component is often affected. The present experiments examine whether such interference is affected by amplitude modulation and the relative phase of modulation of the target and distractors. In all experiments, listeners attempted to discriminate interaural time differences of a target stimulus in the presence of distractor stimuli with ITD=0. In Experiment 1, modulation of the distractors but not the target reduced interference between components. In Experiment 2, synthesized musical notes exhibited little binaural interference when there were slight asynchronies between different streams of notes (31 or 62 ms). The remaining experiments suggested that the reduction in binaural interference in the previous experiments was due neither to the complex spectra of the synthesized notes nor to greater detectability of the target in the presence of modulated distractors. These data suggest that this interference is reduced when components are modulated in ways that result in the target appearing briefly in isolation, not because of segregation cues. These data also suggest that modulation and asynchronies between modulators that might be encountered in real-world listening situations are adequate to reduce binaural interference to inconsequential levels.
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Project description:Sensory information travels along feedforward connections through a hierarchy of cortical areas, which, in turn, send feedback connections to lower-order areas. Feedback has been implicated in attention, expectation, and sensory context, but the mechanisms underlying these diverse feedback functions are unknown. Using specific optogenetic inactivation of feedback connections from the secondary visual area (V2), we show how feedback affects neural responses in the primate primary visual cortex (V1). Reducing feedback activity increases V1 cells' receptive field (RF) size, decreases their responses to stimuli confined to the RF, and increases their responses to stimuli extending into the proximal surround, therefore reducing surround suppression. Moreover, stronger reduction of V2 feedback activity leads to progressive increase in RF size and decrease in response amplitude, an effect predicted by a recurrent network model. Our results indicate that feedback modulates RF size, surround suppression and response amplitude, similar to the modulatory effects of visual spatial attention.
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Project description:For amplitude-modulated sound, the envelope interaural time difference (ITDENV) is a potential cue for sound-source location. ITDENV is encoded in the lateral superior olive (LSO) of the auditory brainstem, by excitatory-inhibitory (EI) neurons receiving ipsilateral excitation and contralateral inhibition. Between human listeners, sensitivity to ITDENV varies considerably, but ultimately decreases with increasing stimulus carrier frequency, and decreases more strongly with increasing modulation rate. Mechanisms underlying the variation in behavioral sensitivity remain unclear. Here, with increasing carrier frequency (4-10 kHz), as we phenomenologically model the associated decrease in ITDENV sensitivity using arbitrarily fewer neurons consistent across populations, we computationally model the variable sensitivity across human listeners and modulation rates (32-800 Hz) as the decreasing range of membrane frequency responses in LSO neurons. Transposed tones stimulate a bilateral auditory-periphery model, driving model EI neurons where electrical membrane impedance filters the frequency content of inputs driven by amplitude-modulated sound, evoking modulation filtering. Calculated from Fisher information in spike-rate functions of ITDENV, for model EI neuronal populations distinctly reflecting the LSO range in membrane frequency responses, just-noticeable differences in ITDENV collectively reproduce the largest variation in ITDENV sensitivity across human listeners. These slow to fast model populations each generally match the best human ITDENV sensitivity at a progressively higher modulation rate, by membrane-filtering and spike-generation properties producing realistically less than Poisson variance. Non-resonant model EI neurons are also sensitive to interaural intensity differences. With peripheral filters centered between carrier frequency and modulation sideband, fast resonant model EI neurons extend ITDENV sensitivity above 500-Hz modulation.
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Project description:Sensitivity to interaural time differences (ITDs) conveyed in the temporal fine structure of low-frequency tones and the modulated envelopes of high-frequency sounds are considered comparable, particularly for envelopes shaped to transmit similar fidelity of temporal information normally present for low-frequency sounds. Nevertheless, discrimination performance for envelope modulation rates above a few hundred Hertz is reported to be poor-to the point of discrimination thresholds being unattainable-compared with the much higher (>1,000 Hz) limit for low-frequency ITD sensitivity, suggesting the presence of a low-pass filter in the envelope domain. Further, performance for identical modulation rates appears to decline with increasing carrier frequency, supporting the view that the low-pass characteristics observed for envelope ITD processing is carrier-frequency dependent. Here, we assessed listeners' sensitivity to ITDs conveyed in pure tones and in the modulated envelopes of high-frequency tones. ITD discrimination for the modulated high-frequency tones was measured as a function of both modulation rate and carrier frequency. Some well-trained listeners appear able to discriminate ITDs extremely well, even at modulation rates well beyond 500 Hz, for 4-kHz carriers. For one listener, thresholds were even obtained for a modulation rate of 800 Hz. The highest modulation rate for which thresholds could be obtained declined with increasing carrier frequency for all listeners. At 10 kHz, the highest modulation rate at which thresholds could be obtained was 600 Hz. The upper limit of sensitivity to ITDs conveyed in the envelope of high-frequency modulated sounds appears to be higher than previously considered.
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