Interaural Level Difference-Dependent Gain Control and Synaptic Scaling Underlying Binaural Computation

SummaryBinaural 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

Noninvasive fMRI investigation of interaural level difference processing the rat auditory subcortex

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Interaural Level Difference Optimization of Binaural Ambisonic Rendering

Ambisonics is a spatial audio technique appropriate for dynamic binaural rendering due to its sound field rotation and transformation capabilities, which has made it popular for virtual reality applications. An issue with low-order Ambisonics is that interaural level differences (ILDs) are often reproduced with lower values when compared to head-related impulse responses (HRIRs), which reduces lateralization and spaciousness. This paper introduces a method of Ambisonic ILD Optimization (AIO), a pre-processing technique to bring the ILDs produced by virtual loudspeaker binaural Ambisonic rendering closer to those of HRIRs. AIO is evaluated objectively for Ambisonic orders up to fifth order versus a reference dataset of HRIRs for all locations on the sphere via estimated ILD and spectral difference, and perceptually through listening tests using both simple and complex scenes. Results conclude AIO produces an overall improvement for all tested orders of Ambisonics, though the benefits are greatest at first and second order

High Frequency Reproduction in Binaural Ambisonic Rendering

Humans can localise sounds in all directions using three main auditory cues: the differences in time and level between signals arriving at the left and right eardrums (interaural time difference and interaural level difference, respectively), and the spectral characteristics of the signals due to reflections and diffractions off the body and ears. These auditory cues can be recorded for a position in space using the head-related transfer function (HRTF), and binaural synthesis at this position can then be achieved through convolution of a sound signal with the measured HRTF. However, reproducing soundfields with multiple sources, or at multiple locations, requires a highly dense set of HRTFs. Ambisonics is a spatial audio technology that decomposes a soundfield into a weighted set of directional functions, which can be utilised binaurally in order to spatialise audio at any direction using far fewer HRTFs. A limitation of low-order Ambisonic rendering is poor high frequency reproduction, which reduces the accuracy of the resulting binaural synthesis. This thesis presents novel HRTF pre-processing techniques, such that when using the augmented HRTFs in the binaural Ambisonic rendering stage, the high frequency reproduction is a closer approximation of direct HRTF rendering. These techniques include Ambisonic Diffuse-Field Equalisation, to improve spectral reproduction over all directions; Ambisonic Directional Bias Equalisation, to further improve spectral reproduction toward a specific direction; and Ambisonic Interaural Level Difference Optimisation, to improve lateralisation and interaural level difference reproduction. Evaluation of the presented techniques compares binaural Ambisonic rendering to direct HRTF rendering numerically, using perceptually motivated spectral difference calculations, auditory cue estimations and localisation prediction models, and perceptually, using listening tests assessing similarity and plausibility. Results conclude that the individual pre-processing techniques produce modest improvements to the high frequency reproduction of binaural Ambisonic rendering, and that using multiple pre-processing techniques can produce cumulative, and statistically significant, improvements

Virtual Reality and Sound Localization

Psychoacoustics is the scientific study of sound perception. Within this field, Virtual Reality is a technique that uses two synthesis speakers to simulate a sine tone coming from anywhere in open space. Using this method it is possible to independently control specific binaural cues in a free-field environment. This study analyzes listener responses to these controlled sine tones to investigate the relative importance of certain binaural cues at different frequencies

Multiplicative Auditory Spatial Receptive Fields Created by a Hierarchy of Population Codes

A multiplicative combination of tuning to interaural time difference (ITD) and interaural level difference (ILD) contributes to the generation of spatially selective auditory neurons in the owl's midbrain. Previous analyses of multiplicative responses in the owl have not taken into consideration the frequency-dependence of ITD and ILD cues that occur under natural listening conditions. Here, we present a model for the responses of ITD- and ILD-sensitive neurons in the barn owl's inferior colliculus which satisfies constraints raised by experimental data on frequency convergence, multiplicative interaction of ITD and ILD, and response properties of afferent neurons. We propose that multiplication between ITD- and ILD-dependent signals occurs only within frequency channels and that frequency integration occurs using a linear-threshold mechanism. The model reproduces the experimentally observed nonlinear responses to ITD and ILD in the inferior colliculus, with greater accuracy than previous models. We show that linear-threshold frequency integration allows the system to represent multiple sound sources with natural sound localization cues, whereas multiplicative frequency integration does not. Nonlinear responses in the owl's inferior colliculus can thus be generated using a combination of cellular and network mechanisms, showing that multiple elements of previous theories can be combined in a single system

Coding of auditory space

Behavioral, anatomical, and physiological approaches can be integrated in the study of sound localization in barn owls. Space representation in owls provides a useful example for discussion of place and ensemble coding. Selectivity for space is broad and ambiguous in low-order neurons. Parallel pathways for binaural cues and for different frequency bands converge on high-order space-specific neurons, which encode space more precisely. An ensemble of broadly tuned place-coding neurons may converge on a single high-order neuron to create an improved labeled line. Thus, the two coding schemes are not alternate methods. Owls can localize sounds by using either the isomorphic map of auditory space in the midbrain or forebrain neural networks in which space is not mapped

A theoretical analysis of sound localisation, with application to amplitude panning

Below 700Hz sound fields can be approximated well over a region of space that encloses the human head, using the acoustic pressure and gradient. With this representation convenient expressions are found for the resulting Interaural Time Difference (ITD) and Interaural Level Difference (ILD). This formulation facilitates the investigation of various head-related phenomena of natural and synthesised fields. As an example, perceived image direction is related to head direction and the sound field description. This result is then applied to a general amplitude panning system, and can be used to create images that are stable with respect to head direction