Neural Models of Subcortical Auditory Processing

An important feature of the auditory system is its ability to distinguish many simultaneous sound sources. The primary goal of this work was to understand how a robust, preattentive analysis of the auditory scene is accomplished by the subcortical auditory system. Reasonably accurate modelling of the morphology and organisation of the relevant auditory nuclei, was seen as being of great importance. The formulation of plausible models and their subsequent simulation was found to be invaluable in elucidating biological processes and in highlighting areas of uncertainty. In the thesis, a review of important aspects of mammalian auditory processing is presented and used as a basis for the subsequent modelling work. For each aspect of auditory processing modelled, psychophysical results are described and existing models reviewed, before the models used here are described and simulated. Auditory processes which are modelled include the peripheral system, and the production of tonotopic maps of the spectral content of complex acoustic stimuli, and of modulation frequency or periodicity. A model of the formation of sequential associations between successive sounds is described, and the model is shown to be capable of emulating a wide range of psychophysical behaviour. The grouping of related spectral components and the development of pitch perception is also investigated. Finally a critical assessment of the work and ideas for future developments are presented. The principal contributions of this work are the further development of a model for pitch perception and the development of a novel architecture for the sequential association of those groups. In the process of developing these ideas, further insights into subcortical auditory processing were gained, and explanations for a number of puzzling psychophysical characteristics suggested.Royal Naval Engineering College, Manadon, Plymout

Adaptation of Binaural Processing in the Adult Brainstem Induced by Ambient Noise

Interaural differences in stimulus intensity and timing are major cues for sound localization. In mammals, these cues are first processed in the lateral and medial superior olive by interaction of excitatory and inhibitory synaptic inputs from ipsi- and contralateral cochlear nucleus neurons. To preserve sound localization acuity following changes in the acoustic environment, the processing of these binaural cues needs neuronal adaptation. Recent studies have shown that binaural sensitivity adapts to stimulation history within milliseconds, but the actual extent of binaural adaptation is unknown. In the current study, we investigated long-term effects on binaural sensitivity using extracellular in vivo recordings from single neurons in the dorsal nucleus of the lateral lemniscus that inherit their binaural properties directly from the lateral and medial superior olives. In contrast to most previous studies, we used a noninvasive approach to influence this processing. Adult gerbils were exposed for 2 weeks to moderate noise with no stable binaural cue. We found monaural response properties to be unaffected by this measure. However, neuronal sensitivity to binaural cues was reversibly altered for a few days. Computational models of sensitivity to interaural time and level differences suggest that upregulation of inhibition in the superior olivary complex can explain the electrophysiological data

The role of sound offsets in auditory temporal processing and perception

Sound-offset responses are distinct to sound onsets in their underlying neural mechanisms, temporal processing pathways and roles in auditory perception following recent neurobiological studies. In this work, I investigate the role of sound offsets and the effect of reduced sensitivity to offsets on auditory perception in humans. The implications of a 'sound-offset deficit' for speech-in-noise perception are investigated, based on a mathematical model with biological significance and independent channels for onset and offset detection. Sound offsets are important in recognising, distinguishing and grouping sounds. They are also likely to play a role in perceiving consonants that lie in the troughs of amplitude fluctuations in speech. The offset influence on the discriminability of model outputs for 48 non-sense vowel-consonant-vowel (VCV) speech stimuli in varying levels of multi-talker babble noise (-12, -6, 0, 6, 12 dB SNR) was assessed, and led to predictions that correspond to known phonetic categories. This work therefore suggests that variability in the offset salience alone can explain the rank order of consonants most affected in noisy situations. A novel psychophysical test battery for offset sensitivity was devised and assessed, followed by a study to find an electrophysiological correlate. The findings suggest that individual differences in sound-offset sensitivity may be a factor contributing to inter-subject variation in speech-in-noise discrimination ability. The promising measures from these results can be used to test between-population differences in offset sensitivity, with more support for objective than psychophysical measures. In the electrophysiological study, offset responses in a duration discrimination paradigm were found to be modulated by attention compared to onset responses. Overall, this thesis shows for the first time that the onset-offset dichotomy in the auditory system, previously explored in physiological studies, is also evident in human studies for both simple and complex speech sounds

Dual Coding of Frequency Modulation in the Ventral Cochlear Nucleus.

Frequency modulation (FM) is a common acoustic feature of natural sounds and is known to play a role in robust sound source recognition. Auditory neurons show precise stimulus-synchronized discharge patterns that may be used for the representation of low-rate FM. However, it remains unclear whether this representation is based on synchronization to slow temporal envelope (ENV) cues resulting from cochlear filtering or phase locking to faster temporal fine structure (TFS) cues. To investigate the plausibility of those encoding schemes, single units of the ventral cochlear nucleus of guinea pigs of either sex were recorded in response to sine FM tones centered at the unit's best frequency (BF). The results show that, in contrast to high-BF units, for modulation depths within the receptive field, low-BF units (<4 kHz) demonstrate good phase locking to TFS. For modulation depths extending beyond the receptive field, the discharge patterns follow the ENV and fluctuate at the modulation rate. The receptive field proved to be a good predictor of the ENV responses for most primary-like and chopper units. The current in vivo data also reveal a high level of diversity in responses across unit types. TFS cues are mainly conveyed by low-frequency and primary-like units and ENV cues by chopper and onset units. The diversity of responses exhibited by cochlear nucleus neurons provides a neural basis for a dual-coding scheme of FM in the brainstem based on both ENV and TFS cues.SIGNIFICANCE STATEMENT Natural sounds, including speech, convey informative temporal modulations in frequency. Understanding how the auditory system represents those frequency modulations (FM) has important implications as robust sound source recognition depends crucially on the reception of low-rate FM cues. Here, we recorded 115 single-unit responses from the ventral cochlear nucleus in response to FM and provide the first physiological evidence of a dual-coding mechanism of FM via synchronization to temporal envelope cues and phase locking to temporal fine structure cues. We also demonstrate a diversity of neural responses with different coding specializations. These results support the dual-coding scheme proposed by psychophysicists to account for FM sensitivity in humans and provide new insights on how this might be implemented in the early stages of the auditory pathway

Relationships between human auditory cortical structure and function

The human auditory cortex comprises multiple areas, largely distributed across the supratemporal plane, but the precise number and configuration of auditory areas and their functional significance have not yet been clearly established. In this paper, we discuss recent research concerning architectonic and functional organisation within the human auditory cortex, as well as architectonic and neurophysiological studies in non-human species, which can provide a broad conceptual framework for interpreting functional specialisation in humans. We review the pattern in human auditory cortex of the functional responses to various acoustic cues, such as frequency, pitch, sound level, temporal variation, motion and spatial location, and we discuss their correspondence to what is known about the organisation of the auditory cortex in other primates. There is some neuroimaging evidence of multiple tonotopically organised fields in humans and of functional specialisations of the fields in the processing of different sound features. It is thought that the primary area, on Heschl's gyrus, may have a larger involvement in processing basic sound features, such as frequency and level, and that posterior non-primary areas on the planum temporale may play a larger role in processing more spectrotemporally complex sounds. Ways in which current knowledge of auditory cortical organisation and different data analysis approaches may benefit future functional neuroimaging studies which seek to link auditory cortical structure and function are discussed

Effects of Hearing Aid Amplification on Robust Neural Coding of Speech

Hearing aids are able to restore some hearing abilities for people with auditory impairments, but background noise remains a significant problem. Unfortunately, we know very little about how speech is encoded in the auditory system, particularly in impaired systems with prosthetic amplifiers. There is growing evidence that relative timing in the neural signals (known as spatiotemporal coding) is important for speech perception, but there is little research that relates spatiotemporal coding and hearing aid amplification. This research uses a combination of computational modeling and physiological experiments to characterize how hearing aids affect vowel coding in noise at the level of the auditory nerve. The results indicate that sensorineural hearing impairment degrades the temporal cues transmitted from the ear to the brain. Two hearing aid strategies (linear gain and wide dynamic-range compression) were used to amplify the acoustic signal. Although appropriate gain was shown to improve temporal coding for individual auditory nerve fibers, neither strategy improved spatiotemporal cues. Previous work has attempted to correct the relative timing by adding frequency-dependent delays to the acoustic signal (e.g., within a hearing aid). We show that, although this strategy can affect the timing of auditory nerve responses, it is unlikely to improve the relative timing as intended. We have shown that existing hearing aid technologies do not improve some of the neural cues that we think are important for perception, but it is important to understand these limitations. Our hope is that this knowledge can be used to develop new technologies to improve auditory perception in difficult acoustic environments

A comparative study of eight human auditory models of monaural processing

A number of auditory models have been developed using diverging approaches, either physiological or perceptual, but they share comparable stages of signal processing, as they are inspired by the same constitutive parts of the auditory system. We compare eight monaural models that are openly accessible in the Auditory Modelling Toolbox. We discuss the considerations required to make the model outputs comparable to each other, as well as the results for the following model processing stages or their equivalents: Outer and middle ear, cochlear filter bank, inner hair cell, auditory nerve synapse, cochlear nucleus, and inferior colliculus. The discussion includes a list of recommendations for future applications of auditory models.Comment: Revision 1 of the manuscrip