Sparse Codes for Speech Predict Spectrotemporal Receptive Fields in the Inferior Colliculus

We have developed a sparse mathematical representation of speech that minimizes the number of active model neurons needed to represent typical speech sounds. The model learns several well-known acoustic features of speech such as harmonic stacks, formants, onsets and terminations, but we also find more exotic structures in the spectrogram representation of sound such as localized checkerboard patterns and frequency-modulated excitatory subregions flanked by suppressive sidebands. Moreover, several of these novel features resemble neuronal receptive fields reported in the Inferior Colliculus (IC), as well as auditory thalamus and cortex, and our model neurons exhibit the same tradeoff in spectrotemporal resolution as has been observed in IC. To our knowledge, this is the first demonstration that receptive fields of neurons in the ascending mammalian auditory pathway beyond the auditory nerve can be predicted based on coding principles and the statistical properties of recorded sounds.Comment: For Supporting Information, see PLoS website: http://www.ploscompbiol.org/article/info%3Adoi%2F10.1371%2Fjournal.pcbi.100259

Efficient coding of spectrotemporal binaural sounds leads to emergence of the auditory space representation

To date a number of studies have shown that receptive field shapes of early sensory neurons can be reproduced by optimizing coding efficiency of natural stimulus ensembles. A still unresolved question is whether the efficient coding hypothesis explains formation of neurons which explicitly represent environmental features of different functional importance. This paper proposes that the spatial selectivity of higher auditory neurons emerges as a direct consequence of learning efficient codes for natural binaural sounds. Firstly, it is demonstrated that a linear efficient coding transform - Independent Component Analysis (ICA) trained on spectrograms of naturalistic simulated binaural sounds extracts spatial information present in the signal. A simple hierarchical ICA extension allowing for decoding of sound position is proposed. Furthermore, it is shown that units revealing spatial selectivity can be learned from a binaural recording of a natural auditory scene. In both cases a relatively small subpopulation of learned spectrogram features suffices to perform accurate sound localization. Representation of the auditory space is therefore learned in a purely unsupervised way by maximizing the coding efficiency and without any task-specific constraints. This results imply that efficient coding is a useful strategy for learning structures which allow for making behaviorally vital inferences about the environment.Comment: 22 pages, 9 figure

Learning Mid-Level Auditory Codes from Natural Sound Statistics

Interaction with the world requires an organism to transform sensory signals into representations in which behaviorally meaningful properties of the environment are made explicit. These representations are derived through cascades of neuronal processing stages in which neurons at each stage recode the output of preceding stages. Explanations of sensory coding may thus involve understanding how low-level patterns are combined into more complex structures. Although models exist in the visual domain to explain how mid-level features such as junctions and curves might be derived from oriented filters in early visual cortex, little is known about analogous grouping principles for mid-level auditory representations. We propose a hierarchical generative model of natural sounds that learns combina- tions of spectrotemporal features from natural stimulus statistics. In the first layer the model forms a sparse convolutional code of spectrograms using a dictionary of learned spectrotemporal kernels. To generalize from specific kernel activation patterns, the second layer encodes patterns of time-varying magnitude of multiple first layer coefficients. Because second-layer features are sensitive to combi- nations of spectrotemporal features, the representation they support encodes more complex acoustic patterns than the first layer. When trained on corpora of speech and environmental sounds, some second-layer units learned to group spectrotemporal features that occur together in natural sounds. Others instantiate opponency between dissimilar sets of spectrotemporal features. Such groupings might be instantiated by neurons in the auditory cortex, providing a hypothesis for mid-level neuronal computation.This work was supported by the Center for Brains, Minds and Machines (CBMM), funded by NSF STC award CCF-1231216

An information theoretic characterisation of auditory encoding.

The entropy metric derived from information theory provides a means to quantify the amount of information transmitted in acoustic streams like speech or music. By systematically varying the entropy of pitch sequences, we sought brain areas where neural activity and energetic demands increase as a function of entropy. Such a relationship is predicted to occur in an efficient encoding mechanism that uses less computational resource when less information is present in the signal: we specifically tested the hypothesis that such a relationship is present in the planum temporale (PT). In two convergent functional MRI studies, we demonstrated this relationship in PT for encoding, while furthermore showing that a distributed fronto-parietal network for retrieval of acoustic information is independent of entropy. The results establish PT as an efficient neural engine that demands less computational resource to encode redundant signals than those with high information content

The Opponent Channel Population Code of Sound Location Is an Efficient Representation of Natural Binaural Sounds

In mammalian auditory cortex, sound source position is represented by a population of broadly tuned neurons whose firing is modulated by sounds located at all positions surrounding the animal. Peaks of their tuning curves are concentrated at lateral position, while their slopes are steepest at the interaural midline, allowing for the maximum localization accuracy in that area. These experimental observations contradict initial assumptions that the auditory space is represented as a topographic cortical map. It has been suggested that a “panoramic” code has evolved to match specific demands of the sound localization task. This work provides evidence suggesting that properties of spatial auditory neurons identified experimentally follow from a general design principle- learning a sparse, efficient representation of natural stimuli. Natural binaural sounds were recorded and served as input to a hierarchical sparse-coding model. In the first layer, left and right ear sounds were separately encoded by a population of complex-valued basis functions which separated phase and amplitude. Both parameters are known to carry information relevant for spatial hearing. Monaural input converged in the second layer, which learned a joint representation of amplitude and interaural phase difference. Spatial selectivity of each second-layer unit was measured by exposing the model to natural sound sources recorded at different positions. Obtained tuning curves match well tuning characteristics of neurons in the mammalian auditory cortex. This study connects neuronal coding of the auditory space with natural stimulus statistics and generates new experimental predictions. Moreover, results presented here suggest that cortical regions with seemingly different functions may implement the same computational strategy-efficient coding.German Science Foundation (Graduate College "InterNeuro"

Functional Sensory Representations of Natural Stimuli: the Case of Spatial Hearing

In this thesis I attempt to explain mechanisms of neuronal coding in the auditory system as a form of adaptation to statistics of natural stereo sounds. To this end I analyse recordings of real-world auditory environments and construct novel statistical models of these data. I further compare regularities present in natural stimuli with known, experimentally observed neuronal mechanisms of spatial hearing. In a more general perspective, I use binaural auditory system as a starting point to consider the notion of function implemented by sensory neurons. In particular I argue for two, closely-related tenets: 1. The function of sensory neurons can not be fully elucidated without understanding statistics of natural stimuli they process. 2. Function of sensory representations is determined by redundancies present in the natural sensory environment. I present the evidence in support of the first tenet by describing and analysing marginal statistics of natural binaural sound. I compare observed, empirical distributions with knowledge from reductionist experiments. Such comparison allows to argue that the complexity of the spatial hearing task in the natural environment is much higher than analytic, physics-based predictions. I discuss the possibility that early brain stem circuits such as LSO and MSO do not \"compute sound localization\" as is often being claimed in the experimental literature. I propose that instead they perform a signal transformation, which constitutes the first step of a complex inference process. To support the second tenet I develop a hierarchical statistical model, which learns a joint sparse representation of amplitude and phase information from natural stereo sounds. I demonstrate that learned higher order features reproduce properties of auditory cortical neurons, when probed with spatial sounds. Reproduced aspects were hypothesized to be a manifestation of a fine-tuned computation specific to the sound-localization task. Here it is demonstrated that they rather reflect redundancies present in the natural stimulus. Taken together, results presented in this thesis suggest that efficient coding is a strategy useful for discovering structures (redundancies) in the input data. Their meaning has to be determined by the organism via environmental feedback