Searchlight Classification Informative Region Mixture Model (SCIM): Identification of Cortical Regions Showing Discriminable BOLD Patterns in Event-Related Auditory fMRI Data.

The investigation of abstract cognitive tasks, e.g., semantic processing of speech, requires the simultaneous use of a carefully selected stimulus design and sensitive tools for the analysis of corresponding neural activity that are comparable across different studies investigating similar research questions. Multi-voxel pattern analysis (MVPA) methods are commonly used in neuroimaging to investigate BOLD responses corresponding to neural activation associated with specific cognitive tasks. Regions of significant activation are identified by a thresholding operation during multivariate pattern analysis, the results of which are susceptible to the applied threshold value. Investigation of analysis approaches that are robust to a large extent with respect to thresholding, is thus an important goal pursued here. The present paper contributes a novel statistical analysis method for fMRI experiments, searchlight classification informative region mixture model (SCIM), that is based on the assumption that the whole brain volume can be subdivided into two groups of voxels: spatial voxel positions around which recorded BOLD activity does convey information about the present stimulus condition and those that do not. A generative statistical model is proposed that assigns a probability of being informative to each position in the brain, based on a combination of a support vector machine searchlight analysis and Gaussian mixture models. Results from an auditory fMRI study investigating cortical regions that are engaged in the semantic processing of speech indicate that the SCIM method identifies physiologically plausible brain regions as informative, similar to those from two standard methods as reference that we compare to, with two important differences. SCIM-identified regions are very robust to the choice of the threshold for significance, i.e., less "noisy," in contrast to, e.g., the binomial test whose results in the present experiment are highly dependent on the chosen significance threshold or random permutation tests that are additionally bound to very high computational costs. In group analyses, the SCIM method identifies a physiologically plausible pre-frontal region, anterior cingulate sulcus, to be involved in semantic processing that other methods succeed to identify only in single subject analyses.

Matching Pursuit Analysis of Auditory Receptive Fields' Spectro-Temporal Properties.

Gabor filters have long been proposed as models for spectro-temporal receptive fields (STRFs), with their specific spectral and temporal rate of modulation qualitatively replicating characteristics of STRF filters estimated from responses to auditory stimuli in physiological data. The present study builds on the Gabor-STRF model by proposing a methodology to quantitatively decompose STRFs into a set of optimally matched Gabor filters through matching pursuit, and by quantitatively evaluating spectral and temporal characteristics of STRFs in terms of the derived optimal Gabor-parameters. To summarize a neuron's spectro-temporal characteristics, we introduce a measure for the "diagonality," i.e., the extent to which an STRF exhibits spectro-temporal transients which cannot be factorized into a product of a spectral and a temporal modulation. With this methodology, it is shown that approximately half of 52 analyzed zebra finch STRFs can each be well approximated by a single Gabor or a linear combination of two Gabor filters. Moreover, the dominant Gabor functions tend to be oriented either in the spectral or in the temporal direction, with truly "diagonal" Gabor functions rarely being necessary for reconstruction of an STRF's main characteristics. As a toy example for the applicability of STRF and Gabor-STRF filters to auditory detection tasks, we use STRF filters as features in an automatic event detection task and compare them to idealized Gabor filters and mel-frequency cepstral coefficients (MFCCs). STRFs classify a set of six everyday sounds with an accuracy similar to reference Gabor features (94% recognition rate). Spectro-temporal STRF and Gabor features outperform reference spectral MFCCs in quiet and in low noise conditions (down to 0 dB signal to noise ratio).

Statistical identification of stimulus-activated network nodes in multi-neuron voltage-sensitive dye optical recordings.

Voltage-Sensitive Dye (VSD) imaging is an optical imaging method that allows measuring the graded voltage changes of multiple neurons simultaneously. In neuroscience, this method is used to reveal networks of neurons involved in certain tasks. However, the recorded relative dye fluorescence changes are usually low and signals are superimposed by noise and artifacts. Therefore, establishing a reliable method to identify which cells are activated by specific stimulus conditions is the first step to identify functional networks. In this paper, we present a statistical method to identify stimulus-activated network nodes as cells, whose activities during sensory network stimulation differ significantly from the un-stimulated control condition. This method is demonstrated based on voltage-sensitive dye recordings from up to 100 neurons in a ganglion of the medicinal leech responding to tactile skin stimulation. Without relying on any prior physiological knowledge, the network nodes identified by our statistical analysis were found to match well with published cell types involved in tactile stimulus processing and to be consistent across stimulus conditions and preparations.

Robust auditory localization using probabilistic inference and coherence-based weighting of interaural cues.

Robust sound source localization is performed by the human auditory system even in challenging acoustic conditions and in previously unencountered, complex scenarios. Here a computational binaural localization model is proposed that possesses mechanisms for handling of corrupted or unreliable localization cues and generalization across different acoustic situations. Central to the model is the use of interaural coherence, measured as interaural vector strength (IVS), to dynamically weight the importance of observed interaural phase (IPD) and level (ILD) differences in frequency bands up to 1.4 kHz. This is accomplished through formulation of a probabilistic model in which the ILD and IPD distributions pertaining to a specific source location are dependent on observed interaural coherence. Bayesian computation of the direction-of-arrival probability map naturally leads to coherence-weighted integration of location cues across frequency and time. Results confirm the model's validity through statistical analyses of interaural parameter values. Simulated localization experiments show that even data points with low reliability (i.e., low IVS) can be exploited to enhance localization performance. A temporal integration length of at least 200 ms is required to gain a benefit; this is in accordance with previous psychoacoustic findings on temporal integration of spatial cues in the human auditory system.

Fast and robust estimation of spectro-temporal receptive fields using stochastic approximations.

BACKGROUND: The receptive field (RF) represents the signal preferences of sensory neurons and is the primary analysis method for understanding sensory coding. While it is essential to estimate a neuron's RF, finding numerical solutions to increasingly complex RF models can become computationally intensive, in particular for high-dimensional stimuli or when many neurons are involved. NEW METHOD: Here we propose an optimization scheme based on stochastic approximations that facilitate this task. The basic idea is to derive solutions on a random subset rather than computing the full solution on the available data set. To test this, we applied different optimization schemes based on stochastic gradient descent (SGD) to both the generalized linear model (GLM) and a recently developed classification-based RF estimation approach. RESULTS AND COMPARISON WITH EXISTING METHOD: Using simulated and recorded responses, we demonstrate that RF parameter optimization based on state-of-the-art SGD algorithms produces robust estimates of the spectro-temporal receptive field (STRF). Results on recordings from the auditory midbrain demonstrate that stochastic approximations preserve both predictive power and tuning properties of STRFs. A correlation of 0.93 with the STRF derived from the full solution may be obtained in less than 10% of the full solution's estimation time. We also present an on-line algorithm that allows simultaneous monitoring of STRF properties of more than 30 neurons on a single computer. CONCLUSIONS: The proposed approach may not only prove helpful for large-scale recordings but also provides a more comprehensive characterization of neural tuning in experiments than standard tuning curves.

Discriminative learning of receptive fields from responses to non-Gaussian stimulus ensembles.

Analysis of sensory neurons' processing characteristics requires simultaneous measurement of presented stimuli and concurrent spike responses. The functional transformation from high-dimensional stimulus space to the binary space of spike and non-spike responses is commonly described with linear-nonlinear models, whose linear filter component describes the neuron's receptive field. From a machine learning perspective, this corresponds to the binary classification problem of discriminating spike-eliciting from non-spike-eliciting stimulus examples. The classification-based receptive field (CbRF) estimation method proposed here adapts a linear large-margin classifier to optimally predict experimental stimulus-response data and subsequently interprets learned classifier weights as the neuron's receptive field filter. Computational learning theory provides a theoretical framework for learning from data and guarantees optimality in the sense that the risk of erroneously assigning a spike-eliciting stimulus example to the non-spike class (and vice versa) is minimized. Efficacy of the CbRF method is validated with simulations and for auditory spectro-temporal receptive field (STRF) estimation from experimental recordings in the auditory midbrain of Mongolian gerbils. Acoustic stimulation is performed with frequency-modulated tone complexes that mimic properties of natural stimuli, specifically non-Gaussian amplitude distribution and higher-order correlations. Results demonstrate that the proposed approach successfully identifies correct underlying STRFs, even in cases where second-order methods based on the spike-triggered average (STA) do not. Applied to small data samples, the method is shown to converge on smaller amounts of experimental recordings and with lower estimation variance than the generalized linear model and recent information theoretic methods. Thus, CbRF estimation may prove useful for investigation of neuronal processes in response to natural stimuli and in settings where rapid adaptation is induced by experimental design.

Temporal variability of spectro-temporal receptive fields in the anesthetized auditory cortex.

Temporal variability of neuronal response characteristics during sensory stimulation is a ubiquitous phenomenon that may reflect processes such as stimulus-driven adaptation, top-down modulation or spontaneous fluctuations. It poses a challenge to functional characterization methods such as the receptive field, since these often assume stationarity. We propose a novel method for estimation of sensory neurons' receptive fields that extends the classic static linear receptive field model to the time-varying case. Here, the long-term estimate of the static receptive field serves as the mean of a probabilistic prior distribution from which the short-term temporally localized receptive field may deviate stochastically with time-varying standard deviation. The derived corresponding generalized linear model permits robust characterization of temporal variability in receptive field structure also for highly non-Gaussian stimulus ensembles. We computed and analyzed short-term auditory spectro-temporal receptive field (STRF) estimates with characteristic temporal resolution 5-30 s based on model simulations and responses from in total 60 single-unit recordings in anesthetized Mongolian gerbil auditory midbrain and cortex. Stimulation was performed with short (100 ms) overlapping frequency-modulated tones. Results demonstrate identification of time-varying STRFs, with obtained predictive model likelihoods exceeding those from baseline static STRF estimation. Quantitative characterization of STRF variability reveals a higher degree thereof in auditory cortex compared to midbrain. Cluster analysis indicates that significant deviations from the long-term static STRF are brief, but reliably estimated. We hypothesize that the observed variability more likely reflects spontaneous or state-dependent internal fluctuations that interact with stimulus-induced processing, rather than experimental or stimulus design.

Beyond novelty detection: incongruent events, when general and specific classifiers disagree.

Unexpected stimuli are a challenge to any machine learning algorithm. Here, we identify distinct types of unexpected events when general-level and specific-level classifiers give conflicting predictions. We define a formal framework for the representation and processing of incongruent events: Starting from the notion of label hierarchy, we show how partial order on labels can be deduced from such hierarchies. For each event, we compute its probability in different ways, based on adjacent levels in the label hierarchy. An incongruent event is an event where the probability computed based on some more specific level is much smaller than the probability computed based on some more general level, leading to conflicting predictions. Algorithms are derived to detect incongruent events from different types of hierarchies, different applications, and a variety of data types. We present promising results for the detection of novel visual and audio objects, and new patterns of motion in video. We also discuss the detection of Out-Of- Vocabulary words in speech recognition, and the detection of incongruent events in a multimodal audiovisual scenario.

Spatio-temporal dynamics in fMRI recordings revealed with complex independent component analysis.

Independent component analysis (ICA) of functional magnetic resonance imaging (fMRI) data is commonly carried out under the assumption that each source may be represented as a spatially fixed pattern of activation, which leads to the instantaneous mixing model. To allow modeling patterns of spatio-temporal dynamics, in particular, the flow of oxygenated blood, we have developed a convolutive ICA approach: spatial complex ICA applied to frequency-domain fMRI data. In several frequency-bands, we identify components pertaining to activity in primary visual cortex (V1) and blood supply vessels. One such component, obtained in the 0.10 Hz band, is analyzed in detail and found to likely reflect flow of oxygenated blood in V1.

Complex independent component analysis of frequency-domain electroencephalographic data.

Independent component analysis (ICA) has proven useful for modeling brain and electroencephalographic (EEG) data. Here, we present a new, generalized method to better capture the dynamics of brain signals than previous ICA algorithms. We regard EEG sources as eliciting spatio-temporal activity patterns, corresponding to, e.g. trajectories of activation propagating across cortex. This leads to a model of convolutive signal superposition, in contrast with the commonly used instantaneous mixing model. In the frequency-domain, convolutive mixing is equivalent to multiplicative mixing of complex signal sources within distinct spectral bands. We decompose the recorded spectral-domain signals into independent components by a complex infomax ICA algorithm. First results from a visual attention EEG experiment exhibit: (1). sources of spatio-temporal dynamics in the data, (2). links to subject behavior, (3). sources with a limited spectral extent, and (4). a higher degree of independence compared to sources derived by standard ICA.