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O'Reilly, J. A.

Publications and source records attributed to O'Reilly, J. A..

9 recordsLinked to original sources

Blind source separation of event-related potentials using a recurrent neural network

Event-related potentials (ERPs) are a superposition of electric potential differences generated by neurophysiological activity associated with psychophysical events. Spatiotemporal dissociation of these signal sources can supplement conventional ERP analysis and improve source localization. However, results from established source separation methods applied to ERPs can be challenging to interpret. Hence, we have developed a recurrent neural network (RNN) method for blind source separation. The RNN transforms input step pulse signals representing events into corresponding ERP difference waveforms. Source waveforms are obtained from penultimate layer units and scalp maps are obtained from feed-forward output layer weights that project these source waveforms onto EEG electrode amplitudes. An interpretable, sparse source representation is achieved by incorporating L1 regularization of signals obtained from the penultimate layer of the network during training. This RNN method was applied to four ERP difference waveforms (MMN, N170, N400, P3) from the open-access ERP CORE database, and independent component analysis (ICA) was applied to the same data for comparison. The RNN decomposed these ERPs into eleven spatially and temporally separate sources that were less noisy, tended to be more ERP-specific, and were less similar to each other than ICA-derived sources. The RNN sources also had less ambiguity between source waveform amplitude, scalp potential polarity, and equivalent current dipole orientation than ICA sources. In conclusion, the proposed RNN blind source separation method can be effectively applied to grand-average ERP difference waves and holds promise for further development as a computational model of event-related neural signals.

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Localized estimation of event-related neural source activity from simultaneous MEG-EEG with a recurrent neural network

Estimating intracranial current sources underlying the electromagnetic signals observed from extracranial sensors is a perennial challenge in non-invasive neuroimaging. Established solutions to this inverse problem treat time samples independently without considering the temporal dynamics of event-related brain processes. This paper describes current source estimation from simultaneously recorded magneto- and electro-encephalography (MEEG) using a recurrent neural network (RNN) that learns sequential relationships from neural data. The RNN was trained in two phases: (1) pre-training and (2) transfer learning with L1 regularization applied to the source estimation layer. Performance of using scaled labels derived from MEEG, magnetoencephalography (MEG), or electroencephalography (EEG) were compared, as were results from volumetric source space with free dipole orientation and surface source space with fixed dipole orientation. Exact low-resolution electromagnetic tomography (eLORETA) and mixed-norm L1/L2 (MxNE) source estimation methods were also applied to these data for comparison with the RNN method. The RNN approach outperformed other methods in terms of output signal-to-noise ratio, correlation and mean-squared error metrics evaluated against ground-truth event-related field (ERF) and event-related potential (ERP) waveforms. Using MEEG labels with fixed-orientation surface sources produced the most consistent estimates. To estimate sources of ERF and ERP waveforms, the RNN generates temporal dynamics within its internal computational units, driven by sequential structure in neural data used as training labels. It thus provides a data-driven model of computational transformations from psychophysiological events into corresponding event-related neural signals, which is unique among MEG and EEG source reconstruction solutions.

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Localized estimation of electromagnetic sources underlying event-related fields using recurrent neural networks

ObjectiveTo use a recurrent neural network (RNN) to reconstruct neural activity responsible for generating noninvasively measured electromagnetic signals. ApproachOutput weights of an RNN were fixed as the lead field matrix from volumetric source space computed using the boundary element method with co-registered structural magnetic resonance images and magnetoencephalography (MEG). Initially, the network was trained to minimize mean-squared-error loss between its outputs and MEG signals, causing activations in the penultimate layer to converge towards putative neural source activations. Subsequently, L1 regularization was applied to the final hidden layer, and the model was fine-tuned, causing it to favour more focused activations. Estimated source signals were then obtained from the outputs of the last hidden layer. We developed and validated this approach with simulations before applying it to real MEG data, comparing performance with three existing methods: beamformers, minimum-norm estimate, and dynamical statistical parametric mapping. Main resultsThe proposed method had higher output signal-to-noise ratios than the others and comparable correlation and error between estimated and simulated sources. Reconstructed MEG signals were also equal or superior to the other methods in terms of their similarity to ground-truth. When applied to MEG data recorded during an auditory roving oddball experiment, source signals estimated with the RNN were generally consistent with expectations from the literature and qualitatively smoother and more reasonable-looking than estimates from the other methods. SignificanceThis work builds on recent developments of RNNs for modelling event-related neural responses by incorporating biophysical constraints from the forward model, thus taking a significant step towards greater biological realism and introducing the possibility of exploring how input manipulations may influence localized neural activity.

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Neural correlates of face perception modeled with a convolutional recurrent neural network

Event-related potential (ERP) sensitivity to faces is predominantly characterized by an N170 peak that has greater amplitude and shorter latency when elicited by human faces than images of other objects. We developed a computational model of visual ERP generation to study this phenomenon which consisted of a convolutional neural network (CNN) connected to a recurrent neural network (RNN). We used open-access data to develop the model, generated synthetic images for simulating experiments, then collected additional data to validate predictions of these simulations. For modeling, visual stimuli presented during ERP experiments were represented as sequences of images (time x pixels). These were provided as inputs to the model. The CNN transformed these inputs into sequences of vectors that were passed to the RNN. The ERP waveforms evoked by visual stimuli were provided to the RNN as labels for supervised learning. The whole model was trained end-to-end using data from the open-access dataset to reproduce ERP waveforms evoked by visual events. Cross-validation model outputs strongly correlated with open-access (r = 0.98) and validation study data (r = 0.78). Open-access and validation study data correlated similarly (r = 0.81). Some aspects of model behavior were consistent with neural recordings while others were not, suggesting promising albeit limited capacity for modeling the neurophysiology of face-sensitive ERP generation.

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Decoding violated sensory expectations from the auditory cortex of anaesthetized mice: Hierarchical recurrent neural network depicts separate 'danger' and 'safety' units

The ability to respond appropriately to sensory information received from the external environment is among the most fundamental capabilities of central nervous systems. In the auditory domain, processes underlying this behaviour are studied by measuring auditory-evoked electrophysiology during sequences of sounds with predetermined regularities. Identifying neural correlates of ensuing auditory novelty responses is supported by research in experimental animals. In the present study, we reanalysed epidural field potential recordings from the auditory cortex of anaesthetised mice during frequency and intensity oddball stimulation. Multivariate pattern analysis (MVPA) and hierarchical recurrent neural network (RNN) modelling were adopted to explore these data with greater resolution than previously considered using conventional methods. Time-wise and generalised temporal decoding MVPA approaches revealed previously underestimated asymmetry between responses to sound-level transitions in the intensity oddball paradigm, in contrast with tone frequency changes. After training, the cross-validated RNN model architecture with four hidden layers produced output waveforms in response to simulated auditory inputs that were strongly correlated with grand-average auditory-evoked potential waveforms (r2 > 0.9). Units in hidden layers were classified based on their temporal response properties and characterised using principal component analysis and sample entropy. These demonstrated spontaneous alpha rhythms, sound onset and offset responses, and putative safety and danger units activated by relatively inconspicuous and salient changes in auditory inputs, respectively. The hypothesised existence of corresponding biological neural sources is naturally derived from this model. If proven, this would have significant implications for prevailing theories of auditory processing.

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Recurrent neural network model of human event-related potentials in response to intensity oddball stimulation

The mismatch negativity (MMN) component of the human event-related potential (ERP) is frequently interpreted as a sensory prediction-error signal. However, there is ambiguity concerning the neurophysiology underlying hypothetical prediction and prediction-error signalling components, and whether these can be dissociated from overlapping obligatory components of the ERP that are sensitive to physical properties of sounds. In the present study, a hierarchical recurrent neural network (RNN) was fitted to ERP data from 38 subjects. After training the model to reproduce ERP waveforms evoked by 80 dB standard and 70 dB deviant stimuli, it was used to simulate a response to 90 dB deviant stimuli. Internal states of the RNN effectively combine to generate synthetic ERPs, where individual hidden units are loosely analogous to population-level sources. Model behaviour was characterised using principal component analysis of stimulus condition, layer, and individual unit responses. Hidden units were categorised according to their temporal response fields, and statistically significant differences among stimulus conditions were observed for amplitudes of units peaking in the 0 to 75 ms (P50), 75 to 125 ms (N1), and 250 to 400 ms (N3) latency ranges, surprisingly not including the measurement window of MMN. The model demonstrated opposite polarity changes in MMN amplitude produced by falling (70 dB) and rising (90 dB) intensity deviant stimuli, consistent with loudness dependence of sensory ERP components. Although perhaps less parsimoniously, these observations could be interpreted within the context of predictive coding theory, as examples of negative and positive prediction errors, respectively.

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Modelling mouse auditory response dynamics along a continuum of consciousness using a deep recurrent neural network

ObjectiveUnderstanding neurophysiological changes that accompany transitions between anaesthetized and conscious states is a key objective of anesthesiology and consciousness science. This study aimed to characterize the dynamics of auditory-evoked potential morphology in mice along a continuum of consciousness. ApproachEpidural field potentials were recorded from above the primary auditory cortices of two groups of laboratory mice: urethane-anaesthetized (A, n = 14) and conscious (C, n = 17). Both groups received auditory stimulation in the form of a repeated pure-tone stimulus, before and after receiving 10 mg/kg i.p. ketamine (AK and CK). Evoked responses were then ordered by ascending sample entropy into AK, A, CK, and C, considered to reflect physiological correlates of awareness. These data were used to train a recurrent neural network (RNN) with an input parameter encoding state. Model outputs were compared with grand-average event-related potential (ERP) waveforms. Subsequently, the state parameter was varied to simulate changes in the ERP that occur during transitions between states, and relationships with dominant peak amplitudes were quantified. Main resultsThe RNN synthesized output waveforms that were in close agreement with grand-average ERPs for each group (r2 > 0.9, p < 0.0001). Varying the input state parameter generated model outputs reflecting changes in ERP morphology predicted to occur between states. Positive peak amplitudes within 25 to 50 ms, and negative peak amplitudes within 50 to 75 ms post-stimulus-onset, were found to display a sigmoidal characteristic during the transition from anaesthetized to conscious states. In contrast, negative peak amplitudes within 0 to 25 ms displayed greater linearity. SignificanceThis study demonstrates a method for modelling changes in ERP morphology that accompany transitions between states of consciousness using a RNN. In future studies, this approach may be applied to human data to support the clinical use of ERPs to predict transition to consciousness.

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More evidence for a long-latency mismatch response in urethane-anaesthetised mice

Prolonged electrophysiological responses to oddball stimuli have recently been observed from anaesthetised rodents. This deviant-related activity is found to extend through 200 to 700 ms post-stimulus; a window typically obstructed from analysis by the response to subsequent stimuli in the auditory sequence. A simple methodological development in terms of difference waveform computation using two adjoining evoked responses has enabled visualisation of this activity over a longer window of analysis than previously available. In the present study, the double-epoch subtraction technique was retroactively applied to data from 13 urethane-anaesthetised mice. Oddball paradigm waveforms were compared with those of a many-standards control sequence, confirming that oddball stimuli evoked long-latency potentials that did not arise from standard or control stimuli. Statistical tests were performed at every time point from 0 to 700 ms post stimuli to highlight regions of significant difference. Oddball-induced mismatch responses were found to display significantly greater long-latency potentials than identical stimuli presented in an equal-probability context. As such, it may be concluded that long-latency potentials were evoked by the oddball condition. How this feature of the anaesthetised rodent mismatch response relates to human mismatch negativity is unclear, although it may be tentatively linked to the human P3a component, which is considered to emerge downstream from mismatch negativity.

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Classical and controlled auditory mismatch responses to multiple physical deviances in anaesthetised and conscious mice

Human mismatch negativity (MMN) is modelled in rodents and other non-human species to examine its underlying neurological mechanisms, primarily described in terms of deviance-detection and adaptation. Using the mouse model, we aim to elucidate subtle dependencies between the mismatch response (MMR) and different physical properties of sound. Epidural field potentials were recorded from urethane-anaesthetised and conscious mice during oddball and many-standards control paradigms; with stimuli varying in duration, frequency, intensity, and inter-stimulus interval. Resulting auditory evoked potentials, classical MMR (oddball - standard), and controlled MMR (oddball - control) waveforms were analysed. Stimulus duration correlated with stimulus-off response peak latency (p < 0.0001). Frequency (p < 0.0001), intensity (p < 0.0001), and inter-stimulus interval (p < 0.0001) correlated with stimulu-son N1 and P1 (conscious only) peak amplitudes. These relationships were instrumental in shaping classical MMR morphology in both anaesthetised and conscious animals, suggesting these waveforms reflect modification of normal auditory processing by different physical properties of stimuli. Controlled MMR waveforms appeared to exhibit habituation to auditory stimulation over time, which was equally observed in response to oddball and standard stimuli. These observations are not consistent with the mechanisms thought to underlie human MMN, which currently do not address differences due to specific physical features of auditory deviance. Thus, no evidence was found to objectively support the deviance-detection or adaptation hypotheses of MMN in relation to anaesthetised or conscious mice.

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