Decreased Beta Power and OFC-STN Phase Synchronization During Reactive Stopping in Freely Behaving Rats.

During natural behavior, an action often needs to be suddenly stopped in response to unexpected sensory input - referred to as reactive stopping. Reactive stopping has been mostly investigated in humans, which led to hypotheses about the involvement of different brain structures, in particular the hyperdirect pathway. Here, we directly investigate the contribution and interaction of two key regions of the hyperdirect pathway, the orbitofrontal cortex (OFC) and subthalamic nucleus (STN), using dual-area, multi-electrode recordings in male rats performing a stop-signal task. In this task rats have to initiate movement to a go-signal, and occasionally stop their movement to the go-signal side after a stop-signal, presented at various stop-signal delays.Both the OFC and STN show near-simultaneous field potential reductions in the beta frequency range (12-30 Hz) compared to the period preceding the go-signal and the movement period. These transient reductions (∼200 ms) only happen during reactive stopping, which is when the stop-signal was received after action initiation, and are well-timed after stop-signal onset and before the estimated time of stopping. Phase synchronization analysis also showed a transient attenuation of synchronization between the OFC and STN in the beta range during reactive stopping.The present results provide the first direct quantification of local neural oscillatory activity in the OFC and STN and interareal synchronization specifically timed during reactive stopping.Significance Statement Different studies observed increases in oscillatory beta activity and suggested increased synchronization between the orbitofrontal cortex (OFC) and subthalamic nucleus (STN) during reactive stopping. However, there has been inconsistency in the timing of beta modulations, and no study has yet investigated phase synchronization during stopping in both the OFC and STN with anatomical and temporal precision. Using dual-area recordings during a stopping task, we observed substantial decreases in beta power in both the OFC and STN at the time of stopping, alongside a decrease in beta phase synchronization. Rather than increased beta-band activity, the OFC and STN appear to facilitate stopping through local and inter-areal desynchronization. This may enable functionally specific neuronal activity to selectively inhibit motor behavior downstream.

Midfrontal theta dynamics index the monitoring of postural stability.

Stepping is a common strategy to recover postural stability and maintain upright balance. Postural perturbations have been linked to neuroelectrical markers such as the N1 potential and theta frequency dynamics. Here, we investigated the role of cortical midfrontal theta dynamics of balance monitoring, driven by balance perturbations at different initial standing postures. We recorded electroencephalography, electromyography, and motion tracking of human participants while they stood on a platform that delivered a range of forward and backward whole-body balance perturbations. The participants' postural threat was manipulated prior to the balance perturbation by instructing them to lean forward or backward while keeping their feet-in-place in response to the perturbation. We hypothesized that midfrontal theta dynamics index the engagement of a behavioral monitoring system and, therefore, that perturbation-induced theta power would be modulated by the initial leaning posture and perturbation intensity. Targeted spatial filtering in combination with mixed-effects modeling confirmed our hypothesis and revealed distinct modulations of theta power according to postural threat. Our results provide novel evidence that midfrontal theta dynamics subserve action monitoring of human postural balance. Understanding of cortical mechanisms of balance control is crucial for studying balance impairments related to aging and neurological conditions (e.g. stroke).

Two modes of midfrontal theta suggest a role in conflict and error processing.

Midfrontal theta increases during scenarios when conflicts are successfully resolved. Often considered a generic signal of cognitive control, its temporal nature has hardly been investigated. Using advanced spatiotemporal techniques, we uncover that midfrontal theta occurs as a transient oscillation or "event" at single trials with their timing reflecting computationally distinct modes. Single-trial analyses of electrophysiological data from participants performing the Flanker (N = 24) and Simon task (N = 15) were used to probe the relationship between theta and metrics of stimulus-response conflict. We specifically investigated "partial errors", in which a small burst of muscle activity in the incorrect response effector occurred, quickly followed by a correction. We found that transient theta events in single trials could be categorized into two distinct theta modes based on their relative timing to different task events. Theta events from the first mode occurred briefly after the task stimulus and might reflect conflict-related processing of the stimulus. In contrast, theta events from the second mode were more likely to occur around the time partial errors were committed, suggesting they were elicited by a potential upcoming error. Importantly, in trials in which a full error was committed, this "error-related theta" occurred too late with respect to the onset of the erroneous muscle response, supporting the role of theta also in error correction. We conclude that different modes of transient midfrontal theta can be adopted in single trials not only to process stimulus-response conflict, but also to correct erroneous responses.

Dynamics of Neural Microstates in the VTA-Striatal-Prefrontal Loop during Novelty Exploration in the Rat.

Neural activity at the large-scale population level has been suggested to be consistent with a sequence of brief, quasistable spatial patterns. These "microstates" and their temporal dynamics have been linked to myriad cognitive functions and brain diseases. Most of this research has been performed using EEG, leaving many questions, such as the existence, dynamics, and behavioral relevance of microstates at the level of local field potentials (LFPs), unaddressed. Here, we adapted the standard EEG microstate analysis to triple-area LFP recordings from 192 electrodes in rats to investigate the mesoscopic dynamics of neural microstates within and across brain regions during novelty exploration. We performed simultaneous recordings from the prefrontal cortex, striatum, and ventral tegmental area in male rats during awake behavior (object novelty and exploration). We found that the LFP data can be accounted for by multiple, recurring microstates that were stable for ∼60-100 ms. The simultaneous microstate activity across brain regions revealed rhythmic patterns of coactivations, which we interpret as a novel indicator of inter-regional, mesoscale synchronization. Furthermore, these rhythmic coactivation patterns across microstates were modulated by behavioral states such as movement and exploration of a novel object. These results support the existence of a functional mesoscopic organization across multiple brain areas and present a possible link of the origin of macroscopic EEG microstates to zero-lag neuronal synchronization within and between brain areas, which is of particular interest to the human research community.SIGNIFICANCE STATEMENT The coordination of neural activity across the entire brain has remained elusive. Here we combine large-scale neural recordings at fine spatial resolution with the analysis of microstates (i.e., short-lived, recurring spatial patterns of neural activity). We demonstrate that the local activity in different brain areas can be accounted for by only a few microstates per region. These microstates exhibited temporal dynamics that were correlated across regions in rhythmic patterns. We demonstrate that these microstates are linked to behavior and exhibit different properties in the frequency domain during different behavioral states. In summary, LFP microstates provide an insightful approach to studying both mesoscopic and large-scale brain activation within and across regions.

A tutorial on generalized eigendecomposition for denoising, contrast enhancement, and dimension reduction in multichannel electrophysiology.

The goal of this paper is to present a theoretical and practical introduction to generalized eigendecomposition (GED), which is a robust and flexible framework used for dimension reduction and source separation in multichannel signal processing. In cognitive electrophysiology, GED is used to create spatial filters that maximize a researcher-specified contrast. For example, one may wish to exploit an assumption that different sources have different frequency content, or that sources vary in magnitude across experimental conditions. GED is fast and easy to compute, performs well in simulated and real data, and is easily adaptable to a variety of specific research goals. This paper introduces GED in a way that ties together myriad individual publications and applications of GED in electrophysiology, and provides sample MATLAB and Python code that can be tested and adapted. Practical considerations and issues that often arise in applications are discussed.

Power spectrum slope confounds estimation of instantaneous oscillatory frequency.

Oscillatory neural dynamics are highly non-stationary and require methods capable of quantifying time-resolved changes in oscillatory activity in order to understand neural function. Recently, a method termed 'frequency sliding' was introduced to estimate the instantaneous frequency of oscillatory activity, providing a means of tracking temporal changes in the dominant frequency within a sub-band of field potential recordings. Here, the ability of frequency sliding to recover ground-truth oscillatory frequency in simulated data is tested while the exponent (slope) of the 1/fx component of the signal power spectrum is systematically varied, mimicking real electrophysiological data. The results show that 1) in the presence of 1/f activity, frequency sliding systematically underestimates the true frequency of the signal, 2) the magnitude of underestimation is correlated with the steepness of the slope, suggesting that, if unaccounted for, slope changes could be misinterpreted as frequency changes, 3) the impact of slope on frequency estimates interacts with oscillation amplitude, indicating that changes in oscillation amplitude alone may also influence instantaneous frequency estimates in the presence of strong 1/f activity; and 4) analysis parameters such as filter bandwidth and location also mediate the influence of slope on estimated frequency, indicating that these settings should be considered when interpreting estimates obtained via frequency sliding. The origin of these biases resides in the output of the filtering step of frequency sliding, whose energy is biased towards lower frequencies precisely because of the 1/f structure of the data. We discuss several strategies to mitigate these biases and provide a proof-of-principle for a 1/f normalization strategy.

Intra- and inter-regional dynamics in cortical-striatal-tegmental networks.

It is increasingly recognized that networks of brain areas work together to accomplish computational goals. However, functional connectivity networks are not often compared between different behavioral states and across different frequencies of electrical oscillatory signals. In addition, connectivity is always defined as the strength of signal relatedness between two atlas-based anatomical locations. Here, we performed an exploratory analysis using data collected from high-density arrays in the prefrontal cortex (PFC), striatum (STR), and ventral tegmental area (VTA) of male rats. These areas have all been implicated in a wide range of different tasks and computations including various types of memory as well as reward valuation, habit formation and execution, and skill learning. Novel intraregional clustering analyses identified patterns of spatially restricted, temporally coherent, and frequency-specific signals that were reproducible across days and were modulated by behavioral states. Multiple clusters were identified within each anatomical region, indicating a mesoscopic scale of organization. Generalized eigendecomposition (GED) was used to dimension-reduce each cluster to a single component time series. Dense intercluster connectivity was modulated by behavioral state, with connectivity becoming reduced when the animals were exposed to a novel object, compared with a baseline condition. Behavior-modulated connectivity changes were seen across the spectrum, with δ, θ, and γ all being modulated. These results demonstrate the brain's ability to reorganize functionally at both the intra- and inter-regional levels during different behavioral states.NEW & NOTEWORTHY We applied novel clustering techniques to discover functional subregional anatomical patches that changed with behavioral conditions but were frequency specific and stable across days. By taking into account these changes in intraregional signal generator location and extent, we were able to reveal a richer picture of inter-regional functional connectivity than would otherwise have been possible. These findings reveal that the brain's functional organization changes with state at multiple levels of scale.

Multiple Midfrontal Thetas Revealed by Source Separation of Simultaneous MEG and EEG.

Theta-band (∼6 Hz) rhythmic activity within and over the medial PFC ("midfrontal theta") has been identified as a distinctive signature of "response conflict," the competition between multiple actions when only one action is goal-relevant. Midfrontal theta is traditionally conceptualized and analyzed under the assumption that it is a unitary signature of conflict that can be uniquely identified at one electrode (typically FCz). Here we recorded simultaneous MEG and EEG (total of 328 sensors) in 9 human subjects (7 female) and applied a feature-guided multivariate source-separation decomposition to determine whether conflict-related midfrontal theta is a unitary or multidimensional feature of the data. For each subject, a generalized eigendecomposition yielded spatial filters (components) that maximized the ratio between theta and broadband activity. Components were retained based on significance thresholding and midfrontal EEG topography. All of the subjects individually exhibited multiple (mean 5.89, SD 2.47) midfrontal components that contributed to sensor-level midfrontal theta power during the task. Component signals were temporally uncorrelated and asynchronous, suggesting that each midfrontal theta component was unique. Our findings call into question the dominant notion that midfrontal theta represents a unitary process. Instead, we suggest that midfrontal theta spans a multidimensional space, indicating multiple origins, but can manifest as a single feature at the sensor level because of signal mixing.SIGNIFICANCE STATEMENT "Midfrontal theta" is a rhythmic electrophysiological signature of the competition between multiple response options. Midfrontal theta is traditionally considered to reflect a single process. However, this assumption could be erroneous because of "mixing" (multiple sources contributing to the activity recorded at a single electrode). We investigated the dimensionality of midfrontal theta by applying advanced multivariate analysis methods to a multimodal MEG/EEG dataset. We identified multiple topographically overlapping neural sources that drove response conflict-related midfrontal theta. Midfrontal theta thus reflects multiple uncorrelated signals that manifest with similar EEG scalp projections. In addition to contributing to the cognitive control literature, we demonstrate both the feasibility and the necessity of signal demixing to understand the narrowband neural dynamics underlying cognitive processes.

Synchronization between Keyboard Typing and Neural Oscillations.

Rhythmic neural activity synchronizes with certain rhythmic behaviors, such as breathing, sniffing, saccades, and speech. The extent to which neural oscillations synchronize with higher-level and more complex behaviors is largely unknown. Here, we investigated electrophysiological synchronization with keyboard typing, which is an omnipresent behavior daily engaged by an uncountably large number of people. Keyboard typing is rhythmic, with frequency characteristics roughly the same as neural oscillatory dynamics associated with cognitive control, notably through midfrontal theta (4-7 Hz) oscillations. We tested the hypothesis that synchronization occurs between typing and midfrontal theta and breaks down when errors are committed. Thirty healthy participants typed words and sentences on a keyboard without visual feedback, while EEG was recorded. Typing rhythmicity was investigated by interkeystroke interval analyses and by a kernel density estimation method. We used a multivariate spatial filtering technique to investigate frequency-specific synchronization between typing and neuronal oscillations. Our results demonstrate theta rhythmicity in typing (around 6.5 Hz) through the two different behavioral analyses. Synchronization between typing and neuronal oscillations occurred at frequencies ranging from 4 to 15 Hz, but to a larger extent for lower frequencies. However, peak synchronization frequency was idiosyncratic across participants, therefore not specific to theta nor to midfrontal regions, and correlated somewhat with peak typing frequency. Errors and trials associated with stronger cognitive control were not associated with changes in synchronization at any frequency. As a whole, this study shows that brain-behavior synchronization does occur during keyboard typing but is not specific to midfrontal theta.

Cortical responses to whole-body balance perturbations index perturbation magnitude and predict reactive stepping behavior.

The goal of this study was to determine whether the cortical responses elicited by whole-body balance perturbations were similar to established cortical markers of action monitoring. Postural changes imposed by balance perturbations elicit a robust negative potential (N1) and a brisk increase of theta activity in the electroencephalogram recorded over midfrontal scalp areas. Because action monitoring is a cognitive function proposed to detect errors and initiate corrective adjustments, we hypothesized that the possible cortical markers of action monitoring during balance control (N1 potential and theta rhythm) scale with perturbation intensity and the eventual execution of reactive stepping responses (as opposed to feet-in-place responses). We recorded high-density electroencephalogram from eleven young individuals, who participated in an experimental balance assessment. The participants were asked to recover balance following anteroposterior translations of the support surface at various intensities, while attempting to maintain both feet in place. We estimated source-resolved cortical activity using independent component analysis. Combining time-frequency decomposition and group-level general linear modeling of single-trial responses, we found a significant relation of the interaction between perturbation intensity and stepping responses with multiple cortical features from the midfrontal cortex, including the N1 potential, and theta, alpha, and beta rhythms. Our findings suggest that the cortical responses to balance perturbations index the magnitude of a deviation from a stable postural state to predict the need for reactive stepping responses. We propose that the cortical control of balance may involve cognitive control mechanisms (i.e., action monitoring) that facilitate postural adjustments to maintain postural stability.

EEG microstates as a continuous phenomenon.

In recent years, EEG microstate analysis has gained popularity as a tool to characterize spatio-temporal dynamics of large-scale electrophysiology data. It has been used in a wide range of EEG studies and the discovered microstates have been linked to cognitive function and brain diseases. EEG microstates are assumed to (1) be winner-take-all, meaning that the topography at any given time point is in one state; and (2) discretely transition from one state into another. In this study we investigated these assumptions by taking a geometric perspective of EEG data, treating microstate topographies as basis vectors for a subspace of the original channel space. We found that within- and across-microstate distance distributions were largely overlapping: for the low GFP (Global Field Power) range (lower 15%), individual time points labeled as one microstate are often equidistant to multiple microstate vectors, challenging the winner-take-all assumption. At high global field power, separability of microstates improved, but remained rather weak. Although many GFP peaks (which are the time points used for defining microstates) occur during high GFP ranges, low GFP ranges associated with poor separability also contain GFP peaks. Furthermore, the geometric analysis suggested that microstates and their transitions appear to be more continuous than discrete. The Analysis of rate of change of trajectory in sensor space suggests gradual microstate transitions as opposed to the classical binary view of EEG microstates. Taken together, our findings suggest that EEG microstates are better conceptualized as spatially and temporally continuous, rather than discrete activations of neural populations.

Midfrontal theta phase coordinates behaviorally relevant brain computations during cognitive control.

Neural oscillations are thought to provide a cyclic time frame for orchestrating brain computations. Following this assumption, midfrontal theta oscillations have recently been proposed to temporally organize brain computations during conflict processing. Using a multivariate analysis approach, we show that brain-behavior relationships during conflict tasks are modulated according to the phase of ongoing endogenous midfrontal theta oscillations recorded by scalp EEG. We found reproducible results in two independent datasets, using two different conflict tasks: brain-behavior relationships (correlation between reaction time and theta power) were theta phase-dependent in a subject-specific manner, and these "behaviorally optimal" theta phases were also associated with fronto-parietal cross-frequency dynamics emerging as theta phase-locked beta power bursts. These effects were present regardless of the strength of conflict. Thus, these results provide empirical evidence that midfrontal theta oscillations are involved in cyclically orchestrating brain computations likely related to response execution during the tasks rather than purely related to conflict processing. More generally, this study supports the hypothesis that phase-based computation is an important mechanism giving rise to cognitive processing.

How the Level of Reward Awareness Changes the Computational and Electrophysiological Signatures of Reinforcement Learning.

The extent to which subjective awareness influences reward processing, and thereby affects future decisions, is currently largely unknown. In the present report, we investigated this question in a reinforcement learning framework, combining perceptual masking, computational modeling, and electroencephalographic recordings (human male and female participants). Our results indicate that degrading the visibility of the reward decreased, without completely obliterating, the ability of participants to learn from outcomes, but concurrently increased their tendency to repeat previous choices. We dissociated electrophysiological signatures evoked by the reward-based learning processes from those elicited by the reward-independent repetition of previous choices and showed that these neural activities were significantly modulated by reward visibility. Overall, this report sheds new light on the neural computations underlying reward-based learning and decision-making and highlights that awareness is beneficial for the trial-by-trial adjustment of decision-making strategies.SIGNIFICANCE STATEMENT The notion of reward is strongly associated with subjective evaluation, related to conscious processes such as "pleasure," "liking," and "wanting." Here we show that degrading reward visibility in a reinforcement learning task decreases, without completely obliterating, the ability of participants to learn from outcomes, but concurrently increases subjects' tendency to repeat previous choices. Electrophysiological recordings, in combination with computational modeling, show that neural activities were significantly modulated by reward visibility. Overall, we dissociate different neural computations underlying reward-based learning and decision-making, which highlights a beneficial role of reward awareness in adjusting decision-making strategies.

A better way to define and describe Morlet wavelets for time-frequency analysis.

Complex Morlet wavelets are frequently used for time-frequency analysis of non-stationary time series data, such as neuroelectrical signals recorded from the brain. The crucial parameter of Morlet wavelets is the width of the Gaussian that tapers the sine wave. This width parameter controls the trade-off between temporal precision and spectral precision. It is typically defined as the "number of cycles," but this parameter is opaque, and often leads to uncertainty and suboptimal analysis choices, as well as being difficult to interpret and evaluate. The purpose of this paper is to present alternative formulations of Morlet wavelets in time and in frequency that allow parameterizing the wavelets directly in terms of the desired temporal and spectral smoothing (expressed as full-width at half-maximum). This formulation provides clarity on an important data analysis parameter, and can facilitate proper analyses, reporting, and interpretation of results. MATLAB code and sample data are provided.

Individual Alpha Peak Frequency Predicts 10 Hz Flicker Effects on Selective Attention.

Rhythmic visual stimulation ("flicker") is primarily used to "tag" processing of low-level visual and high-level cognitive phenomena. However, preliminary evidence suggests that flicker may also entrain endogenous brain oscillations, thereby modulating cognitive processes supported by those brain rhythms. Here we tested the interaction between 10 Hz flicker and endogenous alpha-band (∼10 Hz) oscillations during a selective visuospatial attention task. We recorded EEG from human participants (both genders) while they performed a modified Eriksen flanker task in which distractors and targets flickered within (10 Hz) or outside (7.5 or 15 Hz) the alpha band. By using a combination of EEG source separation, time-frequency, and single-trial linear mixed-effects modeling, we demonstrate that 10 Hz flicker interfered with stimulus processing more on incongruent than congruent trials (high vs low selective attention demands). Crucially, the effect of 10 Hz flicker on task performance was predicted by the distance between 10 Hz and individual alpha peak frequency (estimated during the task). Finally, the flicker effect on task performance was more strongly predicted by EEG flicker responses during stimulus processing than during preparation for the upcoming stimulus, suggesting that 10 Hz flicker interfered more with reactive than proactive selective attention. These findings are consistent with our hypothesis that visual flicker entrained endogenous alpha-band networks, which in turn impaired task performance. Our findings also provide novel evidence for frequency-dependent exogenous modulation of cognition that is determined by the correspondence between the exogenous flicker frequency and the endogenous brain rhythms.SIGNIFICANCE STATEMENT Here we provide novel evidence that the interaction between exogenous rhythmic visual stimulation and endogenous brain rhythms can have frequency-specific behavioral effects. We show that alpha-band (10 Hz) flicker impairs stimulus processing in a selective attention task when the stimulus flicker rate matches individual alpha peak frequency. The effect of sensory flicker on task performance was stronger when selective attention demands were high, and was stronger during stimulus processing and response selection compared with the prestimulus anticipatory period. These findings provide novel evidence that frequency-specific sensory flicker affects online attentional processing, and also demonstrate that the correspondence between exogenous and endogenous rhythms is an overlooked prerequisite when testing for frequency-specific cognitive effects of flicker.

Oscillatory Mechanisms of Response Conflict Elicited by Color and Motion Direction: An Individual Differences Approach.

Goal-directed behavior requires control over automatic behavior, for example, when goal-irrelevant information from the environment captures an inappropriate response and conflicts with the correct, goal-relevant action. Neural oscillations in the theta band (∼6 Hz) measured at midfrontal electrodes are thought to form an important substrate of the detection and subsequent resolution of response conflict. Here, we examined the extent to which response conflict and associated theta-band activity depend on the visual stimulus feature dimension that triggers the conflict. We used a feature-based Simon task to manipulate conflict by motion direction and stimulus color. Analyses were focused on individual differences in behavioral response conflict elicited across different stimulus dimensions and their relationship to conflict-related midfrontal theta. We first confirmed the presence of response conflict elicited by task-irrelevant motion and stimulus color, demonstrating the usefulness of our modified version of the Simon task to assess different sensory origins of response conflict. Despite titrating overall task performance, we observed large individual differences in the behavioral manifestations of response conflict elicited by the different visual dimensions. These behavioral conflict effects were mirrored in a dimension-specific relationship with conflict-related midfrontal theta power, such that, for each dimension, individual midfrontal theta power was generally higher when experienced response conflict was high. Finally, exploratory analyses of interregional functional connectivity suggested a role for phase synchronization between frontal and parietal scalp sites in modulating experienced conflict when color was the task-relevant visual dimension. Highlighting the importance of an individual differences approach in cognitive neuroscience, these results reveal large individual differences in experienced response conflict depending on the source of visual interference, which are predicted by conflict-related midfrontal theta power.

Interactions between frontal and posterior oscillatory dynamics support adjustment of stimulus processing during reinforcement learning.

Reinforcement learning (RL) in humans is subserved by a network of striatal and frontal brain areas. The electrophysiological signatures of feedback evaluation are increasingly well understood, but how those signatures relate to the use of feedback to guide subsequent behavioral adjustment remains unclear. One mechanism for post-feedback behavioral optimization is the modulation of sensory processing. We used source-reconstructed MEG to test whether feedback affects the interactions between sources of oscillatory activity in the learning network and task-relevant stimulus-processing areas. Participants performed a probabilistic RL task in which they learned associations between colored faces and response buttons using trial-and-error feedback. Delta-band (2-4 Hz) and theta-band (4-8 Hz) power in multiple frontal regions were sensitive to feedback valence. Low and high beta-band power (12-20 and 20-30 Hz) in occipital, parietal, and temporal regions differentiated between color and face information. Consistent with our hypothesis, single-trial power-power correlations between frontal and posterior-sensory areas were modulated by the interaction between feedback valence and the relevant stimulus characteristic (color versus identity). These results suggest that long-range oscillatory coupling supports post-feedback updating of stimulus processing.

Using spatiotemporal source separation to identify prominent features in multichannel data without sinusoidal filters.

The number of simultaneously recorded electrodes in neuroscience is steadily increasing, providing new opportunities for understanding brain function, but also new challenges for appropriately dealing with the increase in dimensionality. Multivariate source separation analysis methods have been particularly effective at improving signal-to-noise ratio while reducing the dimensionality of the data and are widely used for cleaning, classifying and source-localizing multichannel neural time series data. Most source separation methods produce a spatial component (that is, a weighted combination of channels to produce one time series); here, this is extended to apply source separation to a time series, with the idea of obtaining a weighted combination of successive time points, such that the weights are optimized to satisfy some criteria. This is achieved via a two-stage source separation procedure, in which an optimal spatial filter is first constructed and then its optimal temporal basis function is computed. This second stage is achieved with a time-delay-embedding matrix, in which additional rows of a matrix are created from time-delayed versions of existing rows. The optimal spatial and temporal weights can be obtained by solving a generalized eigendecomposition of covariance matrices. The method is demonstrated in simulated data and in an empirical electroencephalogram study on theta-band activity during response conflict. Spatiotemporal source separation has several advantages, including defining empirical filters without the need to apply sinusoidal narrowband filters.

Rhythmic entrainment source separation: Optimizing analyses of neural responses to rhythmic sensory stimulation.

Steady-state evoked potentials (SSEPs) are rhythmic brain responses to rhythmic sensory stimulation, and are often used to study perceptual and attentional processes. We present a data analysis method for maximizing the signal-to-noise ratio of the narrow-band steady-state response in the frequency and time-frequency domains. The method, termed rhythmic entrainment source separation (RESS), is based on denoising source separation approaches that take advantage of the simultaneous but differential projection of neural activity to multiple electrodes or sensors. Our approach is a combination and extension of existing multivariate source separation methods. We demonstrate that RESS performs well on both simulated and empirical data, and outperforms conventional SSEP analysis methods based on selecting electrodes with the strongest SSEP response, as well as several other linear spatial filters. We also discuss the potential confound of overfitting, whereby the filter captures noise in absence of a signal. Matlab scripts are available to replicate and extend our simulations and methods. We conclude with some practical advice for optimizing SSEP data analyses and interpreting the results.

Human Frontal-Subcortical Circuit and Asymmetric Belief Updating.

How humans integrate information to form beliefs about reality is a question that has engaged scientists for centuries, yet the biological system supporting this process is not well understood. One of the most salient attributes of information is valence. Whether a piece of news is good or bad is critical in determining whether it will alter our beliefs. Here, we reveal a frontal-subcortical circuit in the left hemisphere that is simultaneously associated with enhanced integration of favorable information into beliefs and impaired integration of unfavorable information. Specifically, for favorable information, stronger white matter connectivity within this system, particularly between the left inferior frontal gyrus (IFG) and left subcortical regions (including the amygdala, hippocampus, thalamus, putamen, and pallidum), as well as insular cortex, is associated with greater change in belief. However, for unfavorable information, stronger connectivity within this system, particularly between the left IFG and left pallidum, putamen, and insular cortex, is associated with reduced change in beliefs. These novel results are consistent with models suggesting that partially separable processes govern learning from favorable and unfavorable information. SIGNIFICANCE STATEMENT: Beliefs of what may happen in the future are important, because they guide decisions and actions. Here, we illuminate how structural brain connectivity is related to the generation of subjective beliefs. We focus on how the valence of information is related to people's tendency to alter their beliefs. By quantifying the extent to which participants update their beliefs in response to desirable and undesirable information and relating those measures to the strength of white matter connectivity using diffusion tensor imaging, we characterize a left frontal-subcortical system that is associated simultaneously with greater belief updating in response to favorable information and reduced belief updating in response to unfavorable information. This neural architecture may allow valence to be incorporated into belief updating.