bioRxiv Science⌕ Search

Biology subjects

Jerbi, K.

Publications and source records attributed to Jerbi, K..

32 records · Page 2Linked to original sources

Class imbalance should not throw you off balance: Choosing classifiers and performance metrics for brain decoding with imbalanced data

Machine learning (ML) is increasingly used in cognitive, computational and clinical neuroscience. The reliable and efficient application of ML requires a sound understanding of its subtleties and limitations. Training ML models on datasets with imbalanced classes is a particularly common problem, and it can have severe consequences if not adequately addressed. With the neuroscience ML user in mind, this paper provides a didactic assessment of the class imbalance problem and illustrates its impact through systematic manipulation of data imbalance ratios in (i) simulated data and (ii) brain data recorded with electroencephalography (EEG) and magnetoencephalography (MEG). Our results illustrate how the widely-used Accuracy (Acc) metric, which measures the overall proportion of successful predictions, yields misleadingly high performances, as class imbalance increases. Because Acc weights the per-class ratios of correct predictions proportionally to class size, it largely disregards the performance on the minority class. A binary classification model that learns to systematically vote for the majority class will yield an artificially high decoding accuracy that directly reflects the imbalance between the two classes, rather than any genuine generalizable ability to discriminate between them. We show that other evaluation metrics such as the Area Under the Curve (AUC) of the Receiver Operating Characteristic (ROC), and the less common Balanced Accuracy (BAcc) metric - defined as the arithmetic mean between sensitivity and specificity, provide more reliable performance evaluations for imbalanced data. Our findings also highlight the robustness of Random Forest (RF), and the benefits of using stratified cross-validation and hyperprameter optimization to tackle data imbalance. Critically, for neuroscience ML applications that seek to minimize overall classification error, we recommend the routine use of BAcc, which in the specific case of balanced data is equivalent to using standard Acc, and readily extends to multi-class settings. Importantly, we present a list of recommendations for dealing with imbalanced data, as well as open-source code to allow the neuroscience community to replicate and extend our observations and explore alternative approaches to coping with imbalanced data.

neuroscience↗

Rhythmic information sampling in the brain during visual recognition

When we fixate an object, visual information is continuously received on the retina. Several studies observed behavioral oscillations in perceptual sensitivity across such stimulus time, and these fluctuations have been linked to brain oscillations. However, whether specific brain areas show oscillations across stimulus presentation time (i.e., different time points of the stimulus being more or less processed, in a rhythmic fashion) has not been investigated. Here, we revealed random areas of face images at random moments across time and recorded the brain activity of human participants (both male and female) using magnetoencephalography (MEG) while they performed two recognition tasks. This allowed us to quantify how each snapshot of visual information coming from the stimulus is processed across time and across the brain. Oscillations across stimulus time (rhythmic sampling) were mostly visible in early visual areas, at theta, alpha and low beta frequencies. We also found that they contributed to brain activity more than previously investigated rhythmic processing (oscillations in the processing of a single snapshot of visual information). Non-rhythmic sampling was also visible at later latencies across the visual cortex, either in the form of a transient processing of early stimulus time points or of a sustained processing of the whole stimulus. Our results suggest that successive cycles of ongoing brain oscillations process stimulus information incoming at successive moments. Together, these results advance our understanding of the oscillatory neural dynamics associated with visual processing and show the importance of considering the temporal dimension of stimuli when studying visual recognition. Significance StatementSeveral behavioral studies have observed oscillations in perceptual sensitivity over the duration of stimulus presentation, and these fluctuations have been linked to brain oscillations. However, oscillations across stimulus time in the brain have not been studied. Here, we developed a MEG paradigm to quantify how visual information received at each moment during fixation is processed through time and across the brain. We showed that different snapshots of a stimulus are distinctly processed in many brain areas and that these fluctuations are oscillatory in early visual areas. Oscillations across stimulus time were more prevalent than previously studied oscillations across processing time. These results increase our understanding of how neural oscillations interact with the visual processing of temporal stimuli.

neuroscience↗

Distinct trajectories in low-dimensional neural oscillation state space track dynamic decision-making in humans

The brain evolved to govern behavior in a dynamic world, in which pertinent information about choices is often in flux. Thus, the commitment to an action choice must reflect a balance between monitoring that information and the necessity to act before opportunities are lost. Here, we investigate the mechanisms of dynamic decision-making in humans using low dimensional space representation of brain wide magnetoencephalography recordings. We show that the principal components (PCs) of alpha (9-13 Hz) and beta power (16-24 Hz) are involved in tracking sensory information evolving over time in the sensorimotor and visual cortex. We also found that alpha PCs reflect the commitment to a particular choice, while beta PCs reflect motor execution. Finally, higher frequency components in subcortical areas reflect the adjustment of speed- accuracy tradeoff policies. These results provide a new detailed characterization of the distributed oscillatory brain processes underlying dynamic decision-making in humans.

neuroscience↗

Aperiodic brain activity and response to anesthesia vary in disorders of consciousness

The analysis of human EEG has traditionally focused on oscillatory power, which is characterized by peaks above an aperiodic component in the power spectral density. This study investigates the aperiodic EEG component of individuals in a disorder of consciousness (DOC); how it changes in response to exposure to anesthesia; and how it relates to the brains information richness and criticality. High-density EEG was recorded from 43 individuals in a DOC, with 16 of these individuals undergoing a protocol of propofol anesthesia. The aperiodic component was defined by the spectral slope of the power spectral density. Our results demonstrate that the EEG aperiodic component is more informative about the participants level of consciousness than the oscillatory component. Importantly, the pharmacologically induced change in the spectral slope from 30-45 Hz positively correlated with individuals pre-anesthetic level of consciousness. The pharmacologically induced loss of information-richness and criticality was associated with individuals pre-anesthetic aperiodic component. During exposure to anesthesia, the aperiodic component was correlated with 3-month recovery status for individuals with DOC. The aperiodic EEG component has been historically neglected; this research highlights the necessity of considering this measure for the assessment of individuals in DOC and future research that seeks to understand the neurophysiological underpinnings of consciousness.

neuroscience↗

Altered brain criticality in Schizophrenia: New insights from MEG

Schizophrenia has a complex etiology and symptomatology that is difficult to untangle. After decades of research, important advancements towards a central biomarker are still lacking. One of the missing pieces is a better understanding of how non-linear neural dynamics are altered in this patient population. In this study, the resting-state neuromagnetic signals of schizophrenia patients and healthy controls were analyzed in the framework of criticality. When biological systems like the brain are in a state of criticality, they are thought to be functioning at maximum efficiency (e.g., optimal communication and storage of information) and with maximum adaptability to incoming information. Here, we assessed the self-similarity and multifractality of resting-state brain signals recorded with magnetoencephalography in patients with schizophrenia patients and in matched controls. Our analysis showed a clear ascending, rostral to caudal gradient of self-similarity values in healthy controls, and an opposite gradient for multifractality (descending values, rostral to caudal). Schizophrenia patients had similar, although attenuated, gradients of self-similarity and multifractality values. Statistical tests showed that patients had higher values of self-similarity than controls in fronto-temporal regions, indicative of more regularity and memory in the signal. In contrast, patients had less multifractality than controls in the parietal and occipital regions, indicative of less diverse singularities and reduced variability in the signal. In addition, supervised machine-learning, based on logistic regression, successfully discriminated the two groups using measures of self-similarity and multifractality as features. Our results provide new insights into the baseline cognitive functioning of schizophrenia patients by identifying key alterations of criticality properties in their resting-state brain data.

neuroscience↗

Neural oscillations track natural but not artificial fast speech: Novel insights from speech-brain coupling using MEG

Speech processing is supported by the synchronization of cortical oscillations to its rhythmic components, including syllable rate. This has been shown to be the case for normal rate speech as well as artificially accelerated speech. However, the case of natural speech rate variations, which are among the most ubiquitous sources of variability in speech, has been largely overlooked. Here, we directly compared changes in the properties of cortico-acoustic coupling when speech naturally shifts from normal to fast rate and when it is artificially accelerated. Neuromagnetic brain signals of 24 normal-hearing adults were recorded with magnetoencephalography (MEG) while they listened to natural normal ([~]6 syllables/s), natural fast ([~]9 syllables/s) and time-compressed ([~]9 syllables/s) sentences, as well as to envelope-matched amplitude-modulated noise. We estimated coherence between the envelope of the acoustic input and MEG source time-series at frequencies corresponding to the mean syllable rates of the normal and fast speech stimuli. We found that listening to natural speech at normal and fast rates was associated with coupling between speech signal envelope and neural oscillations in right auditory and (pre)motor cortices. This oscillatory alignment occurred at [~]6.25 Hz for normal rate sentences and shifted up to [~]8.75 Hz for naturally-produced fast speech, mirroring the increase in syllable rate between the two conditions. Unexpectedly, despite being generated at the same rate as naturally-produced fast speech, the time-compressed sentences did not lead to significant cortico-acoustic coupling at [~]8.75 Hz. Interestingly, neural activity in putative right articulatory cortex exhibited stronger tuning to natural fast rather than to artificially accelerated speech, as well as stronger phase-coupling with left temporo-parietal and motor regions. This may reflect enhanced tracking of articulatory features of naturally-produced speech. Altogether, our findings provide new insights into the oscillatory brain signature underlying the perception of natural speech at different rates and highlight the importance of using naturally-produced speech when probing the dynamics of brain-to-speech coupling.

neuroscience↗

Tensorpac : an open-source Python toolbox for tensor-based Phase-Amplitude Couplingmeasurement in electrophysiological brain signals

Despite being the focus of a thriving field of research, the biological mechanisms that underlie information integration in the brain are not yet fully understood. A theory that has gained a lot of traction in recent years suggests that multi-scale integration is regulated by a hierarchy of mutually interacting neural oscillations. In particular, there is accumulating evidence that phase-amplitude coupling (PAC), a specific form of cross-frequency interaction, plays a key role in numerous cognitive processes. Current research in the field is not only hampered by the absence of a gold standard for PAC analysis, but also by the computational costs of running exhaustive computations on large and high-dimensional electrophysiological brain signals. In addition, various signal properties and analyses parameters can lead to spurious PAC. Here, we present Tensorpac, an open-source Python toolbox dedicated to PAC analysis of neurophysiological data. The advantages of Tensorpac include (1) higher computational efficiency thanks to software design that combines tensor computations and parallel computing, (2) the implementation of all most widely used PAC methods in one package, (3) the statistical analysis of PAC measures, and (4) extended PAC visualization capabilities. Tensorpac is distributed under a BSD-3-Clause license and can be launched on any operating system (Linux, OSX and Windows). It can be installed directly via pip or downloaded from Github (https://github.com/EtienneCmb/tensorpac). By making Tensorpac available, we aim to enhance the reproducibility and quality of PAC research, and provide open tools that will accelerate future method development in neuroscience.

bioinformatics↗

NeuroPycon: A free Python toolbox for fast multi-modal and reproducible brain connectivity pipelines

Recent years have witnessed a massive push towards reproducible research in neuroscience. Unfortunately, this endeavor is often challenged by the large diversity of tools used, project-specific custom code and the difficulty to track all user-defined parameters. NeuroPycon is an open-source multi-modal brain data analysis toolkit which provides Python-based template pipelines for advanced multi-processing of MEG, EEG, functional and anatomical MRI data, with a focus on connectivity and graph theoretical analyses. Importantly, it provides shareable parameter files to facilitate replication of all analysis steps. NeuroPycon is based on the NiPype framework which facilitates data analyses by wrapping many commonly-used neuroimaging software tools into a common Python environment. In other words, rather than being a brain imaging software with is own implementation of standard algorithms for brain signal processing, NeuroPycon seamlessly integrates existing packages (coded in python, Matlab or other languages) into a unified python framework. Importantly, thanks to the multi-threaded processing and computational efficiency afforded by NiPype, NeuroPycon provides an easy option for fast parallel processing, which critical when handling large sets of multi-dimensional brain data. Moreover, its flexible design allows users to easily configure analysis pipelines by connecting distinct nodes to each other. Each node can be a Python-wrapped module, a user-defined function or a well-established tool (e.g. MNE-Python for MEG analysis, Radatools for graph theoretical metrics, etc.). Last but not least, the ability to use NeuroPycon parameter files to fully describe any pipeline is an important feature for reproducibility, as they can be shared and used for easy replication by others. The current implementation of NeuroPycon contains two complementary packages: The first, called ephypype, includes pipelines for electrophysiology analysis and a command-line interface for on the fly pipeline creation. Current implementations allow for MEG/EEG data import, pre-processing and cleaning by automatic removal of ocular and cardiac artefacts, in addition to sensor or source-level connectivity analyses. The second package, called graphpype, is designed to investigate functional connectivity via a wide range of graph-theoretical metrics, including modular partitions. The present article describes the philosophy, architecture, and functionalities of the toolkit and provides illustrative examples through interactive notebooks. NeuroPycon is available for download via github (https://github.com/neuropycon) and the two principal packages are documented online (https://neuropycon.github.io/ephypype/index.html. and https://neuropycon.github.io/graphpype/index.html). Future developments include fusion of multi-modal data (eg. MEG and fMRI or intracranial EEG and fMRI). We hope that the release of NeuroPycon will attract many users and new contributors, and facilitate the efforts of our community towards open source tool sharing and development, as well as scientific reproducibility.

neuroscience↗

Decoding the neural dynamics of oculomotor decision making in humans

How do we choose a particular action among equally valid alternatives? Non-human primate findings have shown that decision-making implicates modulations in unit firing rates and local field potentials (LFPs) across frontal and parietal cortices. Yet the electrophysiological brain mechanisms that underlie free choice in humans remain ill defined. Here, we address this question using rare intracerebral EEG recordings in surgical epilepsy patients performing a delayed oculomotor decision task. We find that the temporal dynamics of high gamma (HG, 60-140 Hz) neural activity in distinct frontal and parietal brain areas robustly discriminate free choice from instructed saccade planning at the level of single trials. Classification analysis was applied to the LFP signals to isolate decision-related activity from sensory and motor planning processes. Compared to instructed saccades, free choice trials exhibited delayed and longer-lasting HG activity. The temporal dynamics of these sustained decision-related responses distinguished deliberation-related from working memory processes. Taken together, these findings provide the first direct electrophysiological evidence in humans for the role of sustained high-frequency neural activation in fronto-parietal cortex in mediating the intrinsically driven process of freely choosing among competing behavioral alternatives. HighlightsO_LIFirst intracerebral recordings in humans performing an oculomotor decision-making task C_LIO_LIMachine learning analytics unravel underlying spectral and temporal brain dynamics C_LIO_LIFree choice trials exhibit sustained fronto-parietal high gamma (HG) activity during the delay C_LIO_LIMaking a decision and maintaining it in working memory are associated with distinct sustained HG dynamics C_LI

neuroscience↗

The processing of continuously received visual information

Visual object recognition seems to occur almost instantaneously. However, not only does it require hundreds of milliseconds of processing, but our eyes also typically fixate the object for hundreds of milliseconds. Consequently, information reaching our eyes at different moments is processed in the brain together. Moreover, information received at different moments during fixation is likely to be processed differently, notably because different features might be selectively attended at different moments. Here, we introduce a novel reverse correlation paradigm that allows us to uncover with millisecond precision the processing time course of specific information received on the retina at specific moments. Using faces as stimuli, we observed that processing at several electrodes and latencies was different depending on the moment at which information was received. Some of these variations were caused by a disruption occurring 160-200 ms after the face onset, suggesting a role of the N170 ERP component in gating information processing; others hinted at temporal compression and integration mechanisms. Importantly, the observed differences were not explained by simple adaptation or repetition priming, they were modulated by the task, and they were correlated with differences in behavior. These results suggest that top-down routines of information sampling are applied to the continuous visual input, even within a single eye fixation.

neuroscience↗

Estimation of task-based modulations in functional connectivity with MEG: a comparison of methods

For the assessment of functional interactions between distinct brain regions there is a great variety of mathematical techniques, with well-known properties, relative merits and shortcomings; however, the methods that deal specifically with task-based fluctuations in interareal coupling are scarce, and their relative performance is unclear. In the present article, we compare two approaches used in the estimation of correlation changes between the envelope amplitudes of narrowband brain activity obtained from magnetoencephalography (MEG) recordings. One approach is an implementation of semipartial canonical correlation analysis (SP-CCA), which is formally equivalent to the psychophysiological interactions technique successfully applied to functional magnetic resonance data. The other approach, which has been used in recent electrophysiology studies, consists of simply computing linear correlation coefficients of signals from two experimental conditions and taking their differences. We compared the two approaches with simulations and with multi-subject MEG signals acquired during a visuomotor coordination study. The analyses with simulated activity showed that computing differences in correlation coefficients (DCC) provided better discrimination between true coupling changes and spurious effects; on the other hand, SP-CCA resulted in significant effects around the reference location which were not found with DCC, and which may be due to field spread. Based on our findings, we recommend the use of DCC for the detection of task-based changes in connectivity, as it provided better performance than SP-CCA.

neuroscience↗

Estimation of narrowband amplitude and phase from electrophysiology signals for phase-amplitude coupling studies: a comparison of methods

Many functional connectivity studies based on electrophysiological measurements, such as electro- and magnetoencephalography (EEG/MEG), start their investigations by extracting a narrowband representation of brain activity time series, and then computing their envelope amplitudes and instantaneous phases, which serve as inputs to subsequent data processing. The two most popular approaches for obtaining these narrowband amplitudes and phases are: bandpass filtering followed by Hilbert transform (we call this the Hilbert approach); and convolution with wavelet kernels (the wavelet approach). In this work, we investigate how these two approaches perform in detecting the phenomenon of phase-amplitude coupling (PAC), whereby the amplitude of a high-frequency signal is driven by the phase of a low-frequency signal. The comparison of both approaches is carried out by means of simulated brain activity, from which we run receiver operating characteristic (ROC) analyses, and of experimental MEG data from a visuomotor coordination study. The ROC analyses show that both approaches have comparable accuracy, except in the presence of interfering signals with frequencies near the high-frequency band. As for the visuomotor data, the most noticeable impact of the choice of approach was observed when evaluating task-based changes in PAC between the delta (2-5 Hz) and the high-gamma (60-90 Hz) frequency bands, as we were able to identify widespread brain areas with statistically significant effects only with the Hilbert approach. These results provide preliminary evidence of the advantages of the Hilbert approach over the wavelet approach, at least in the context of PAC estimates.

neuroscience↗

Ghost interactions in MEG/EEG source space: A note of caution on inter-areal coupling measures

When combined with source modeling, magneto- (MEG) and electroencephalography (EEG) can be used to study long-range interactions among cortical processes non-invasively. Estimation of such inter-areal connectivity is nevertheless hindered by instantaneous field spread and volume conduction, which artificially introduce linear correlations and impair source separability in cortical current estimates. To overcome the inflating effects of linear source mixing inherent to standard interaction measures, alternative phase- and amplitude-correlation based connectivity measures, such as imaginary coherence and orthogonalized amplitude correlation have been proposed. Being by definition insensitive to zero-lag correlations, these techniques have become increasingly popular in the identification of correlations that cannot be attributed to field spread or volume conduction. We show here, however, that while these measures are immune to the direct effects of linear mixing, they may still reveal large numbers of spurious false positive connections through field spread in the vicinity of true interactions. This fundamental problem affects both region-of-interest-based analyses and all-to-all connectome mappings. Most importantly, beyond defining and illustrating the problem of spurious, or \"ghost\" interactions, we provide a rigorous quantification of this effect through extensive simulations. Additionally, we further show that signal mixing also significantly limits the separability of neuronal phase and amplitude correlations. We conclude that spurious correlations must be carefully considered in connectivity analyses in MEG/EEG source space even when using measures that are immune to zero-lag correlations.\n\nHighlights{checkmark} Reliable estimation of neuronal coupling with MEG and EEG is challenged by signal mixing\n{checkmark}A number of coupling techniques attempt to overcome this limitation by excluding zero-lag interactions\n{checkmark}Contrary to what is commonly admitted, our simulations illustrate that such interaction metrics will still yield false positives\n{checkmark}Spurious, or \"ghost\", interactions are generally detected between sources in the vicinity of true phase-lagged interacting sources\n{checkmark}Signal mixing also severely affects the mutual separability of phase and amplitude correlations

neuroscience↗

Power and shift invariant detection of dynamically coupled networks (PSIICOS) from non-invasive MEG data

MEG reflects electrical activity of neuronal assemblies that coalesce and decoalesce in time providing for massively parallel and dynamic flow of information exchange in the brain. There is a growing evidence that our behavior is mediated by simultaneous activity of several interacting and simultaneously active cortical networks. Not only the location of network nodes but also the temporal profiles of such interaction are of interest. The existing methods are primarily concerned with proposing novel measures of synchrony to be applied to the cortical region activity signals extracted from the MEG data using beamforming or other spatial filtering approaches.\n\nIn this work we propose to render the task of estimating network topology and temporal profiles as a source estimation problem but in the space of interacting topographies. The proposed point of view has a number of benefits as it allows to use the wealth of techniques and intuition accumulated in the community when dealing with source estimation problems. Operating in the interacting topographies space we first propose subspace projection method to significantly reduce the spatial leakage effect in the cross-spectral matrix. We then use subspace matching metrics to extract a set of networks with their synchronicity profiles that explain the variance in the sensor space cross-spectral matrix, much like regular dipoles and their activations explain the variance in the regular evoked responses.\n\nThe preliminary results of application of this method to a simple self-paced voluntary finger movement paradigm comparing allowed to observe functional coupling between primary motor and supplementary motor area and establish that this coupling can be characterized with near-zero phase delay between the oscillatory activity of these two regions. This observed synchrony pattern is physiologically plausible and is in agreement with the current hypothesis about the neural mechanics of the motor acts.

neuroscience↗