Deep learning-based EEG analysis: investigating P3 ERP components

The neural processing of incoming stimuli can be analysed from the electroencephalogram (EEG) through event-related potentials (ERPs). The P3 component is largely investigated as it represents an important psychophysiological marker of psychiatric disorders. This is composed by several subcomponents, such as P3a and P3b, reflecting distinct but interrelated sensory and cognitive processes of incoming stimuli. Due to the low EEG signal-to-noise-ratio, ERPs emerge only after an averaging procedure across trials and subjects. Thus, this canonical ERP analysis lacks in the ability to highlight EEG neural signatures at the level of single-subject and single-trial. In this study, a deep learning-based workflow is investigated to enhance EEG neural signatures related to P3 subcomponents already at single-subject and at single-trial level. This was based on the combination of a convolutional neural network (CNN) with an explanation technique (ET). The CNN was trained using two different strategies to produce saliency representations enhancing signatures shared across subjects or more specific for each subject and trial. Cross-subject saliency representations matched the signatures already emerging from ERPs, i.e., P3a and P3b-related activity within 350–400 ms (frontal sites) and 400–650 ms (parietal sites) post-stimulus, validating the CNN+ET respect to canonical ERP analysis. Single-subject and single-trial saliency representations enhanced P3 signatures already at the single-trial scale, while EEG-derived representations at single-subject and single-trial level provided no or only mildly evident signatures. Empowering the analysis of P3 modulations at single-subject and at single-trial level, CNN+ET could be useful to provide insights about neural processes linking sensory stimulation, cognition and behaviour

The Sensory-Cognitive Interplay: Insights into Neural Mechanisms and Circuits

Senses are our interface for acting in the external world. Consequently, sensory-motor information grounds and drives our higher cognitive processes. At the same time, we are impinged by a multitude of sensory inputs with variable saliency. It is therefore crucial that the process- ing of sensory inputs and motor signals is modulated by cognitive and executive mechanisms such as expectation, memory, attention, emotion, planning, monitoring. This is needed to highlight sensory information that is currently rel- evant for task goals, and to adapt motor control and behav- ior accordingly. The strict intertwining of sensory, motor, and cognitive functions is evidenced in aging and in neuro- logical disorders. Indeed, sensory-motor dysfunctions of- ten accompany higher-level dysfunctions in older popula- tions [1] and in neurological subjects (e.g., in dyslexia, at- tention deficit hyperactivity disorders, or autism spectrum disorders) [2,3] [...

A Semantic Model to Study Neural Organization of Language in Bilingualism

A neural network model of object semantic representation is used to simulate learning of new words from a foreign language. The network consists of feature areas, devoted to description of object properties, and a lexical area, devoted to words representation. Neurons in the feature areas are implemented as Wilson-Cowan oscillators, to allow segmentation of different simultaneous objects via gamma-band synchronization. Excitatory synapses among neurons in the feature and lexical areas are learned, during a training phase, via a Hebbian rule. In this work, we first assume that some words in the first language (L1) and the corresponding object representations are initially learned during a preliminary training phase. Subsequently, second-language (L2) words are learned by simultaneously presenting the new word together with the L1 one. A competitive mechanism between the two words is also implemented by the use of inhibitory interneurons. Simulations show that, after a weak training, the L2 word allows retrieval of the object properties but requires engagement of the first language. Conversely, after a prolonged training, the L2 word becomes able to retrieve object per se. In this case, a conflict between words can occur, requiring a higher-level decision mechanism

A Lightweight Multi-Scale Convolutional Neural Network for P300 Decoding: Analysis of Training Strategies and Uncovering of Network Decision

Convolutional neural networks (CNNs), which automatically learn features from raw data to approximate functions, are being increasingly applied to the end-to-end analysis of electroencephalographic (EEG) signals, especially for decoding brain states in brain-computer interfaces (BCIs). Nevertheless, CNNs introduce a large number of trainable parameters, may require long training times, and lack in interpretability of learned features. The aim of this study is to propose a CNN design for P300 decoding with emphasis on its lightweight design while guaranteeing high performance, on the effects of different training strategies, and on the use of post-hoc techniques to explain network decisions. The proposed design, named MS-EEGNet, learned temporal features in two different timescales (i.e., multi-scale, MS) in an efficient and optimized (in terms of trainable parameters) way, and was validated on three P300 datasets. The CNN was trained using different strategies (within-participant and within-session, within-participant and cross-session, leave-one-subject-out, transfer learning) and was compared with several state-of-the-art (SOA) algorithms. Furthermore, variants of the baseline MS-EEGNet were analyzed to evaluate the impact of different hyper-parameters on performance. Lastly, saliency maps were used to derive representations of the relevant spatio-temporal features that drove CNN decisions. MS-EEGNet was the lightest CNN compared with the tested SOA CNNs, despite its multiple timescales, and significantly outperformed the SOA algorithms. Post-hoc hyper-parameter analysis confirmed the benefits of the innovative aspects of MS-EEGNet. Furthermore, MS-EEGNet did benefit from transfer learning, especially using a low number of training examples, suggesting that the proposed approach could be used in BCIs to accurately decode the P300 event while reducing calibration times. Representations derived from the saliency maps matched the P300 spatio-temporal distribution, further validating the proposed decoding approach. This study, by specifically addressing the aspects of lightweight design, transfer learning, and interpretability, can contribute to advance the development of deep learning algorithms for P300-based BCIs

Alpha and theta mechanisms operating in internal-external attention competition

Attention is the ability to prioritize a set of information at expense of others and can be internally- or externally-oriented. Alpha and theta oscillations have been extensively implicated in attention. However, it is unclear how these oscillations operate when sensory distractors are presented continuously during task-relevant internal processes, in close-to-real-life conditions. Here, EEG signals from healthy participants were obtained at rest and in three attentional conditions, characterized by the execution of a mental math task (internal attention), presentation of pictures on a monitor (external attention), and task execution under the distracting action of picture presentation (internal-external competition). Alpha and theta power were investigated at scalp level and at some cortical regions of interest (ROIs); moreover, functional directed connectivity was estimated via spectral Granger Causality. Results show that frontal midline theta was distinctive of mental task execution and was more prominent during competition compared to internal attention alone, possibly reflecting higher executive control; anterior cingulate cortex appeared as mainly involved and causally connected to distant (temporal/ occipital) regions. Alpha power in visual ROIs strongly decreased in external attention alone, while it assumed values close to rest during competition, reflecting reduced visual engagement against distractors; connectivity results suggested that bidirectional alpha influences between frontal and visual regions could contribute to reduce visual interference in internal attention. This study can help to understand how our brain copes with internal-external attention competition, a condition intrinsic in the human sensory-cognitive interplay, and to elucidate the relationships between brain oscillations and attentional functions/dysfunctions in daily tasks

Modulations of Cortical Power and Connectivity in Alpha and Beta Bands during the Preparation of Reaching Movements

Planning goal-directed movements towards different targets is at the basis of common daily activities (e.g., reaching), involving visual, visuomotor, and sensorimotor brain areas. Alpha (8-13 Hz) and beta (13-30 Hz) oscillations are modulated during movement preparation and are implicated in correct motor functioning. However, how brain regions activate and interact during reaching tasks and how brain rhythms are functionally involved in these interactions is still limitedly explored. Here, alpha and beta brain activity and connectivity during reaching preparation are investigated at EEG-source level, considering a network of task-related cortical areas. Sixty-channel EEG was recorded from 20 healthy participants during a delayed center-out reaching task and projected to the cortex to extract the activity of 8 cortical regions per hemisphere (2 occipital, 2 parietal, 3 peri-central, 1 frontal). Then, we analyzed event-related spectral perturbations and directed connectivity, computed via spectral Granger causality and summarized using graph theory centrality indices (in degree, out degree). Results suggest that alpha and beta oscillations are functionally involved in the preparation of reaching in different ways, with the former mediating the inhibition of the ipsilateral sensorimotor areas and disinhibition of visual areas, and the latter coordinating disinhibition of the contralateral sensorimotor and visuomotor areas

EEG alpha power is modulated by attentional changes during cognitive tasks and virtual reality immersion

Variations in alpha rhythm have a significant role in perception and attention. Recently, alpha decrease has been associated with externally directed attention, especially in the visual domain, whereas alpha increase has been related to internal processing such as mental arithmetic. However, the role of alpha oscillations and how the different components of a task (processing of external stimuli, internal manipulation/representation, and task demand) interact to affect alpha power are still unclear. Here, we investigate how alpha power is differently modulated by attentional tasks depending both on task difficulty (less/more demanding task) and direction of attention (internal/external). To this aim, we designed two experiments that differently manipulated these aspects. Experiment 1, outside Virtual Reality (VR), involved two tasks both requiring internal and external attentional components (intake of visual items for their internal manipulation) but with different internal task demands (arithmetic vs. reading). Experiment 2 took advantage of the VR (mimicking an aircraft cabin interior) to manipulate attention direction: it included a condition of VR immersion only, characterized by visual external attention, and a condition of a purely mental arithmetic task during VR immersion, requiring neglect of sensory stimuli. Results show that: (1) In line with previous studies, visual external attention caused a significant alpha decrease, especially in parieto-occipital regions; (2) Alpha decrease was significantly larger during the more demanding arithmetic task, when the task was driven by external visual stimuli; (3) Alpha dramatically increased during the purely mental task in VR immersion, whereby the external stimuli had no relation with the task. Our results suggest that alpha power is crucial to isolate a subject from the environment, and move attention from external to internal cues. Moreover, they emphasize that the emerging use of VR associated with EEG may have important implications to study brain rhythms and support the design of artificial systems

An Emergent Model of Multisensory Integration in Superior Colliculus Neurons

Neurons in the cat superior colliculus (SC) integrate information from different senses to enhance their responses to cross-modal stimuli. These multisensory SC neurons receive multiple converging unisensory inputs from many sources; those received from association cortex are critical for the manifestation of multisensory integration. The mechanisms underlying this characteristic property of SC neurons are not completely understood, but can be clarified with the use of mathematical models and computer simulations. Thus the objective of the current effort was to present a plausible model that can explain the main physiological features of multisensory integration based on the current neurological literature regarding the influences received by SC from cortical and subcortical sources. The model assumes the presence of competitive mechanisms between inputs, nonlinearities in NMDA receptor responses, and provides a priori synaptic weights to mimic the normal responses of SC neurons. As a result, it provides a basis for understanding the dependence of multisensory enhancement on an intact association cortex, and simulates the changes in the SC response that occur during NMDA receptor blockade. Finally, it makes testable predictions about why significant response differences are obtained in multisensory SC neurons when they are confronted with pairs of cross-modal and within-modal stimuli. By postulating plausible biological mechanisms to complement those that are already known, the model provides a basis for understanding how SC neurons are capable of engaging in this remarkable process

Bottom-up vs. top-down connectivity imbalance in individuals with high-autistic traits: An electroencephalographic study

Brain connectivity is often altered in autism spectrum disorder (ASD). However, there is little consensus on the nature of these alterations, with studies pointing to either increased or decreased connectivity strength across the broad autism spectrum. An important confound in the interpretation of these contradictory results is the lack of information about the directionality of the tested connections. Here, we aimed at disambiguating these confounds by measuring differences in directed connectivity using EEG resting-state recordings in individuals with low and high autistic traits. Brain connectivity was estimated using temporal Granger Causality applied to cortical signals reconstructed from EEG. Between-group differences were summarized using centrality indices taken from graph theory (in degree, out degree, authority, and hubness). Results demonstrate that individuals with higher autistic traits exhibited a significant increase in authority and in degree in frontal regions involved in high-level mechanisms (emotional regulation, decision-making, and social cognition), suggesting that anterior areas mostly receive information from more posterior areas. Moreover, the same individuals exhibited a significant increase in the hubness and out degree over occipital regions (especially the left and right pericalcarine regions, where the primary visual cortex is located), suggesting that these areas mostly send information to more anterior regions. Hubness and authority appeared to be more sensitive indices than the in degree and out degree. The observed brain connectivity differences suggest that, in individual with higher autistic traits, bottom-up signaling overcomes top-down channeled flow. This imbalance may contribute to some behavioral alterations observed in ASD

Mathematical modeling and parameter estimation of levodopa motor response in patients with Parkinson disease

Parkinson disease (PD) is characterized by a clear beneficial motor response to levodopa (LD) treatment. However, with disease progression and longer LD exposure, drug-related motor fluctuations usually occur. Recognition of the individual relationship between LD concentration and its effect may be difficult, due to the complexity and variability of the mechanisms involved. This work proposes an innovative procedure for the automatic estimation of LD pharmacokinetics and pharmacodynamics parameters, by a biologically-inspired mathematical model. An original issue, compared with previous similar studies, is that the model comprises not only a compartmental description of LD pharmacokinetics in plasma and its effect on the striatal neurons, but also a neurocomputational model of basal ganglia action selection. Parameter estimation was achieved on 26 patients (13 with stable and 13 with fluctuating LD response) to mimic plasma LD concentration and alternate finger tapping frequency along four hours after LD administration, automatically minimizing a cost function of the difference between simulated and clinical data points. Results show that individual data can be satisfactorily simulated in all patients and that significant differences exist in the estimated parameters between the two groups. Specifically, the drug removal rate from the effect compartment, and the Hill coefficient of the concentration-effect relationship were significantly higher in the fluctuating than in the stable group. The model, with individualized parameters, may be used to reach a deeper comprehension of the PD mechanisms, mimic the effect of medication, and, based on the predicted neural responses, plan the correct management and design innovative therapeutic procedures