Explainability of Recurrent Neural Networks for Enhancing P300-based Brain-Computer Interfaces

EEG foundation models such as BIOT, LaBraM, and EEGMamba have achieved remarkable performance in neural signal decoding, but their black-box nature limits clinical trust and neuroscientific validation. We propose a unified attribution framework for interpreting EEG foundation models across heterogeneous architectures. The framework integrates gradient-, perturbation-, and activation-based explanation methods to analyze model behavior in spatial, temporal, and frequency dimensions. Spatially, it identifies critical EEG channels and visualizes their distributions using topographic maps. Temporally, it highlights decision-relevant signal segments through attribution heatmaps. In the frequency domain, it quantifies the contributions of canonical EEG rhythms via spectral perturbation analysis. To assess explanation reliability, we introduce a population-level evaluation combining Area Over the Perturbation Curve (AOPC) and cross-method consistency analysis. The framework further leverages Large Language Models (LLMs) to transform structured attribution outputs into natural-language reports, bridging low-level neural representations and high-level semantic reasoning. Experiments on benchmark datasets, including Mumtaz2016 and TUAB, demonstrate that the generated explanations are consistent with established neurophysiological markers, validating meaningful neural representations while exposing potential dependencies on artifacts and spurious patterns. The proposed framework provides a standardized approach for evaluating the interpretability, reliability, and physiological plausibility of EEG foundation models.

Recent advances in deep learning have enabled increasingly accurate electroencephalography (EEG)-based classification of Major Depressive Disorder (MDD), but the decision-making processes of high-capacity models remain difficult to interpret. This study investigates multiple post-hoc explainability methods applied to an InceptionTime architecture trained for EEG-based MDD detection. The analysis includes Shapley-based, gradient-based, and perturbation-based attribution approaches: DeepSHAP, Integrated Gradients, GradCAM, Occlusion, and Permutation Feature Importance. Explainability analysis was performed within a subject-level stratified 5-fold cross-validation framework using global attribution aggregation across EEG segments and subjects. The evaluated methods revealed partially convergent attribution patterns, with recurring emphasis on frontal, temporal, and posterior EEG regions, particularly in the right hemisphere. Quantitative comparison demonstrated substantial agreement between gradient- and perturbation-based approaches, while DeepSHAP produced comparatively distinct attribution distributions. At the same time, variability between explainability methods highlighted the influence of methodological assumptions on the resulting explanations. Overall, the results suggest that different post-hoc explainability approaches capture partially overlapping relevance structures in EEG-based deep learning models for depression detection. Although the observed attribution patterns are broadly consistent with several previous EEG studies of MDD, the analysis should be interpreted as exploratory rather than evidence of definitive neurophysiological biomarkers or clinical applicability. The study highlights both the usefulness and limitations of post-hoc explainability for interpreting black-box EEG classifiers in psychiatric applications.

Emerging foundation models (FMs) in electroencephalography (EEG) promise a path to scale deep learning in diagnostics and brain-computer interfaces despite data scarcity, yet their opaque nature remains a barrier to wider adoption. We investigate attention-aware Layer-wise relevance propagation (LRP) as a post-hoc attribution method for EEG-FMs, extending LRP's use on convolutional neural network (CNN)-based EEG models to the Transformer architectures that current FMs are based on. We find that LRP can both verify EEG-FM decisions and surface novel, biologically plausible hypotheses from them. In motor imagery, it unmasks 'Clever Hans' behavior where models prioritize task correlated ocular signals over the intended motor correlates. In a naturalistic paradigm for affect prediction, it reveals a recurring reliance on a central electrode cluster, suggesting a candidate sensorimotor signature of arousal. Though heatmap interpretation remains ambiguous in this complex domain, the results position LRP as a tool for both verification and exploration of EEG-FMs, a role that will grow in both importance and discovery potential as the underlying models mature.