Working memory (WM) is an essential cognitive function. Manipulating WM load and observing corresponding brain responses is a commonly applied technique for uncovering the neural mechanisms of WM. In this study, we applied multivariate pattern analysis (MVPA) to two high-density EEG datasets from healthy human volunteers performing verbal WM tasks with three different levels of memory load to examine the formation and development of WM representations in the brain. By testing at which moment WM-specific neural representations become decodable from scalp EEG, we characterized the temporal dynamics of WM representations. Moreover, we observed evidence for dynamic coding of the memory-specific information as evidenced by the current source density (CSD)-based decoding. Critically, we observed evidence suggesting that WM could be maintained in the format of an activity-silent neural state via the activity-silent synaptic mechanisms. Using CSD connectivity-based decoding, we could decode the neural representation about the contents in WM from the so-called activity-silent period. It is quite remarkable that the patterned hidden state in WM networks can be detected at the scalp level using whole-head EEG CSD functional connectivity. This provides important proof-of-principle evidence for the feasibility of exploring hidden neural states with scalp EEG, with important implications for WM.
High-density EEG data was recorded while the subject was performing two working memory Experiments. (A) Structure of a trial of Experiment 1. (B) Time periods of interest including encoding, retention, and retrieval for Experiment 1. (C) Structure of Experiment 2. (D) Time periods of interest for Experiment 2.
Decoding accuracy time courses averaged across subjects. (A) CSD based decoding and (B) CSD connectivity based decoding (only load 2 vs. load 6 is depicted) for Experiment 1. (C) CSD based decoding and (D) CSD connectivity based decoding (only load 1 vs. load 5 is depicted) for Experiment 2. The horizontal dark line marks chance classification accuracy at 50%. The shaded area is the standard error of the mean across subjects. Colored lines above the time axis indicate clusters where decoding performance is significantly higher than chance.
Temporal generalization of CSD value-based decoding performance. CSD-based decoding generalization accuracy matrix averaged over subjects for Experiment 1 (A) and Experiment 2 (C). Binary map of time-point pairs where the CSD-based decoding generalization was significantly above chance level (red area) for Experiment 1 (B) and Experiment 2 (D) (Wilcoxon signed rank test, controlled for multiple comparisons using FDR).
Temporal generalization of CSD connectivity-based decoding performance. CSD connectivity-based decoding generalization accuracy matrix averaged over subjects for Experiment 1 (A) and Experiment 2 (C). Binary map of time point pairs where the CSD connectivity-based decoding generalization was significantly above chance level (red area) for Experiment 1 (B) and Experiment 2 (D) (Wilcoxon signed rank test, controlled for multiple comparisons using FDR).