Simultaneous EEG-fMRI reveals a temporal cascade of task-related and default-mode activations during a simple target detection task.
- Authors
- Walz, Jennifer M; Goldman, Robin I; Carapezza, Michael; Muraskin, Jordan; Brown, Truman R; Sajda, Paul
- Year
- 2014
- Journal
- NeuroImage
- PMID
- 23962956
- DOI
- 10.1016/j.neuroimage.2013.08.014
- PMCID
- PMC3926909
Focused attention continuously and inevitably fluctuates, and to completely understand the mechanisms responsible for these modulations it is necessary to localize the brain regions involved. During a simple visual oddball task, neural responses measured by electroencephalography (EEG) modulate primarily with attention, but source localization of the correlates is a challenge. In this study we use single-trial analysis of simultaneously-acquired scalp EEG and functional magnetic resonance image (fMRI) data to investigate the blood oxygen level dependent (BOLD) correlates of modulations in task-related attention, and we unravel the temporal cascade of these transient activations. We hypothesize that activity in brain regions associated with various task-related cognitive processes modulates with attention, and that their involvements occur transiently in a specific order. We analyze the fMRI BOLD signal by first regressing out the variance linked to observed stimulus and behavioral events. We then correlate the residual variance with the trial-to-trial variation of EEG discriminating components for identical stimuli, estimated at a sequence of times during a trial. Post-stimulus and early in the trial, we find activations in right-lateralized frontal regions and lateral occipital cortex, areas that are often linked to task-dependent processes, such as attentional orienting, and decision certainty. After the behavioral response we see correlates in areas often associated with the default-mode network and introspective processing, including precuneus, angular gyri, and posterior cingulate cortex. Our results demonstrate that during simple tasks both task-dependent and default-mode networks are transiently engaged, with a distinct temporal ordering and millisecond timescale.
Method for constructing fMRI regressors from simultaneously-acquired EEG dataA. For each trial (i), select a training window of EEG data (Xi) with offset τ from the stimulus (or behavioral response). B. Train linear classifier on EEG data within the time window to estimate a set of spatial weights (w) that maximize discrimination of the two conditions (shown using only 2 EEG channels for visualization purposes). C. In addition to traditional event-related average response (ERAR) and reaction time (RT) regressors, construct single-trial EEG variability (STV) regressors by modulating boxcar height with classifier output (y) for each trial. D. This technique is repeated for multiple window offsets spanning the epoch to view temporal progression of discriminating components spanning the trial.
Stimulus-locked and response-locked ERPs recorded at the Pz electrodeOur definition of early, middle, and late window ranges (see main text for discussion) are indicated with shading.
Group mean averages and standard errors of single-trial EEG discrimination performanceResults of both the stimulus-locked (blue) and response-locked (green) analyses are shown, aligned by mean RT. Since we are interested in the BOLD correlates of single-trial EEG variability, we only consider EEG components with discrimination that is both significant (Az > 0.66, p < 0.01) and substantial (Az > 0.75). Windows resulting in significant positive and negative BOLD correlations are indicated with magenta diamonds and orange circles, respectively. Early, middle, and late windows (as grouped for discussion) are indicated with shading.
Significant clusters correlating with EEG single-trial variability early in the trialShown for stimulus-locked (SL, top) and response-locked (RL, bottom) windows. Corresponding EEG discriminant component scalp projections (forward models) are also shown.
Significant clusters correlating with EEG single-trial variability in middle windowsThese activations occurred near the behavioral response time and in the range of the P3, shown for stimulus-locked (SL, top) and response-locked (RL, bottom) windows. Corresponding EEG discriminant component scalp projections (forward models) are also shown.
Significant clusters correlating with EEG single-trial variability late in the trialThese activations occurred after the subject has made his/her response, for stimulus-locked (SL, top) and response-locked (RL, bottom) windows. Corresponding EEG discriminant component scalp projections (forward models) are also shown.
DMN determined independently with EEG single-trial variability and ICA525 ms stimulus-locked window EEG single-trial variability (STV) negative correlates in medial frontal gyrus, bilateral angular gyri, and posterior cingulate (in blue, with hue representing p-value in the range 0.005−0.001), overlaid with the mean default-mode network (DMN) as determined using ICA (pink). STV results shown are multiple-comparison corrected at p < 0.05. An additional cluster in right middle frontal gyrus can also be seen.
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