We employed Granger causality (GC) to characterize causal interactions among LFP time series from the recorded regions. GC is based on the Wiener-Granger concept of causality61. First, if knowledge of the past of a time series A improves the prediction of the future of another time series B better than what could be accomplished by knowing the past of B alone, then signal A “Granger-causes” signal B. This statistical concept is increasingly popular in the neuroscience community and offers a directional measure of neural functional connectivity. We used the multivariate Granger causality (MVGC) Matlab toolbox provided by Barnett and Seth44 to calculate the multivariate conditional GC in the frequency domain over time, which is more appropriate for neural time series.
First, we fit a multivariate autoregressive (MVAR) model to the LFP data from the 4 regions studied in all trials, using the ordinary least squares method with model order estimated by the Akaike Information Criterion (limited to 20). Next, we obtained a MVAR model and calculated the autocovariance sequence for each sliding window (150 ms length, 10 ms step), which was then used to calculate the pairwise-conditional frequency-domain MVGC. The GC results was standard scored by the baseline (−3 s to −1 s) of each frequency. We considered as significant values those higher than baseline mean plus two times its standard deviation. The spectral GC maps were smoothed with a Gaussian filter (mask size = [20 70], standard deviation sigma = 30).
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