In plots of power spectral density (Figure 2C), each rat had peak

In plots of power spectral density (Figure 2C), each rat had peak BG beta frequency slightly below 20 Hz (range: 17.9–19.5 Hz) with cortical frequency consistently a touch higher (18.4 Hz

to 20.4 Hz). If beta oscillations represent a distinct, network-wide coordinated BG state, this should CH5424802 in vivo be apparent in analyses of phase and power relationships between structures. Coherence between all BG structures consistently showed a peak at ∼20 Hz for all rats (Figures 2D and Figures S3B). By contrast, we have previously shown that coherence between striatum and dorsal hippocampus in behaving rats is close to zero at 20 Hz (Berke et al., 2004 and Berke, 2009). We next constructed comodulograms, which illustrate the extent to which moment-to-moment oscillatory power covaries between structures (Buzsáki et al., 2003). Coordinated power changes within the BG network were observed especially at ∼20 Hz (Figures 2D and Figures S3B).

Modulation of beta power relative to behavioral events was essentially identical throughout the BG, and similar between BG and ECoG (Figure S2B). There was no consistent difference in the modulation of beta power for ipsilateral RG7204 order versus contralateral movements (Figure S4). We have previously reported that striatal LFPs show mutually exclusive dynamic states, characterized by combinations of either ∼20 Hz beta and ∼50 Hz low-gamma rhythms, or ∼8 Hz theta and ∼80 Hz high-gamma rhythms, respectively (Berke, 2009; see also Dejean et al., 2011). These distinct states were visible in our current comodulograms: in STR, GP, and STN ∼50 Hz power was positively correlated with beta power SB-3CT and negatively correlated with ∼80 Hz activity. These relationships were absent or diminished for SNr. BG beta rhythms were tightly coordinated between structures, but not identical in all respects. We consistently observed a significant difference in beta phase between simultaneously recorded subregions (28 pairwise comparisons, p < 0.05 in every case; see Experimental Procedures).

Although the specific set of recording regions varied between subjects, for all four rats we were able to compare beta phases between frontal ECoG, STR, and GP (Figure 2E). STR beta was always phase-advanced relative to the ECoG, (by an average of 97°), and GP was always slightly phase-advanced relative to the striatum (by an average of 4.8°). These results rule out some nonphysiological explanations for coordinated beta rhythms throughout the BG—for example, if the beta oscillations were on the common reference electrode, they would show no phase shift across regions. However, phase differences do not necessarily indicate where an ERS/ERD occurs first, especially as beta has a different phase at the cortical surface compared to deep layers (Murthy and Fetz, 1992).

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