, 2009; Karlsson

and Frank, 2009). CA3-CA1 gamma coherence varied as a function of replay quality, with high quality (higher R2; Figure 7A) and more significant (lower p value; Figure 7B) replay events displaying the strongest levels of gamma coherence following SWR detection. The magnitude and the duration of gamma coherence appeared to decrease for lower quality (lower R2) and less significant replay events (higher p value). We then compared CA3-CA1 gamma coherence for significant (p < 0.05; n = 454 SWRs) and nonsignificant (p > 0.05; n = 477 SWRs) candidate SWRs. Gamma coherence was significantly greater for significant as compared to nonsignificant candidate SWRs for the 50–300 ms following Autophagy inhibitor SWR detection (Figure 7C; permutation test; significant > nonsignificant p < 0.001). Similarly, increases in CA3-CA1 gamma phase locking were predictive of the quality of memory replay. Highly significant replay events showed the largest increase in phase locking for the longest duration. In contrast, less sequential and nonsignificant candidate events showed the smallest increase in phase locking for the shortest duration (Figures

7D and 7E). CA3-CA1 gamma phase locking was significantly different for significant and nonsignificant candidate SWRs for the 50–250 ms after SWR detection (Figure 7F; permutation test; significant selleck chemical > nonsignificant p < 0.001). The increase in gamma coherence and phase locking observed for significant as compared to nonsignificant SWRs persisted when

we controlled for gamma power, SWR magnitude, SWR duration, the number of spikes in each event and the number of cells participating in each event (Figure S7). Finally, we noted that awake replay has been reported both during SWRs (Foster and Wilson, 2006; Diba and Buzsáki, 2007; Davidson et al., 2009; Karlsson and Frank, 2009; Gupta et al., 2010) and during periods associated with theta rhythmicity (Johnson and Redish, 2007), which occurs during attentive behaviors and movement. We therefore asked whether the events we examined included a subset with high theta power, as might be expected if there were two distinct types of replay events. As theta is thought to reflect a relatively long duration state of hippocampal information processing and the sharp-wave in each SWR has power in the Parvulin 6–12 Hz theta band (Buzsáki, 1986), we examined theta power during the 400 ms before SWR detection. There was a unimodal distribution of theta power during this period, suggesting that all of the events we examined occurred in a similar network state (Figure S7). Finally, neither theta power nor theta coherence in the 400 ms before each SWR was related to replay quality (Spearman correlation; replay p value versus theta power; CA1: ρ = −0.06, p > 0.1; CA3: ρ = 0.01, p > 0.5; replay p value versus theta coherence; ρ = −0.03, p > 0.4).