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In this condition the frequency of the gamma oscillations was reduced and their amplitude increased by raising gI and lowering gE (Figure 3Bb). With a further increase in noise to 300 pA, gamma activity remained entrained to theta cycles, but became less ordered (Figure 3Ca,b). Figure 3. Differential sensitivity of gamma oscillations and grid fields to changes in the strength of E and I synapses. To explore gamma activity across a wider range of gI and gE we automated quantification of the strength and frequency of oscillatory input to E cells (see ��Materials and methods��). In the absence of noise gamma frequency activity only occurred for a narrow range of gI and gE (Figure 3D). Strikingly, following addition of moderate noise the region of parameter space that supports gamma activity was massively expanded (Figure 3E). Within this space, the amplitude of gamma increased with increasing inhibition, whereas the frequency Carfilzomib mw was reduced. As noise is increased further the amplitude and frequency of gamma oscillations Selleck VX-809 are reduced (Figure 3F). We found a similar dependence of gamma oscillations on noise, gE and gI in networks with probabilistic connectivity (Figure 3��figure supplement 1). Thus, intrinsic noise modifies the amplitude and frequency of nested gamma oscillations. To determine whether there is a systematic relationship between values of gE and gI that generate gamma and grid firing we compared the gridness score and gamma scores for each circuit configuration (Figure 3G, Figure 3��figure supplements 2, 3). We found this relationship to be complex and highly sensitive to noise. However, we did not find any evidence for strong linear relationships between gamma amplitude or gamma frequency and grid score (R2 Cefaloridine and amplitude of gamma oscillations varied smoothly as a function of noise (Figure 3H). Thus, neither the frequency nor the power of gamma appears to be a good predictor of grid firing. When we considered only regions of parameter space that generate robust grid fields (grid score >0.5), we found circuits generating almost the complete observed range of gamma amplitudes (0.02