In a subset of experiments (n = 8), we used red-fluorescent muscimol Vorinostat manufacturer to monitor the extent of muscimol diffusion. Postmortem, in all cases we found that muscimol diffusion remained restricted to the infragranular layers (6 mice, see Figure 6C and the fluorescence intensity profiles along the depth of the cortex in Figure S6). These combined data argue that there were no direct effects of muscimol in the supragranular layers after deeplayer injection, and thus that we were able to selectively inhibit the infragranular layers. Infragranular layer blockade with both normal and

fluorescent muscimol abolished SHs in overlying L2/3Ps (Figure 6E; n = 16, 14 mice; −3.5 ± 0.3 versus 0.3 ± 0.7 mV, p < 0.001; data from animals injected with normal and fluorescent muscimol

were cumulated as they were statistically undistinguishable: 0.3 ± 1.2 versus 0.4 ± 0.8 mV; p = 0.9). Thus, both local GABA blockade and silencing of layer 5 effectively counteracted SHs in V1 L2/3Ps. Overall, the data argue that translaminar (infragranular to supragranular) inhibition is important for the generation of SHs in L2/3Ps of V1. What is the impact of sound-driven IPSPs on sub- and suprathreshold visual responses of V1 neurons? Based on the observed latency of SHs, we presented the noise burst so that the SH peak would coincide with the peak of the synaptic visual response evoked by optimally oriented moving bars (Figure 7A). Combining the auditory and visual stimulation Selleck BMS777607 in this way significantly reduced the amplitude of visually driven depolarizations (Figure 7B; n = 9, 5 mice; 14.4 ± 1.8 versus 9.7 ± 1.7 mV, p < 0.001). Combined

auditory and visual stimulation also reduced action potential (AP) responses compared to pure visual stimulation, in terms of both peak and total number of spikes per stimulus (Figure 7B; medians: 6.6 versus 1.2 Hz and 0.48 versus 0.05 APs, respectively; p < 0.05). Moreover, bimodal stimulation reduced the reliability Levetiracetam of visually driven spiking, as indicated by an increase of the coefficient of variation for APs counts on single trials (Figure 7B; medians: 1.74 versus 2.71, p < 0.05). Based on these results, one could expect that a noise burst would degrade visual perception. We tested this prediction by comparing the behavioral response to a simple visual stimulus presented alone or with a simultaneous noise burst (Figure 8A). Mice were first conditioned by pairing the visual stimulus (50 ms flash, 25% luminance change) with an electric foot-shock occurring 250 ms later. This caused the emergence of a visually driven conditioned motor response (V-CMR). V-CMR was expressed as the normalized peak of locomotor activity, measured around the expected time of the electrical shock (200–400 ms, see Supplemental Experimental Procedures).