, 2001, Goard and Dan, 2009, Haider et al , 2007, Haider and McCo

, 2001, Goard and Dan, 2009, Haider et al., 2007, Haider and McCormick, 2009, Harris and Thiele, 2011, Hasenstaub et al., 2007 and Marguet and Harris, 2011). In this study, we focus on the contributions of motor cortex activity http://www.selleckchem.com/products/incb28060.html to sensory processing in the mouse whisker system. One potentially important pathway for providing contextual signals in the

whisker system is the corticocortical feedback projection from the vibrissal portion of primary motor cortex (vM1) to the vibrissal representation in primary somatosensory cortex (S1) (Miyashita et al., 1994, Porter and White, 1983 and Veinante and Deschênes, 2003). As vM1 neuronal activity correlates with whisking and other task-related parameters (Carvell et al., 1996, Erlich et al., 2011, Friedman et al., 2012, Hill et al., 2011, Huber et al., 2012 and Petreanu et al., 2012), this pathway has been hypothesized to distribute the motor plan throughout the cortical whisker system (Kleinfeld et al., 1999 and Kleinfeld et al., 2006). Recent studies have characterized responses of S1 neurons to vM1 stimulation in vitro (Petreanu et al., 2009 and Rocco and Brumberg, 2007) and in vivo (Lee et al., 2008), demonstrating an

excitatory effect of vM1 inputs most prominently learn more onto infragranular S1 neurons. It is not fully understood, however, how vM1 feedback activity

modulates S1 network dynamics, or how these signals integrate with sensory inputs and contribute to sensory processing. Tolmetin We demonstrate that motor cortex activity can dramatically influence network dynamics in S1, during both whisking and nonwhisking conditions. This modulation of network dynamics is rapid, exhibits target specificity, and is mediated at least in part by the direct corticocortical feedback pathway. Furthermore, we demonstrate that altering the network state directly influences sensory responses and can modulate network response reliability and discrimination. We describe a cortical mechanism that directly links motor cortex activity to changes in somatosensory cortex network state and may enhance representation of sensory inputs during active exploration. We recorded network activity simultaneously from ipsilateral vM1 and S1 in waking mice that had been habituated to head fixation (n = 9 mice; recordings in LV of vM1 and S1). As previously described in S1 recordings (Crochet and Petersen, 2006 and Petersen et al., 2003), we found that network activity in vM1 and S1 was highly variable and correlated with behavioral state (Figures 1A and 1C and Figure S1A available online). When the mice were not whisking, we often observed prominent slow, rhythmic LFP fluctuations at low frequencies (3–5 Hz).

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