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Layer V pyramidal neuron - Dendrites sample the entire cortical column. - Axons project locally, and distally to basal ganglia, brainstem, & spinal cord. - Receives local inputs (from the same cortical column) and distal inputs (other cortical areas and thalamus). From Lübke et al 1996 Excitatory connections between layer V pyramidal cells Full connectivity predicted from morphology vs. Sparse (10 %) functional connectivity Kalisman et al 2005 Touch # axo-dendritic touches The solution: All-to-All Potential Connectivity Unconnected Whole-cell recordings & 6 Connected Confocal connection analysis 5 4 3 2 1 Bouton # boutons 0 3 2 1 0 Connections between interneurons & pyramidal neurons Summary - An inhibitory feedback loop is described for layer V pyramidal cells. - The loop is frequency dependent, and is more prevalent than monosynaptic excitation. - It is mediated by a facilitating excitatory connection onto GABAergic inhibitory neurons. - The inhibition is targeted to the dendrites of pyramidal neurons (low reversal & dendritic patch experiments). - Multi-patch recordings of PCs and Interneurons revealed that the loop was mediated by dendritic-targeting Martinotti cells.