Types of dendritic inhibition. (A) Ionotropic, GABAA inhibition shunts the dendrites and spines via conductance of Cl− and HCO3 ions. GABAB receptors on the other hand, which operate metabotropically, have a range of actions and locations. Presynaptically, one isoform of the receptor tends to block Ca2+ channels responsible for triggering the release mechanism. Postsynaptically, a different isoform of the GABAB receptor opens G-protein activated inwardly rectifying K+ channels and blocks Ca2+ and NMDA channels that control dendritic electrogenesis. GABAB receptors are found extrasynaptically where they are involved in “volume transmission” and tonic inhibition. (B) Martinotti neurons have been shown to mediate disynaptic inhibition between neocortical pyramidal neurons. Their apical dendrite targets the dendritic initiation zones of nearby pyramidal neurons (Silberberg and Markram, 2007). (C) Top, repetitive activation of Martinotti neurons causes brief and small hyperpolarizing potentials in the dendrites of pyramidal neurons but Ca2+ spikes generated by local dendritic depolarization (middle) are still powerfully blocked by GABAA-mediated dendritic inhibition (bottom) because of the profound block of the underlying mechanisms for Ca2+ spikes (Murayama et al., 2009). (D) Late-spiking neurogliaform neurons of layer 1 also target the dendrites of cortical pyramidal neurons (Chu et al., 2003). (E) Neurogliaform cells mediate a large fraction of their inhibitory action on the dendrites through GABAB receptors (Oláh et al., 2007).