Top-down Dendritic Input Increases the Gain of Layer 5 Pyramidal Neurons

The cerebral cortex is organized so that an important component of feedback input from higher to lower cortical areas arrives at the distal apical tufts of pyramidal neurons. Yet, distal inputs are predicted to have much less impact on firing than proximal inputs. Here we show that even weak asynchronous dendritic input to the distal tuft region can significantly increase the gain of layer 5 pyramidal neurons and thereby the output of columns in the primary somatosensory cortex of the rat. Noisy currents injected in ramps at different dendritic locations showed that the initial slope of the frequency-current (f/I) relationship increases with the distance of the current injection from the soma. The increase was due to the interaction of dendritic depolarization with back-propagating APs which activated dendritic calcium conductances. Gain increases were accompanied by a change of firing mode from isolated spikes to bursting where the timing of bursts coded the presence of coincident somatic and dendritic inputs. We propose that this dendritic gain modulation and the timing of bursts may serve to associate top-down and bottom-up input on different time scales.