Physiology of Layer 5 Pyramidal Neurons in Mouse Primary Visual Cortex: Coincidence Detection through Bursting

L5 pyramidal neurons are the only neocortical cell type with dendrites reaching all six layers of cortex, casting them as one of the main integrators in the cortical column. What is the nature and mode of computation performed in mouse primary visual cortex (V1) given the physiology of L5 pyramidal neurons? First, we experimentally establish active properties of the dendrites of L5 pyramidal neurons of mouse V1 using patch-clamp recordings. Using a detailed multi-compartmental model, we show this physiological setup to be well suited for coincidence detection between basal and apical tuft inputs by controlling the frequency of spike output. We further show how direct inhibition of calcium channels in the dendrites modulates such coincidence detection. To establish the singe-cell computation that this biophysics supports, we show that the combination of frequency-modulation of somatic output by tuft input and (simulated) calcium-channel blockage functionally acts as a composite sigmoidal function. Finally, we explore how this computation provides a mechanism whereby dendritic spiking contributes to orientation tuning in pyramidal neurons.Author Summary: Neurons in the brain have elaborate dendritic morphologies, hosting a variety of nonlinear channels that give way to single cell computation. In this study, we perform patch clamp recordings in the apical dendrites to establish the spatial distribution of nonlinear channels and the signals they support in the dendrites of layer 5 pyramidal neurons of the mouse primary visual cortex. Using this data, we create a detailed single cell model and simulate synaptic input. We then summarize the results of the simulations using a simple abstracted model, that ultimately describes the computation layer 5 pyramidal neurons perform on synaptic input. We find that this computation is a form of nonlinear frequency-modulation that works in a dendritic-spike dependent manner. Finally, we show how this computation allows dendritic spikes to contribute to the orientation tuning of pyramidal neurons in the visual cortex.