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The timing and location of action potential initiation in layer V pyramidal cells of the rat somatosensory cortex depends on different ligand-gated and intrinsic conductances. In order to assess the shunting effect of GABA A  receptor activation and  I  h  on the summative properties of these neurons, we recorded simultaneously from their somata and dendrites and induced combined excitatory and inhibitory postsynaptic potentials using extracellular stimulation. Three major consequences of GABA A  receptor activation on the integration properties were found: (i) the action potential could be initiated in the distal apical dendrite, if GABA A  receptors were simultaneously activated in the soma; (ii) GABA A  receptor activation narrowed the time window for suprathreshold summation of two coincident excitatory synaptic inputs by a factor of 3.4; and (iii) uneven weights of the GABAergic components between two inputs could lead to unbalanced shunting and consequently to a time shift in the peak of the coincidence detection window. Blockade of  I  h  practically abolished coincidence detection within a time window of 30 ms. Location of spike initiation and summation of coincident inputs in pyramidal cells is determined by at least two shunting mechanisms: activation of the presynaptic inhibitory network and modulation of  I  h  as a function of the postsynaptic membrane potential.

Timing and Precision of Spike Initiation in Layer V Pyramidal Cells of the Rat Somatosensory Cortex