Dendritic Hyperpolarization-Activated Currents Modify the Integrative Properties of Hippocampal CA1 Pyramidal Neurons

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Abstract
Step hyperpolarizations evoked slowly activating, noninactivating, and slowly deactivating inward currents from membrane patches recorded in the cell-attached patch configuration from the soma and apical dendrites of hippocampal CA1 pyramidal neurons. The density of these hyperpolarization-activated currents (Ih) increased over sixfold from soma to distal dendrites. Activation curves demonstrate that a significant fraction ofIhchannels is active near rest and that the range is hyperpolarized relatively more in the distal dendrites.Ihactivation and deactivation kinetics are voltage-and temperature-dependent, with time constants of activation and deactivation decreasing with hyperpolarization and depolarization, respectively.Ihdemonstrated a mixed Na+–K+conductance and was sensitive to low concentrations of external CsCl. Dual whole-cell recordings revealed regional differences in input resistance (Rin) and membrane polarization rates (τmem) across the somatodendritic axis that are attributable to the spatial gradient ofIhchannels. As a result of these membrane effects the propagation of subthreshold voltage transients is directionally specific. The elevated dendriticIhdensity decreases EPSP amplitude and duration and reduces the time window over which temporal summation takes place. The backpropagation of action potentials into the dendritic arborization was impacted only slightly by dendriticIh, with the most consistent effect being a decrease in dendritic action potential duration and an increase in afterhyperpolarization. Overall,Ihacts to dampen dendritic excitability, but its largest impact is on the subthreshold range of membrane potentials where the integration of inhibitory and excitatory synaptic inputs takes place.