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The Journal of Neurophysiology Vol. 85 No. 2 February 2001, pp. 855-868
Copyright ©2001 by the American Physiological Society
1Institute of Physiology, University of Bern, CH-3012 Bern, Switzerland; and 2Abteilung Zellphysiologie, Max-Planck-Institut für Medizinische Forschung, D-69120 Heidelberg, Germany
Berger, Thomas,
Matthew E. Larkum, and
Hans-R. Lüscher.
High Ih Channel Density in the Distal
Apical Dendrite of Layer V Pyramidal Cells Increases Bidirectional
Attenuation of EPSPs. J. Neurophysiol. 85: 855-868, 2001. Despite the wealth of recent research on
active signal propagation along the dendrites of layer V neocortical
pyramidal neurons, there is still little known regarding the traffic of
subthreshold synaptic signals. We present a study using three
simultaneous whole cell recordings on the apical dendrites of these
cells in acute rat brain slices to examine the spread and attenuation
of spontaneous excitatory postsynaptic potentials (sEPSPs). Equal current injections at each of a pair of sites separated by ~500 µm
on the apical dendrite resulted in equal voltage transients at the
other site ("reciprocity"), thus disclosing linear behavior of the
neuron. The mean apparent "length constants" of the apical dendrite
were 273 and 446 µm for somatopetal and somatofugal sEPSPs, respectively. Trains of artificial EPSPs did not show temporal summation. Blockade of the hyperpolarization-activated cation current
(Ih) resulted in less attenuation by
17% for somatopetal and by 47% for somatofugal sEPSPs. A pronounced
location-dependent temporal summation of EPSP trains was seen. The
subcellular distribution and biophysical properties of
Ih were studied in cell-attached patches. Within less than ~400 µm of the soma, a low density of ~3 pA/µm2 was found, which increased to ~40
pA/µm2 in the apical distal dendrite.
Ih showed activation and deactivation kinetics with time constants faster than 40 ms and half-maximal activation at
95 mV. These findings suggest that integration of
synaptic input to the apical tuft and the basal dendrites occurs spatially independently. This is due to a high
Ih channel density in the apical tuft
that increases the electrotonic distance between these two compartments
in comparison to a passive dendrite.
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