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The Journal of Neuroscience, August 15, 2002, 22(16):6991-7005
Signaling of Layer 1 and Whisker-Evoked Ca2+ and
Na+ Action Potentials in Distal and Terminal Dendrites of
Rat Neocortical Pyramidal Neurons In Vitro and In
Vivo
Matthew E.
Larkum1 and
J. Julius
Zhu1, 2, 3
1 Department of Cell Physiology, Max Planck Institute
for Medical Research, Heidelberg D-69120, Germany, 2 Cold
Spring Harbor Laboratory, Cold Spring Harbor, New York 11724, and
3 Department of Pharmacology, University of Virginia School
of Medicine, Charlottesville, Virginia 22908
Dendritic regenerative potentials play an important role in
integrating and amplifying synaptic inputs. To understand how distal
synaptic inputs are integrated and amplified, we made multiple simultaneous (double, triple, or quadruple) and sequential (4-12 paired) recordings from different locations of single tufted layer 5 pyramidal neurons in the cortex in vitro and studied the
spatial and temporal properties of their dendritic regenerative
potential initial zone. Recordings from the soma and from trunk,
primary, secondary, tertiary, and quaternary tuft branches of the
apical dendrite of these neurons reveal a spatially restricted
low-threshold zone ~550-900 µm from the soma for
Ca2+-dependent regenerative potentials. Dendritic
regenerative potentials initiated in this zone have a clearly defined
threshold and a refractory period, and they can propagate actively
along the dendrite before evoking somatic action potentials. The
detailed biophysical characterization of this dendritic action
potential initiation zone allowed for the further investigation of
dendritic potentials in the intact brain and their roles in information
processing. By making whole-cell recordings from the soma and varied
locations along the apical dendrite of 53 morphologically identified
layer 5 pyramidal neurons in anesthetized rats, we found that three of
the dendritic potentials characterized in vitro could be
induced by spontaneous or whisker inputs in vivo. Thus
layer 5 pyramidal neurons of the rat neocortex have a spatially
restricted low-threshold zone in the apical dendrite, the activation or
interaction of which with the axonal action potential initiation zone
is responsible for multiple forms of regenerative potentials critical
for integrating and amplifying sensory and modulatory inputs.
Key words:
rat; somatosensory; excitation; inhibition; synaptic
integration; development; attention; synaptic plasticity
Copyright © 2002 Society for Neuroscience 0270-6474/02/22166991-15$05.00/0
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