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First published online on April 11, 2003.
Copyright © 2003 by The Physiological Society
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Received December 13, 2002
Accepted after revision March 18, 2003

Synaptically activated Ca2+ waves in layer 2/3 and layer 5 rat neocortical pyramidal neurons

Matthew Larkum1, Shigeo Watanabe2, Takeshi Nakamura3, Nechama Lasser-Ross2, and W. Ross2*

1 Abteilung Zellphysiologie, Max-Planck-Institüt f ür medizinische Forschung, D-69120 Heidelberg, Germany,
2 Department of Physiology, New York Medical College, Valhalla, NY 10595, USA
3 Department of Molecular Neurobiology, University of Tokyo, Tokyo 108-8639, Japan

* To whom correspondence should be addressed. E-mail: ross{at}nymc.edu.

Calcium waves in layer 2/3 and layer 5 neocortical somatosensory pyramidal neurons were examined in slices from 2- to 8-week-old rats. Repetitive synaptic stimulation evoked a delayed, all-or-none [Ca2+]i increase primarily on the main dendritic shaft. This component was blocked by 1 mM MCPG, 10 µM ryanodine, 1 mg ml-1 internal heparin, and was not blocked by 400 µM internal Ruthenium Red, indicating that it was due to Ca2+ release from internal stores by inositol 1,4,5-trisphosphate (IP3) mobilized via activation of metabotropic glutamate receptors. Calcium waves were initiated on the apical shaft at sites between the soma to around the main branch point, mostly at insertion points of oblique dendrites, and spread in both directions along the shaft. In the proximal dendrites the peak amplitude of the resulting [Ca2+]i change was much larger than that evoked by a train of Na+ spikes. In distal dendrites the peak amplitude was comparable to the [Ca2+]i change due to a Ca2+ spike. IP3-mediated Ca2+ release also was observed in the presence of the metabotropic agonists t-ACPD and carbachol when backpropagating spikes were generated. Ca2+ entry through NMDA receptors was observed primarily on the oblique dendrites. The main differences between waves in neocortical neurons and in previously described hippocampal pyramidal neurons were, (a) Ca2+ waves in L5 neurons could be evoked further out along the main shaft, (b) Ca2+ waves extended slightly further out into the oblique dendrites and (c) higher concentrations of bath-applied t-ACPD and carbachol were required to generate Ca2+ release events by backpropagating action potentials.




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