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J Neurophysiol (December 10, 2008). doi:10.1152/jn.90382.2008
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Submitted on March 20, 2008
Revised on November 19, 2008
Accepted on November 19, 2008

Axonal Na+ channels ensure fast spike activation and back-propagation in cerebellar granule cells

Shyam Diwakar1, Jacopo Magistretti1, Mitchell Goldfarb2, Giovanni Naldi3, and Egidio D'Angelo1*

1 University of Pavia
2 Hunter College of City University
3 University of Milan

* To whom correspondence should be addressed. E-mail: dangelo{at}unipv.it.

In most neurons, Na+ channels in the axon are complemented by others localized in the soma and dendrites to ensure spike back-propagation. However, cerebellar granule cells are neurons with simplified architecture, in which the dendrites are short and unbranched and a single thin ascending axon travels toward the molecular layer before bifurcating into parallel fibers. Following previous indications (Magistretti et al., 2006; Goldfarb et al., 2007), here we show that, in cerebellar granule cells, Na+ channels are enriched in the axon, especially in the hillock, but almost absent from soma and dendrites. The impact of this channel distribution on neuronal electroresponsiveness was investigated by multi-compartmental modeling. Numerical simulations indicated that granule cells have a compact electrotonic structure allowing EPSPs to diffuse with little attenuation from dendrites to axon. The spike arose almost simultaneously along the whole axonal ascending branch and invaded the hillock, whose activation promoted spike back-propagation with marginal delay (<200 µs) and attenuation (<20 mV) into the somato-dendritic compartment. These properties allow granule cells to perform sub-millisecond coincidence detection of pre- and postsynaptic activity and to rapidly activate Purkinje cells contacted by the axonal ascending branch.







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