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The Journal of Neuroscience, January 12, 2005, 25(2):464-472; doi:10.1523/JNEUROSCI.3871-04.2005

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Cellular/Molecular
Determinants of Action Potential Propagation in Cerebellar Purkinje Cell Axons

Pablo Monsivais, ^* Beverley A. Clark, ^* Arnd Roth, and Michael Häusser

Wolfson Institute for Biomedical Research and Department of Physiology, University College London, London WC1E 6BT, United Kingdom

Axons have traditionally been viewed as highly faithful transmitters of action potentials. Recently, however, experimental evidence has accumulated to support the idea that under some circumstances axonal propagation may fail. Cerebellar Purkinje neurons fire highfrequency simple spikes, as well as bursts of spikes in response to climbing fiber activation (the "complex spike"). Here we have visualized the axon of individual Purkinje cells to directly investigate the relationship between somatic spikes and axonal spikes using simultaneous somatic whole-cell and cell-attached axonal patch-clamp recordings at 200-800 µm from the soma. We demonstrate that sodium action potentials propagate at frequencies up to 260 Hz, higher than simple spike rates normally observed in vivo. Complex spikes, however, did not propagate reliably, with usually only the first and last spikes in the complex spike waveform being propagated. On average, only 1.7 ± 0.2 spikes in the complex spike were propagated during resting firing, with propagation limited to interspike intervals above 4 msec. Hyperpolarization improved propagation efficacy without affecting total axonal spike number, whereas strong depolarization could abolish propagation of the complex spike. These findings indicate that the complex spike waveform is not faithfully transmitted to downstream synapses and that propagation of the climbing fiber response may be modulated by background activity.

Key words: axon; climbing fiber; excitability; refractory; sodium channel; burst

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Received Sep 17, 2004; revised November 25, 2004; accepted November 29, 2004.

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