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J Neurophysiol (December 29, 2004). doi:10.1152/jn.00636.2004
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Submitted on June 23, 2004
Accepted on December 22, 2004

Persistent sodium currents in Mesencephalic V Neurons participate in burst generation and control of membrane excitability

Nanping Wu1*, Akifumi Enomoto2, Susumu Tanaka3, Chie-Fang Hsiao1, Duane Q. Nykamp4, Eugene Izhikevich5, and Scott H. Chandler1

1 Physiological Science, University of California at Los Angeles, Los Angeles, CA, USA
2 Physiological Science, University of California at Los Angeles, Los Angeles, CA, USA; 1st Department of Oral and Maxillofacial Surgery, Graduate School of Dentistry, Osaka University, Suita, Osaka, Japan
3 Physiological Science, University of California at Los Angeles, Los Angeles, CA, USA; Department of Oral and Maxillofacial Surgery, Matsumoto Dental University, Matsumoto, Nagano, Japan
4 School of Mathematics, University of Minnesota, Minneapolis, MN, USA
5 Neurosciences Institute, San Diego, CA, USA

* To whom correspondence should be addressed. E-mail: wnanping{at}mednet.ucla.edu.

The functional and biophysical properties of a persistent sodium current (INaP) previously proposed to participate in the generation of subthreshold oscillations and burst discharge in Mesencephalic trigeminal sensory neurons (Mes V) were investigated in brainstem slices (rats p7-12) using whole-cell patch clamp methods. INaP activated around -76 mV and peaked at -48 mV with V1/2 of -58.7 mV. Ramp voltage clamp protocols demonstrated that INaP undergoes time- as well as voltage-dependent inactivation and recovery from inactivation in the range of several seconds ({tau}onset = 2.04 sec, {tau}recov = 2.21 sec). Riluzole (≤5 µM) substantially reduced INaP, membrane resonance, postinhibitory rebound (PIR), subthreshold oscillations and completely blocked bursting but produced modest effects on the fast transient Na+ current (INaT). Prior to complete cessation, burst cycle duration was increased substantially, while modest and inconsistent changes in burst duration were observed. The properties of the INaT were obtained and revealed that the amplitude and voltage dependence of the resulting "window current" were not consistent with those of the observed INaP recorded in the same neurons. This suggests an additional mechanism for the origin of INaP. A neuronal model was constructed using realistic Hodgkin-Huxley parameters for Na+ and K+ currents that simulated the experimentally observed membrane resonance, subthreshold oscillations, bursting and PIR. Alterations in the model gNaP parameters indicate that INaP is critical for control of both subthreshold and suprathreshold Mes V neuron membrane excitability and burst generation.




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