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The Journal of Neurophysiology Vol. 83 No. 5 May 2000, pp. 2562-2579
Copyright ©2000 by the American Physiological Society
1Department of Neurology and Neurosurgery, Montreal Neurological Institute and McGill University, Montreal, Quebec H3A 2B4, Canada; 2Department of Experimental Neurophysiology, Istituto Nazionale Neurologico C. Besta, Milan 510, 20133 Italy; 3Department of Numerical Analysis and Computing Science, Royal Institute of Technology, S-100 44 Stockholm, Sweden; and 4Department of Psychology, Boston University, Boston, Massachusetts 02215
Dickson, Clayton T.,
Jacopo Magistretti,
Mark H. Shalinsky,
Erik Fransén,
Michael E. Hasselmo, and
Angel Alonso.
Properties and Role of Ih in the
Pacing of Subthreshold Oscillations in Entorhinal Cortex Layer II
Neurons. J. Neurophysiol. 83: 2562-2579, 2000. Various subsets of brain neurons express a
hyperpolarization-activated inward current
(Ih) that has been shown to be instrumental in pacing oscillatory activity at both a single-cell and a network level. A characteristic feature of the stellate cells (SCs) of entorhinal cortex (EC) layer II, those neurons giving rise to the main
component of the perforant path input to the hippocampal formation, is
their ability to generate persistent, Na+-dependent
rhythmic subthreshold membrane potential oscillations, which are
thought to be instrumental in implementing theta rhythmicity in the
entorhinal-hippocampal network. The SCs also display a robust
time-dependent inward rectification in the hyperpolarizing direction
that may contribute to the generation of these oscillations. We
performed whole cell recordings of SCs in in vitro slices to investigate the specific biophysical and pharmacological properties of
the current underlying this inward rectification and to clarify its potential role in the genesis of the subthreshold oscillations. In
voltage-clamp conditions, hyperpolarizing voltage steps evoked a slow,
noninactivating inward current, which also deactivated slowly on
depolarization. This current was identified as
Ih because it was resistant to extracellular
Ba2+, sensitive to Cs+, completely and
selectively abolished by ZD7288, and carried by both Na+
and K+ ions. Ih in the SCs had
an activation threshold and reversal potential at approximately 
45
and 20 mV, respectively. Its half-activation voltage was 77 mV.
Importantly, bath perfusion with ZD7288, but not Ba2+,
gradually and completely abolished the subthreshold oscillations, thus
directly implicating Ih in their generation.
Using experimentally derived biophysical parameters for
Ih and the low-threshold persistent Na+ current (INaP) present in
the SCs, a simplified model of these neurons was constructed and their
subthreshold electroresponsiveness simulated. This indicated that the
interplay between INaP and Ih can sustain persistent subthreshold
oscillations in SCs. INaP and
Ih operate in a "push-pull" fashion
where the delay in the activation/deactivation of
Ih gives rise to the oscillatory process.
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