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J Neurophysiol (August 24, 2005). doi:10.1152/jn.00661.2005
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Submitted on June 24, 2005
Accepted on August 17, 2005

An NMDA receptor / nitric oxide cascade is involved in cerebellar LTD but is not localized to the parallel fiber terminal

Jung Hoon Shin and David J. Linden1*

1 Neuroscience, Johns Hopkins University, Baltimore, MD, USA

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

Long-term depression (LTD) of the parallel fiber-Purkinje cell synapse in the cerebellum is a cellular model system which has been suggested to underlie certain forms of motor learning. Induction of cerebellar LTD requires a postsynaptic kinase limb involving activation of mGluR1, protein kinase C{alpha} (PKC{alpha}) and phosphorylation of ser-880 on the AMPA receptor subunit GluR2. Several lines of evidence have also implicated a complementary phosphatase limb in which NMDA receptor-mediated Ca2+ influx activates neuronal nitric oxide synthase (nNOS), the ultimate consequences of which are mediated by nitric oxide (NO), cGMP and inhibition of postsynaptic protein phosphatases. However, the cellular localization of an NMDA/NO cascade has been complicated by the fact that neither functional NMDA receptors nor nNOS are expressed in Purkinje cells. This has lead to a proposal in which NMDA receptors activate nNOS in parallel fibers (Casado et al. 2000; 2002). Here, we confirm that pharmacological blockade of NMDA receptor or NO signaling blocks induction of LTD. However, no evidence was found for functional NMDA receptors in parallel fiber terminals: blockade of NMDA receptors did not alter either presynaptic Ca2+ transients or the frequency of mEPSCs. NMDA receptor blockade did abolish a slow depolarization evoked by burst stimulation of parallel fiber-stellate cell synapses. The application of NMDA evoked a Ca2+ transient in stellate cell terminals but not in parallel fiber terminals. These results are consistent with the hypothesis that an NMDA receptor / NO cascade involved in cerebellar LTD is localized to interneurons rather than parallel fibers.




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