PMC:1255741 / 39848-43962 JSONTXT

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This notion will be further tested in mice with piriform cortex-specific GluR-B ablation.\nA direct contribution to the phenotype by Ca2+ influx through genetically modified AMPA channels seems to be likely, since mice expressing Q/R site-unedited GluR-B showed even better odor learning and discrimination performance compared with mice with depleted GluR-B. In both mouse models Ca2+ influx through AMPA channels is increased, whereas the effect on the macroscopic AMPA conductance differs. AMPAR currents are reduced in mice not expressing GluR-B [51], possibly due to fewer synaptic AMPA channels and impaired AMPAR trafficking and recycling. In mice expressing the unedited form of GluR-B(Q), macroscopic conductance in whole-cell patches of CA1 pyramidals is increased [60], but excitatory transmission in CA3-to-CA1 cell synapses is somewhat reduced, in spite of a lower threshold for generating a population spike [58], indicating increased synaptic excitability by sustained GluR-B(Q) expression, in line with the seizure-prone phenotype. Yet, both mouse models yield enhanced discrimination and learning capability. The milder phenotype upon GluR-B depletion could be attributed to reduced AMPAR densities at synapses [40,41] and therefore probably to a lesser extent of Ca2+ influx. This is further supported by kainate-induced Co2+ uptake in acute brain slices, revealing considerably higher Co2+ uptake via Ca2+-permeable AMPARs in hippocampal pyramidal neurons of mice expressing GluR-B(Q), than those lacking GluR-B (DRS, RS, and PHS, unpublished data).\nNormally, GluR-B-containing AMPARs are prominently localized at the dendrodendritic synapse between mitral and granule cells in the olfactory bulb [64,65]. As Ca2+ influx through glutamate receptors is thought to contribute to lateral and recurrent inhibition ([38] see also [39]), the absence of GluR-B, or the presence of GluR-B(Q) in the olfactory bulb, is likely to result in increased inhibition between the principal neurons in this structure. Notably, the improved olfactory discrimination capabilities were apparent even upon relatively small reductions in GluR-B levels. It will be interesting to see if GluR-B+/neo mice also exhibit enhanced odor discrimination, which is indeed likely given that low levels of GluR-B(Q), and hence a small increase in Ca2+-permeable AMPARs, arise from the attenuated GluR-Bneo allele [60], and, moreover, that GluR-B levels are decreased in these mice due to the single copy of the GluR-B+ allele.\nIn virtually all models of the olfactory bulb, lateral (and, in fewer models, also recurrent) inhibition plays a dominant role, either in establishing spatiotemporal dynamics [32,66–69], or in directly enhancing contrast and therefore simplifying discrimination and learning of similar odorants [24,28,29,36]. Increased inhibition will thus in general improve discriminability, consistent with the behavioral improvements observed with GluR-B(Q) expression, the GluR-B knockout, and the “rescue” mice. A direct test of this link would require the quantitative measurement of inhibition in the intact preparation, a task that might be feasible with further improved in vivo electrophysiological techniques, such as targeted recordings [70,71] or simultaneous pre- and postsynaptic intracellular 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