The mitral-granule reciprocal synapse shapes the response of the olfactory bulb to odour stimuli by mediating lateral and reciprocal inhibition. We investigated the short-term plasticity of both the mitral-to-granule excitatory synapse and the granule-to-mitral inhibitory synapse in rat olfactory bulb slices, using whole-cell patch clamp recordings. The granule-to-mitral inhibitory synapse invariably exhibited paired-pulse depression at interstimulus intervals of less than a second, while the mitral-to-granule excitatory synapse showed heterogeneous responses, which on average yielded a moderate facilitation. Trains of stimuli led to a much greater depression at the granule-to-mitral synapse than at the mitral-to-granule synapse. Since mitral cells commonly respond to odours by burst firing with each inhalation cycle, we used bursts of stimuli to study recovery from depression. We found that recovery from depression induced by fast trains of stimuli was more rapid at the mitral-to-granule synapse than at the granule-to-mitral synapse. In addition, depression was enhanced by higher calcium concentrations, suggesting at least partial contribution of presynaptic mechanisms to short-term depression. The observed short-term plasticity could enable mitral cells to overcome autoinhibition and increase action potential propagation along lateral dendrites by burst firing.
In mice, most studies of the organization of the spinal central pattern generator (CPG) for locomotion, and its component neuron classes, have been performed on neonatal [postnatal day (P)2-P4] animals. While the neonatal spinal cord can generate a basic locomotor pattern, it is often argued that the CPG network is in an immature form whose detailed properties mature with postnatal development. Here, we compare intrinsic properties and serotonergic modulation of the V2a class of excitatory spinal interneurons in behaviorally mature (older than P43) mice to those in neonatal mice. Using perforated patch recordings from genetically tagged V2a interneurons, we revealed an age-dependent increase in excitability. The input resistance increased, the rheobase values decreased, and the relation between injected current and firing frequency (F/I plot) showed higher excitability in the adult neurons, with almost all neurons firing tonically during a current step. The adult action potential (AP) properties became narrower and taller, and the AP threshold hyperpolarized. While in neonates the AP afterhyperpolarization was monophasic, most adult V2a interneurons showed a biphasic afterhyperpolarization. Serotonin increased excitability and depolarized most neonatal and adult V2a interneurons. However, in ∼30% of adult V2a interneurons, serotonin additionally elicited spontaneous intrinsic membrane potential bistability, resulting in alternations between hyperpolarized and depolarized states with a dramatically decreased membrane input resistance and facilitation of evoked plateau potentials. This was never seen in younger animals. Our findings indicate a significant postnatal development of the properties of locomotor-related V2a interneurons, which could alter their interpretation of synaptic inputs in the locomotor CPG.
.2010. In rhythmic neural circuits, a neuron often fires action potentials with a constant phase to the rhythm, a timing relationship that can be functionally significant. To characterize these phase preferences in a large-scale, cell type-specific manner, we adapted multitaper coherence analysis for two-photon calcium imaging. Analysis of simulated data showed that coherence is a simple and robust measure of rhythmicity for calcium imaging data. When applied to the neonatal mouse hindlimb spinal locomotor network, the phase relationships between peak activity of Ͼ1,000 ventral spinal interneurons and motor output were characterized. Most interneurons showed rhythmic activity that was coherent and in phase with the ipsilateral motor output during fictive locomotion. The phase distributions of two genetically identified classes of interneurons were distinct from the ensemble population and from each other. There was no obvious spatial clustering of interneurons with similar phase preferences. Together, these results suggest that cell type, not neighboring neuron activity, is a better indicator of an interneuron's response during fictive locomotion. The ability to measure the phase preferences of many neurons with cell type and spatial information should be widely applicable for studying other rhythmic neural circuits.
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