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Modeling Spike-Train Processing in the Cerebellum Granular Layer and Changes in Plasticity Reveal Single Neuron Effects in Neural Ensembles

DOI: 10.1155/2012/359529

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The cerebellum input stage has been known to perform combinatorial operations on input signals. In this paper, two types of mathematical models were used to reproduce the role of feed-forward inhibition and computation in the granular layer microcircuitry to investigate spike train processing. A simple spiking model and a biophysically-detailed model of the network were used to study signal recoding in the granular layer and to test observations like center-surround organization and time-window hypothesis in addition to effects of induced plasticity. Simulations suggest that simple neuron models may be used to abstract timing phenomenon in large networks, however detailed models were needed to reconstruct population coding via evoked local field potentials (LFP) and for simulating changes in synaptic plasticity. Our results also indicated that spatio-temporal code of the granular network is mainly controlled by the feed-forward inhibition from the Golgi cell synapses. Spike amplitude and total number of spikes were modulated by LTP and LTD. Reconstructing granular layer evoked-LFP suggests that granular layer propagates the nonlinearities of individual neurons. Simulations indicate that granular layer network operates a robust population code for a wide range of intervals, controlled by the Golgi cell inhibition and is regulated by the post-synaptic excitability. 1. Introduction Decoding neural activity is the key to understand spatiotemporal patterns that the brain receives as sensory information regarding the world. Time-scale of operation is closely correlated to the activity of the neural circuit and decoding such activity reveals principles regarding the function. One of the main circuits in the cerebellum is the large input layer circuit formed of granule and Golgi cells. Spatiotemporal information is one of the unique functional characteristics observed in the cerebellar input layer network [1, 2]. Cerebellar granular layer forms the input stage of the cerebellum in which information coming from the peripheral and central systems converge through the mossy fibers. The granular layer has by far the smallest (~5?μm) and the most numerous neurons (~1011) in humans. Understanding how the granular layer process information appears critical to understand the cerebellar function, since signals coming into upper cortical layers are provided by the granular layer. The granule cells form the largest neuronal population in the mammalian brain and regulate information transfer along the major afferent systems to the cerebellum. The granule layer receives

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