In decerebrated, nonanesthetized cats, we made intracellular whole-cell recordings and extracellular cell-attached recordings from granule cells in the cerebellar C3 zone. Spontaneous EPSPs had large, relatively constant peak amplitudes, whereas IPSPs were small and did not appear to contribute substantially to synaptic integration at a short time scale. In many cases, the EPSPs of individual mossy fiber synapses appeared to be separable by their peak amplitudes. A substantial proportion of our granule cells had small receptive fields on the forelimb skin. Skin stimulation evoked explosive responses in which the constituent EPSPs were analyzed. In the rising phase of the response, our analyses indicated a participation of three to four different mossy fiber synapses, corresponding to the total number of mossy fiber afferents. The cutaneous receptive fields of the driven EPSPs overlapped, indicating an absence of convergence of mossy fibers activated from different receptive fields. Also in granule cells activated by joint movements did we find indications that different afferents were driven by the same type of input. Regardless of input type, the temporal patterns of granule cell spike activity, both spontaneous and evoked, appeared to primarily follow the activity in the presynaptic mossy fibers, although much of the nonsynchronized mossy fiber input was filtered out. In contrast to the prevailing theories of granule cell function, our results suggest a function of granule cells as signal-to-noise enhancing threshold elements, rather than as sparse coding pattern discriminators or temporal pattern generators.
Initial investigations of the cerebellar microcircuit inspired the Marr-Albus theoretical framework of cerebellar function. We review recent developments in the experimental understanding of cerebellar microcircuit characteristics and in the computational analysis of Marr-Albus models. We conclude that many Marr-Albus models are in effect adaptive filters, and that evidence for symmetrical long-term potentiation and long-term depression, interneuron plasticity, silent parallel fibre synapses and recurrent mossy fibre connectivity is strikingly congruent with predictions from adaptive-filter models of cerebellar function. This congruence suggests that insights from adaptive-filter theory might help to address outstanding issues of cerebellar function, including both microcircuit processing and extra-cerebellar connectivity.
The highly specific relationships between parallel fiber (PF) and climbing fiber (CF) receptive fields in Purkinje cells and interneurons suggest that normal PF receptive fields are established by CF-specific plasticity. To test this idea, we used PF stimulation that was either paired or unpaired with CF activity. Conspicuously, unpaired PF stimulation that induced long-lasting, very large increases in the receptive field sizes of Purkinje cells induced long-lasting decreases in receptive field sizes of their afferent interneurons. In contrast, PF stimulation paired with CF activity that induced long-lasting decreases in the receptive fields of Purkinje cells induced long-lasting, large increases in the receptive fields of interneurons. These properties, and the fact the mossy fiber receptive fields were unchanged, suggest that the receptive field changes were due to bidirectional PF synaptic plasticity in Purkinje cells and interneurons.
The cutaneous parallel fiber (PF) receptive fields of cerebellar stellate and basket cells in the cerebellar C3 zone in vivo are normally very small but can be dramatically enlarged by climbing fiber (CF)-dependent plasticity. To analyze the effects of this receptive field plasticity, we present for the first time whole-cell patch-clamp recordings from these interneurons during natural and electrical activation of cutaneously driven synaptic input. In "naive" interneurons, peripheral input nearly exclusively activated a few (two to eight) large PF EPSPs from a specific small skin area that overlapped the receptive field of the local CF input. After conjunctive PF and CF stimulation, numerous small and large EPSPs and ramp-like depolarizations could be activated from the entire forelimb skin. These findings therefore confirm previous suggestions that conjunctive PF and CF activation leads to a long-lasting potentiation of PF synaptic input to interneurons. The CF response, which is crucial for the induction of the PF synaptic potentiation, was strong but variable and very different from the conventional EPSPs evoked by PFs.
In cats decerebrated at the intercollicular level, the cutaneous parallel fibre receptive fields of Purkinje cells, molecular layer interneurons and Golgi cells in the cerebellar C3 zone were delineated by natural stimulation of the skin during extracellular unitary recordings. The locations of these receptive fields were compared with the climbing fibre receptive field of the local Purkinje cell and with the receptive fields of other neurons located along a beam of parallel fibres. The parallel fibre receptive fields of these neurons were highly specific to the local climbing fibre receptive field. In Purkinje cells, the parallel fibre receptive fields were located outside the climbing fibre receptive field of the same cell. In contrast, the parallel fibre receptive fields of interneurons were similar to the receptive field of the locally terminating climbing fibres. In both types of neurons, the parallel fibre receptive fields were small and had distinct borders. The location on the skin of the parallel fibre receptive fields differed conspicuously between neighbouring Purkinje cells and between neighbouring interneurons along a beam as well as between Purkinje cells and interneurons in the same electrode tracks. The remarkable specificity between the parallel fibre receptive fields in Purkinje cells and interneurons and the receptive field of the local climbing fibre is most easily explained by different forms of parallel fibre synaptic plasticity.
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