Convergence of muscle spindle afferents on single neurons of the cat dorsal spino-cerebellar tract and their synaptic efficacy

By means of tungsten microelectrodes, action potentials from axons within the dorsal spino-cerebellar tract (DSCT) and from muscle spindle afferents were recorded. A quantitative study was performed in monomuscular DSCT neurons which were excited predominantly by Ia fibers originating in the gastrocnemius muscles. In some experiments single Ia fibers were stimulated electrically while the impulse sequence of a DSCT neuron postsynaptic to the respective afferent fiber was recorded. The gastrocnemius DSCT neurons receive excitatory inputs from 10-18 Ia muscle spindle afferents. The efficacy of each of these inputs is very similar. Thus the neuronal activation decreased approximately linearly with the number of the excitatory afferents cut. Cross-correlograms between the impulse sequence of a Ia gastrocnemius muscle spindle afferent and a DSCT neuron postsynaptic to it exhibited an increased discharge probability of the DSCT neuron from 3-4 ms to 10 ms after the Ia action potential. With increasing impulse rates of the Ia afferent fibers, the excitatory efficacy of the single action potential decreased, but the overall excitation increased with the presynaptic discharge frequency, according to a hyperbolic function. This effect was tested by electrical stimulation of a single Ia axon exciting the DSCT neuron recorded. Interval histograms computed from DSCT neuron impulse trains at steady stretch conditions were predominantly monomodal. They can be well approximated by a Gaussian distribution. The coefficient of variation was independent of the mean activity. At impulse rates above 25 imp X s-1 a negative correlation between successive intervals was observed in first order joint interval diagrams. With an increasing mean discharge rate this correlation (expressed as the serial linear correlation coefficient of the first order r1,2) became stronger up to--0.62 at 90 imp X s-1. Only in a few neurons did the higher order linear correlation coefficients deviate significantly from zero. In 15% of the observed histograms double discharging (mean interval 3-5 ms) produced bimodal distributions. Under steady-state conditions the response of Ia-activated DSCT cells are linearly related to muscle stretch within a middle range of extensions. The differences between Ia impulse pattern and DSCT neuron impulse pattern at steady stretch are discussed. The number of large dendrites of the principal cells in the nucleus dorsalis (Clarke's column) corresponds to the number of excitatory afferent muscle fibers. It is assumed that each excitatory Ia axon sends one axon collateral to the DSCT neuron, forming a climbing type terminal mainly on one of the large dendrites of a DSCT cell.