Mechanisms underlying firing in healthy and sick human motoneurons

Abstract

International audienceIn an address to the British Association for the Advancement of Science in Cambridge, Professor Sherrington introduced the terms " motor neurone " and " the final common path, " the latter term implying that all motor commands converge onto the motoneuron which integrates the incoming information and passes the net information to the muscle for contraction (Sherrington, 1904). The relative ease of access of the spinal motoneuron made it feasible to set up techniques for investigating the physiological, biophysical and molecular properties of these neurons. It became the most investigated neuron of the CNS in the twentieth century and the information gained from studies on motoneurons formed the basis for examining the other neurons of the CNS. Since the compound action potential of a muscle unit is strictly related one-to-one to the action potential arriving from the innervating motoneuron, the statistical analysis of muscle unit action potentials provides an investigator with an elegant way to probe the properties of motoneurons in behaving humans. In the following review the terms motoneuron and motor unit might be used interchangeably. Different aspects of human motoneuron investigations in health and disease are presented in 16 articles of this topic which are summarized below. An increase in the net excitatory synaptic input to the motoneuron pool results in an increase in the level of muscle contraction by recruitment of additional motor units (MUs) and an increase in firing rates of the already recruited units (Milner-Brown et al., 1973; Henneman et al., 1974). The principle of orderly recruitment of motoneurons by size was originally proposed by Henneman (1957) but was later questioned by other researchers presenting examples of selective, rather than orderly recruitment (e.g., Smith et al., 1980). These controversies are assessed by Bawa et al. (2014), and the opinion unifying the concept of orderly recruitment is presented. In humans, increases in firing rates of motor units have been shown to follow the " onion skin " pattern at lower levels of contraction, meaning that the lower-threshold motor units discharge with higher rates than higher-threshold ones. However, studies performed on the whole range of muscle forces indicated that for higher force levels the motor unit firing rate follows a " reverse onion skin " pattern. Hu et al. (2014) decided to approach this problem using small surface electrodes and step increases in force instead of the " ramp and hold " protocols used by previous authors. They showed that the " onion skin " pattern was preserved until 15% of maximal voluntary contraction, and from their results predict this pattern to be valid for the whole range of muscle forces, which is not supported by the previous published works. However, the reported rate saturation of the MUs discharging with higher rates implies that at the higher forces the " reverse onion skin " pattern may be expected. In another paper, Duchateau and Baudry (2014) show that during ballistic c...