Neural pathways of movement fractionation

Abstract

Stroke is a common neurological event which often results in motor deficits of the hand and arm. The reticulospinal tract (RST) may partly underlie residual hand and arm movement ability after a stroke but remains poorly characterised. A greater understanding of the RST could inform work to improve motor recovery. Additionally, the development of non-invasive methods of probing the RST in humans should allow comparison of the characteristics of the RST across species. It has been suggested that the RST is involved in mediating muscle responses to auditory startle, experimentally known as the StartReact paradigm. However, it was not clear how this pathway was involved. A human experiment presented here suggests that the RST comprises the final pathway in the StartReact effect, confirming it as a technique to probe the RST in humans. Other factors such as habituation and the validity of a marker of the StartReact effect were also further explored; these findings may inform future use of the technique. The output divergence, co-activation patterns, level of fractionation and synergies produced by the RST were further characterised in macaques and baboons; these factors had previously been mostly unexplored. In macaques, two subdivisions of primary motor cortex (M1) were also characterised in order to compare to the RST. These subdivisions are based upon the presence of corticomotoneuronal (CM) cells, and consist of ‘old’ (CM cells absent) and ‘new’ (CM cells present) M1. Stimulation of new M1 produced a higher level of fractionation of movement than stimulation of old M1 and the reticular formation (RF). The RF is suggested to produce slightly more fractionated behaviour than old M1, though the baboon RF responses may be less fractionated than those from macaque old M1. Output divergence of the RF as well as old and new M1 was also explored. However, methodological limitations may have biased the results towards muscles with more excitable motoneurons, or monosynaptic connections. Abstract ii In baboons, threshold stimulation elicited responses in upper limb and axial muscles only, with higher stimulation intensities or trains of pulses required to activate leg muscles. In contrast to long-held beliefs about RST output, distal upper limb muscles were more commonly activated than proximal ones. Previously reported attempts to record natural electromyography (EMG) data from macaques were limited to controlled experimental settings, and hence may have differed from EMG observed during truly natural behaviours. Here, EMG was recorded from 18 muscles in one macaque over several hours of natural, untrained activity in her home cage. Two matrix decomposition algorithms extracted three to four dominant synergies from the data. This number is comparable to that previously described for ‘natural’ behaviour in more controlled conditions, suggesting that it accurately reflects the dominant synergies used across both conditions. Future work should aim to delineate the respective contributions of the RST and corticospinal tract to natural movement and to develop approaches to manipulate RST projections in humans to improve post-stroke motor outcomes.