The CPGs for Limbed Locomotion–Facts and Fiction

The neuronal networks that generate locomotion are well understood in swimming animals such as the lamprey, zebrafish and tadpole. The networks controlling locomotion in tetrapods remain, however, still enigmatic with an intricate motor pattern required for the control of the entire limb during the support, lift off, and flexion phase, and most demandingly when the limb makes contact with ground again. It is clear that the inhibition that occurs between bursts in each step cycle is produced by V2b and V1 interneurons, and that a deletion of these interneurons leads to synchronous flexor–extensor bursting. The ability to generate rhythmic bursting is distributed over all segments comprising part of the central pattern generator network (CPG). It is unclear how the rhythmic bursting is generated; however, Shox2, V2a and HB9 interneurons do contribute. To deduce a possible organization of the locomotor CPG, simulations have been elaborated. The motor pattern has been simulated in considerable detail with a network composed of unit burst generators; one for each group of close synergistic muscle groups at each joint. This unit burst generator model can reproduce the complex burst pattern with a constant flexion phase and a shortened extensor phase as the speed increases. Moreover, the unit burst generator model is versatile and can generate both forward and backward locomotion.

Experimental proof that multivariate patterns among muscle attachments (entheses) can reflect repetitive muscle use

Reconstructions of habitual activity in past populations and extinct human groups is a primary goal of paleoanthropological research. Muscle attachment scars (entheses) are widely considered as indicators of habitual activity and many attempts have been made to use them for this purpose. However, their interpretation remains equivocal due to methodological limitations and a paucity of empirical data supporting an interaction between systematic muscle forces and entheseal morphology. We have recently addressed the first issue with precise three-dimensional measuring protocols and rigorous multivariate analysis focusing on the patterns among different entheses rather than comparing each entheseal structure separately. In a previous study, the resulting entheseal correlations reflected synergistic muscle groups that separated individuals according to their lifelong occupational activities. Here we address the second issue by applying this methodology to existing micro-computed tomography data from rats that have undergone muscle stimulation under experimental conditions. In contrast to previous animal studies, we relied on blind analytical procedures across two research institutions and controlled for most factors of interindividual variability. Results demonstrated that the multivariate associations among different entheseal surfaces can directly reflect repetitive muscle recruitment and provide essential information on muscle use.

Neuromuscular adjustments of gait associated with unstable conditions

A compact description of coordinated muscle activity is provided by the factorization of electromyographic (EMG) signals. With the use of this approach, it has consistently been shown that multimuscle activity during human locomotion can be accounted for by four to five modules, each one comprised of a basic pattern timed at a different phase of gait cycle and the weighting coefficients of synergistic muscle activations. These modules are flexible, in so far as the timing of patterns and the amplitude of weightings can change as a function of gait speed and mode. Here we consider the adjustments of the locomotor modules related to unstable walking conditions. We compared three different conditions, i.e., locomotion of healthy subjects on slippery ground (SL) and on narrow beam (NB) and of cerebellar ataxic (CA) patients on normal ground. Motor modules were computed from the EMG signals of 12 muscles of the right lower limb using non-negative matrix factorization. The unstable gait of SL, NB, and CA showed significant changes compared with controls in the stride length, stride width, range of angular motion, and trunk oscillations. In most subjects of all three unstable conditions, >70% of the overall variation of EMG waveforms was accounted for by four modules that were characterized by a widening of muscle activity patterns. This suggests that the nervous system adopts the strategy of prolonging the duration of basic muscle activity patterns to cope with unstable conditions resulting from either slippery ground, reduced support surface, or pathology.