
A second line of indirect evidence comes from induced rhythmic activities in paralyzed leg muscles by tonic epidural electrical stimulation (EES) of the lumbar spinal cord 16, 17. A common denominator in the individuals expressing this type of spinal myoclonus was the presence of additional musculoskeletal pathologies below the level of the lesion, particularly of the hip 12, 13, 14, 15. Despite the lack of a locomotor pattern, it was suggested that this type of self-sustained rhythmic activity was due to a partial release of a CPG 12, 13, 14. The electromyographic patterns largely involved synchronous bursting across muscles. First, spontaneously emerging rhythmic spinal myoclonus at a highly reproducible rate of 0.3–0.6 Hz and involving multiple lower-limb muscles was described in six individuals with chronic, clinically complete SCI 12, 13, 14. Indirect evidence of spinally generated rhythmic activity comes from specific motor phenomena in individuals with spinal cord injury (SCI) 6. In humans, the experimental procedures necessary for the demonstration of fictive locomotion cannot be applied.
#Natural phenomena full
Yet, full fictive locomotion was not evoked by any single condition. If combined, the component fictive patterns obtained under different experimental conditions would resemble a true fictive locomotor pattern. In adult marmoset monkeys, electrical stimulation of the brainstem in the decerebrate preparation or administration of different pharmacological agents (clonidine, NMDA, serotonin) following spinalization generated various rhythmic activities that presented “component fictive patterns” with alternating as well as synchronous bursts in flexor and extensor nerves 11.

Attempts to elicit fictive locomotion by L-Dopa and dorsal column stimulation failed in acutely spinalized and immobilized macaque monkeys 10. Such explicit evidence is sparse in non-human primates. Fictive locomotion was expressed, inter alia, in the immobilized (neuromuscular block by curarization), acute spinal cat pretreated with L-Dopa by electrical dorsal root or dorsal column stimulation 7, in the immobilized decerebrate cat by electrical stimulation of brainstem structures 8, or in the isolated neonatal mouse spinal cord through the administration of excitatory amino acids and serotonin 9.
#Natural phenomena generator
This central pattern generator (CPG) capability of the lumbar spinal circuits was directly demonstrated in various animal models by the generation of so-called fictive locomotion, i.e., rhythmic motor output recorded from ventral roots or peripheral nerves with flexor-extensor and left-right alternations in the absence of descending neural inputs or peripheral feedback cues 6. In reduced animal preparations, the neural control exerted by these circuits was shown to be necessary and sufficient to generate the basic motor patterns underlying locomotion 3, 4, 5. In the motor control system for hindlimb locomotion in quadrupedal mammals, specialized neural circuits in the lumbar spinal cord represent the final network 1, 2. Spinal myoclonus and the EES-induced activity demonstrate that the human lumbar spinal cord contains distinct mechanisms for generating rhythmic multi-muscle patterns. These motor deletions, with preserved cycle frequency and period when rhythmic activity resumed, were previously reported only in animal studies and suggest a separation between rhythm generation and pattern formation. The EES-induced patterns were fundamentally different in that they included flexor-extensor and left-right alternations, hallmarks of locomotor CPGs, and showed spontaneous errors in rhythmicity.

Analysis of muscle activation patterns suggested that the myoclonus tapped into spinal circuits that generate muscle spasms, rather than reflecting locomotor CPG activity as previously thought. Here, we investigated a case of a male individual with complete thoracic spinal cord injury who presented with a rare form of self-sustained rhythmic spinal myoclonus in the legs and rhythmic activities induced by epidural electrical stimulation (EES). The existence and function of CPGs in humans have remained controversial. Lumbar central pattern generators (CPGs) control the basic rhythm and coordinate muscle activation underlying hindlimb locomotion in quadrupedal mammals.
