Higher Responsiveness of Pattern Generation Circuitry to Sensory Stimulation in Healthy Humans Is Associated with a Larger Hoffmann Reflex.

The state and excitability of pattern generators are attracting the increasing interest of neurophysiologists and clinicians for understanding the mechanisms of the rhythmogenesis and neuromodulation of the human spinal cord. It has been previously shown that tonic sensory stimulation can elicit non-voluntary stepping-like movements in non-injured subjects when their limbs were placed in a gravity-neutral unloading apparatus. However, large individual differences in responsiveness to such stimuli were observed, so that the effects of sensory neuromodulation manifest only in some of the subjects. Given that spinal reflexes are an integral part of the neuronal circuitry, here we investigated the extent to which spinal pattern generation excitability in response to the vibrostimulation of muscle proprioceptors can be related to the H-reflex magnitude, in both the lower and upper limbs. For the H-reflex measurements, three conditions were used: stationary limbs, voluntary limb movement and passive limb movement. The results showed that the H-reflex was considerably higher in the group of participants who demonstrated non-voluntary rhythmic responses than it was in the participants who did not demonstrate them. Our findings are consistent with the idea that spinal reflex measurements play important roles in assessing the rhythmogenesis of the spinal cord.

Muscle Responses to Passive Joint Movements in Infants During the First Year of Life.

Muscle tone represents one of the important concepts for characterizing changes in the state of the developing nervous system. It can be manifested in the level of activity of flexors and extensors and in muscle reactions to its passive stretching (StR) or shortening (ShR). Here we investigated such reactions in a cohort of healthy infants aged from 2 weeks to 12 months. We examined the presence and the characteristics of StR and ShR during slow passive cyclic flexion/extension movements (T~3 s) in the hip, knee, ankle, and elbow joints while awake infants were in the supine position. The results showed that most infants demonstrated prominent ShRs in response to passive joint rotations, although the StR was observed more frequently, suggesting that the ShR is an important component of adaptive motor behavior already at an early developmental stage. Interestingly, the occurrence of both StR and ShR in most muscles significantly decreased throughout the first year of life. Passive cyclic flexion/extension movements could also evoke rhythmic muscle responses in other joints or in the contralateral limb, however, such responses were predominantly observed in younger infants (<6 months). A noticeable manifestation of muscle reactions at an early developmental stage, along with spontaneous motor activity in this period of life, may reflect the processes underlying a formation of appropriate muscle tone and the self-organization of neural circuits. A substantial reduction of ipsilateral and contralateral muscle responses to passive movements with age is consistent with the idea of a functional reorganization of the motor circuitry during early development.

Tonic and Rhythmic Spinal Activity Underlying Locomotion.

In recent years, many researches put significant efforts into understanding and assessing the functional state of the spinal locomotor circuits in humans. Various techniques have been developed to stimulate the spinal cord circuitries, which may include both diffuse and quite specific tuning effects. Overall, the findings indicate that tonic and rhythmic spinal activity control are not separate phenomena but are closely integrated to properly initiate and sustain stepping. The spinal cord does not simply transmit information to and from the brain. Its physiologic state determines reflex, postural and locomotor control and, therefore, may affect the recovery of the locomotor function in individuals with spinal cord and brain injuries. This review summarizes studies that examine the rhythmogenesis capacity of cervical and lumbosacral neuronal circuitries in humans and its importance in developing central pattern generator-modulating therapies.

Rhythmic wrist movements facilitate the soleus H-reflex and non-voluntary air-stepping in humans.

Neural coupling between the upper and lower limbs during human walking is supported by modulation of cross-limb reflexes and the presence of rhythmic activity in the proximal arm muscles. Nevertheless, the involvement of distal arm muscles in cyclic movements and sensorimotor neuromodulation is also suggested given their step-synchronized activation in many locomotor-related tasks (e.g., swimming, skiing, climbing, cycling, crawling, etc.). Here we investigated the effect of rhythmic wrist movements, separately and in conjunction with arm swinging, on the characteristics of non-voluntary cyclic leg movements evoked by muscle vibration in a gravity neutral position and on the soleus H-reflex of the stationary legs. For the H-reflex modulation, five conditions were compared: stationary arms, voluntary alternating upper limb swinging, combined upper limb and wrist motion, wrist movements only and motion of the upper limbs with addition of load. Rhythmic wrist movements significantly facilitated the amplitude of non-voluntary leg oscillations, including ankle joint oscillations, and the H-reflex. The latter effect was related to rhythmicity of wrist motion rather than to a simple extra tension in the upper limb muscles (a kind of the Jendrassik manoeuvre) since adding resistance to arm oscillations (without flexion-extension in the wrist joint) had an opposite inhibitory effect on the H-reflex. Our results further support the existence of connections between the distal parts of the upper and lower extremities at the neural level, suggesting that wrist joint movements can be an important component of motor neurorehabilitation.

Tapping into rhythm generation circuitry in humans during simulated weightlessness conditions.

An ability to produce rhythmic activity is ubiquitous for locomotor pattern generation and modulation. The role that the rhythmogenesis capacity of the spinal cord plays in injured populations has become an area of interest and systematic investigation among researchers in recent years, despite its importance being long recognized by neurophysiologists and clinicians. Given that each individual interneuron, as a rule, receives a broad convergence of various supraspinal and sensory inputs and may contribute to a vast repertoire of motor actions, the importance of assessing the functional state of the spinal locomotor circuits becomes increasingly evident. Air-stepping can be used as a unique and important model for investigating human rhythmogenesis since its manifestation is largely facilitated by a reduction of external resistance. This article aims to provide a review on current issues related to the "locomotor" state and interactions between spinal and supraspinal influences on the central pattern generator (CPG) circuitry in humans, which may be important for developing gait rehabilitation strategies in individuals with spinal cord and brain injuries.

Activation of spinal locomotor circuits in the decerebrated cat by spinal epidural and/or intraspinal electrical stimulation.

The present study was designed to further compare the stepping-like movements generated via epidural (ES) and/or intraspinal (IS) stimulation. We examined the ability to generate stepping-like movements in response to ES and/or IS of spinal lumbar segments L1-L7 in decerebrate cats. ES (5-10 Hz) of the dorsal surface of the spinal cord at L3-L7 induced hindlimb stepping-like movements on a moving treadmill belt, but with no rhythmic activity in the forelimbs. IS (60 Hz) of the dorsolateral funiculus at L1-L3 (depth of 0.5-1.0mm from the dorsal surface of the spinal cord) induced quadrupedal stepping-like movements. Forelimb movements appeared first, followed by stepping-like movements in the hindlimbs. ES and IS simultaneously enhanced the rhythmic performance of the hindlimbs more robustly than ES or IS alone. The differences in the stimulation parameters, site of stimulation, and motor outputs observed during ES vs. IS suggest that different neural mechanisms were activated to induce stepping-like movements. The effects of ES may be mediated more via dorsal structures in the lumbosacral region of the spinal cord, whereas the effects of IS may be mediated via more ventral propriospinal networks and/or brainstem locomotor areas. Furthermore, the more effective facilitation of the motor output during simultaneous ES and IS may reflect some convergence of pathways on the same interneuronal populations involved in the regulation of locomotion.