Postural control in the elephant.

As the largest extant legged animals, elephants arguably face the most extreme challenge for stable standing. In this study, we investigated the displacement of the centre of pressure of 12 elephants during quiet standing. We found that the average amplitude of the oscillations in the lateral and fore-aft directions was less than 1.5 cm. Such amplitudes for postural oscillation are comparable with those of dogs and other species, suggesting that some aspects of sensorimotor postural control do not scale with size.

Increasing muscle activity correlations during spontaneous movements in the first six months of life.

Spontaneous muscle activity in the first months of life is an important prerequisite to developing voluntary motor skills and to adapting sensorimotor circuits and muscle tone to body and environmental changes. Even though high variability is a characteristic of early development, several studies have reported significant correlations of limb movements. These assessments were typically made based on kinematics, while the analysis of lower and upper limb muscle activity may provide additional information about maturation of the neuromuscular control. To this end, we examined the electromyographic activity of 12 muscles of the upper and lower limbs in full-term healthy infants (n = 40) aged from 1 week to six months. An increase of ipsilateral and contralateral limb muscle activity correlations with age was found in both flexors and extensors and may reflect a progressive emergence of elements of coordinative neuromuscular behaviour. Correlations between arm and leg muscle responses also increased during passive leg movements. Overall, the findings are consistent with maturation of physiologically relevant neuromuscular network connections during the course of transition from spontaneous-like to voluntary goal-directed movements during early development.

Spinal motoneurons of the human newborn are highly synchronized during leg movements.

Motoneurons of neonatal rodents show synchronous activity that modulates the development of the neuromuscular system. However, the characteristics of the activity of human neonatal motoneurons are largely unknown. Using a noninvasive neural interface, we identified the discharge timings of individual spinal motoneurons in human newborns. We found highly synchronized activities of motoneurons of the tibialis anterior muscle, which were associated with fast leg movements. Although neonates' motor units exhibited discharge rates similar to those of adults, their synchronization was significantly greater than in adults. Moreover, neonatal motor units showed coherent oscillations in the delta band, which is directly translated into force generation. These results suggest that motoneuron synchronization in human neonates might be an important mechanism for controlling fast limb movements, such as those of primitive reflexes. In addition to help revealing mechanisms of development, the proposed neural interface might monitor children at risk of developing motor disorders.

Locomotor patterns during obstacle avoidance in children with cerebral palsy.

We investigated how early injuries to developing brain affect the interaction of locomotor patterns with the voluntary action required by obstacle clearance. This task requires higher cognitive load and specific anticipatory sensorimotor integration than more automated steady-state gait. To this end, we compared the adaptive gait patterns during obstacle clearance in 40 children with cerebral palsy (CP) (24 diplegic, 16 hemiplegic, 2-12 yr) and 22 typically developing (TD) children (2-12 yr) by analyzing gait kinematics, joint moments during foot elevation, electromyographic (EMG) activity of 11 pairs of bilateral muscles, and muscle modules evaluated by factorization of the EMG signals. The results confirmed generally slower task performance, plus difficulty in motor planning and control in CP. Thus ~30% of diplegic children failed to perform the task. Children with CP demonstrated higher foot lift, smaller range of motion of distal segments, difficulties in properly activating the hamstring muscles at liftoff, and a modified hip strategy when elevating the trailing limb. Basic muscle modules were generally roughly similar to TD patterns, though they showed a limited adaptation. Thus a distinct activation burst in the adaptable muscle module timed to the voluntary task (liftoff) was less evident in CP. Children with CP also showed prolonged EMG burst durations. Impaired obstacle task performance may reflect impaired or less adaptable supraspinal and spinal control of gait when a locomotor task is superimposed with the voluntary movement. Neurorehabilitation of gait in CP may thus be beneficial by adding voluntary tasks such as obstacle clearance during gait performance.NEW & NOTEWORTHY Previous studies mainly evaluated the neuromuscular pattern generation in cerebral palsy (CP) during unobstructed gait. Here we characterized impairments in the obstacle task performance associated with a limited adaptation of the task-relevant muscle module timed to the foot lift during obstacle crossing. Impaired task performance in children with CP may reflect basic developmental deficits in the adaptable control of gait when the locomotor task is superimposed with the voluntary movement.

Quasi-Herglotz functions and convex optimization.

We introduce the set of quasi-Herglotz functions and demonstrate that it has properties useful in the modelling of non-passive systems. The linear space of quasi-Herglotz functions constitutes a natural extension of the convex cone of Herglotz functions. It consists of differences of Herglotz functions and we show that several of the important properties and modelling perspectives are inherited by the new set of quasi-Herglotz functions. In particular, this applies to their integral representations, the associated integral identities or sum rules (with adequate additional assumptions), their boundary values on the real axis and the associated approximation theory. Numerical examples are included to demonstrate the modelling of a non-passive gain medium formulated as a convex optimization problem, where the generating measure is modelled by using a finite expansion of B-splines and point masses.

Neuromechanical adjustments when walking with an aiding or hindering horizontal force.

PURPOSE: Walking against a constant horizontal traction force which either hinders or aids the motion of the centre of mass of the body (COM) will create a discrepancy between the positive and negative work being done by the muscles and may thus affect the mechanics and energetics of walking. We aimed at investigating how this imbalance affects the exchange between potential and kinetic energy of the COM and how its dynamics is related to specific spatiotemporal organisation of motor pool activity in the spinal cord. To understand if and how the spinal cord activation may be associated with COM dynamics, we also compared the neuromechanical adjustments brought on by a horizontal force with published data about those brought on by a slope. METHODS: Ten subjects walked on a treadmill at different speeds with different traction forces. We recorded kinetics, kinematics, and electromyographic activity of 16 lower-limb muscles and assessed the spinal locomotor output by mapping them onto the rostrocaudal location of the motoneuron pools. RESULTS: When walking with a hindering force, the major part of the exchange between potential and kinetic energy of the COM occurs during the first part of stance, whereas with an aiding force exchanges increase during the second part of stance. Those changes occur since limb and trunk orientations remain aligned with the average orientation of the ground reaction force vector. Our results also show the sacral motor pools decreased their activity with an aiding force and increased with a hindering one, whereas the lumbar motor pools increased their engagement both with an aiding and a hindering force. CONCLUSION: Our findings suggest that applying a constant horizontal force results in similar modifications of COM dynamics and spinal motor output to those observed when walking on slopes, consistent with common principles of motor pool functioning and biomechanics of locomotion.

Pelvic movements during walking throughout gestation - the relationship between morphology and kinematic parameters.

BACKGROUND: Many researchers emphasize adaptations following pregnancy. Our purpose was to get more insight into how morphology interacts with the pelvic walking pattern - the segment most prone to the adaptation following altered body demands. METHODS: Thirty women were enrolled. Three experimental sessions were arranged according to the same protocol in the first, second and third trimesters of pregnancy. First, the anthropometric measures were taken, then walking trials at a self-selected speed were registered. At the end of the experimental session the subjects were asked to fill out a questionnaire on pain. FINDINGS: The sagittal plane pelvic range of motion (RoM) significantly increased throughout pregnancy. There were significant positive correlations between pelvic anthropometric dimensions and pelvic tilt and rotation primarily in the third trimester of pregnancy. Significant positive correlations were found between pelvic RoM and thigh circumference. Indicators associated with body mass increase were positively correlated with pelvic obliquity in the second trimester and pelvic tilt and rotation in late pregnancy. It is also worth noting that the individual differences were not related to back pain and that the reported correlations were observed in some but not in all trimesters. INTERPRETATION: Morphological changes following the fetus growth induced increased pelvic tilt and rotation, however, pelvis movements were not associated with back pain. Overall, the results highlight correlations between morphology and pelvis kinematic patterns in some but not in all trimesters.

Differential activation of lumbar and sacral motor pools during walking at different speeds and slopes.

Organization of spinal motor output has become of interest for investigating differential activation of lumbar and sacral motor pools during locomotor tasks. Motor pools are associated with functional grouping of motoneurons of the lower limb muscles. Here we examined how the spatiotemporal organization of lumbar and sacral motor pool activity during walking is orchestrated with slope of terrain and speed of progression. Ten subjects walked on an instrumented treadmill at different slopes and imposed speeds. Kinetics, kinematics, and electromyography of 16 lower limb muscles were recorded. The spinal locomotor output was assessed by decomposing the coordinated muscle activation profiles into a small set of common factors and by mapping them onto the rostrocaudal location of the motoneuron pools. Our results show that lumbar and sacral motor pool activity depend on slope and speed. Compared with level walking, sacral motor pools decrease their activity at negative slopes and increase at positive slopes, whereas lumbar motor pools increase their engagement when both positive and negative slope increase. These findings are consistent with a differential involvement of the lumbar and the sacral motor pools in relation to changes in positive and negative center of body mass mechanical power production due to slope and speed.NEW & NOTEWORTHY In this study, the spatiotemporal maps of motoneuron activity in the spinal cord were assessed during walking at different slopes and speeds. We found differential involvement of lumbar and sacral motor pools in relation to changes in positive and negative center of body mass power production due to slope and speed. The results are consistent with recent findings about the specialization of neuronal networks located at different segments of the spinal cord for performing specific locomotor tasks.

Locomotor coordination in patients with Hereditary Spastic Paraplegia.

Locomotion is a complex behaviour that requires the coordination of multiple body segments and muscle groups. Here we investigated how the weakness and spasticity in individuals with Hereditary Spastic Paraplegia (HSP) affect the coordination patterns of the lower limbs. We analysed kinematics and electromyographic (EMG) activity from 12 leg muscles in 21 persons with HSP and 20 control subjects at matched walking speeds. To assess the locomotor coordination, we examined the covariation between thigh, shank and foot elevation angles by means of principal component analysis and the modular organization of EMG patterns using the non-negative matrix factorization algorithm. The characteristic features of the HSP gait consisted in changes of the elevation angles covariation, the shape of the gait loop, reduced range of motion of the distal segments and significantly lower foot lift. The EMG factorization analysis revealed a comparable structure of the motor output between HSP and control groups, but significantly wider basic temporal patterns associated with muscles innervated from the sacral spinal segments in HSP. Overall, the applied methodology highlighted the impact of the corticospinal degeneration and spasticity on the coordination of distal limb segments and basic muscle modules associated with distal spinal segments.

Synergistic influences of sensory and central stimuli on non-voluntary rhythmic arm movements.

In recent years, neuromodulation of the cervical spinal circuitry has become an area of interest for investigating rhythmogenesis of the human spinal cord and interaction between cervical and lumbosacral circuitries, given the involvement of rhythmic arm muscle activity in many locomotor tasks. We have previously shown that arm muscle vibrostimulation can elicit non-voluntary upper limb oscillations in unloading body conditions. Here we investigated the excitability of the cervical spinal circuitry by applying different peripheral and central stimuli in healthy humans. The rationale for applying combined stimuli is that the efficiency of only one stimulus is generally limited. We found that low-intensity electrical stimulation of the superficial arm median nerve can evoke rhythmic arm movements. Furthermore, the movements were enhanced by additional peripheral stimuli (e.g., arm muscle vibration, head turns or passive rhythmic leg movements). Finally, low-frequency transcranial magnetic stimulation of the motor cortex significantly facilitated rhythmogenesis. The findings are discussed in the general framework of a brain-spinal interface for developing adaptive central pattern generator-modulating therapies.

Progressive changes in walking kinematics throughout pregnancy-A follow up study.

BACKGROUND: Progressive weight gain and changes in its distribution following pregnancy may be challenging for the gravidas' ability to move in a stable way. RESEARCH QUESTION: How is gait kinematics changing throughout pregnancy and to what extend is it affected by physical activity level and energy balance? METHODS: 30 women were enrolled. Three experimental sessions were arranged according to the same protocol in the first, second and third trimesters of pregnancy. Walking kinematics at a self-selected speed was registered. The total physical activity (TPA) was assessed from the subjects' questionnaires. Energy balance ('positive', 'balanced' or 'negative') was estimated as the difference between dietary energy intake and energy expenditure during 7 days. RESULTS: No significant differences were found in the spatiotemporal variables between experimental sessions. However, the gait analysis revealed significant increments in the single support and base of support (BoS) measures. Generally, the sagittal plane mobility of the lower limb joints did not differ, however, the pelvic tilt increased in late pregnancy. The hip and pelvis angles were significantly different over the gait cycle throughout gestation. The 'balanced' energy was dominant in the first trimester although the relative number of participants with negative balance increased over pregnancy. Overall, gait parameters were independent of the energy balance. However, significant correlation was found between gait parameters, such as BoS, velocity, stride length, and TPA in the advanced pregnancy. SIGNIFICANCE: The longitudinal assessment of walking kinematics demonstrates few changes adopted to accommodate for pregnancy. The enlargement of BoS is considered as a strategy to provide safety and stability. The increased pelvic tilt is likely to compensate for changes in the body mass distribution. The physical activity correlates with the BoS measures and stride length and thus may be important for enhancing gait stability.

Kinematic patterns while walking on a slope at different speeds.

During walking, the elevation angles of the thigh, shank, and foot (i.e., the angle between the segment and the vertical) covary along a characteristic loop constrained on a plane. Here, we investigate how the shape of the loop and the orientation of the plane, which reflect the intersegmental coordination, change with the slope of the terrain and the speed of progression. Ten subjects walked on an inclined treadmill at different slopes (between -9° and +9°) and speeds (from 0.56 to 2.22 m/s). A principal component analysis was performed on the covariance matrix of the thigh, shank, and foot elevation angles. At each slope and speed, the variance accounted for by the two principal components was >99%, indicating that the planar covariation is maintained. The two principal components can be associated to the limb orientation (PC1*) and the limb length (PC2*). At low walking speeds, changes in the intersegmental coordination across slopes are characterized mainly by a change in the orientation of the covariation plane and in PC2* and to a lesser extent, by a change in PC1*. As speed increases, changes in the intersegmental coordination across slopes are more related to a change in PC1 *, with limited changes in the orientation of the plane and in PC 2*. Our results show that the kinematic patterns highly depend on both slope and speed. NEW & NOTEWORTHY In this paper, changes in the lower-limb intersegmental coordination during walking with slope and speed are linked to changes in the trajectory of the body center of mass. Modifications in the kinematic pattern with slope depend on speed: at slow speeds, the net vertical displacement of the body during each step is related to changes in limb length and orientation. When speed increases, the vertical displacement is mostly related to a change in limb orientation.

Early manifestation of arm-leg coordination during stepping on a surface in human neonates.

The accomplishment of mature locomotor movements relies upon the integrated coordination of the lower and upper limbs and the trunk. Human adults normally swing their arms and a quadrupedal limb coordination persists during bipedal walking despite a strong corticospinal control of the upper extremities that allows to uncouple this connection during voluntary activities. Here we investigated arm-leg coordination during stepping responses on a surface in human neonates. In eight neonates, we found the overt presence of alternating arm-leg oscillations, the arms moving up and down in alternation with ipsilateral lower limb movements. These neonates moved the diagonal limbs together, and the peak of the arm-to-trunk angle (i.e., maximum vertical excursion of the arm) occurred around the end of the ipsilateral stance phase, as it occurs during typical adult walking. Although episodes of arm-leg coordination were sporadic in our sample of neonates, their presence provides significant evidence for a neural coupling between the upper and lower limbs during early ontogenesis of locomotion in humans.

Differential changes in the spinal segmental locomotor output in Hereditary Spastic Paraplegia.

OBJECTIVE: A comprehensive treatment of Hereditary Spastic Paraplegia (HSP) should consider the specific pathophysiological changes in the spinal cord. Here we reported a detailed characterization of the spinal motoneuronal output in HSP during locomotion. METHODS: We recorded kinematics and electromyographic (EMG) activity of 12 leg muscles in 29 patients with pure forms of HSP and compared them with 30 controls while walking at matched speeds. We assessed the spinal locomotor output by evaluating EMG patterns and by mapping them onto the rostrocaudal location of the spinal motoneuron pools. RESULTS: The activity profiles of muscles innervated from the sacral segments were significantly wider in patients. Similarly, spinal maps revealed a tendency for spreading the main loci of activation, involving initially the sacral segments and, at more severe stages, the lumbar segments. CONCLUSIONS: The degeneration of the corticospinal tract in HSP is associated with a widening of spinal locomotor output spreading from caudal to rostral segments. SIGNIFICANCE: The findings highlight pathophysiologically relevant differential changes in the spinal locomotor output in HSP related to the specific innervation of muscles in the spinal cord, and might be helpful for developing future therapeutic strategies and identifying physiological markers of the disease.

Corrigendum: Foot Placement Characteristics and Plantar Pressure Distribution Patterns during Stepping on Ground in Neonates.

[This corrects the article on p. 784 in vol. 8, PMID: 29066982.].

Foot Placement Characteristics and Plantar Pressure Distribution Patterns during Stepping on Ground in Neonates.

Stepping on ground can be evoked in human neonates, though it is rather irregular and stereotyped heel-to-toe roll-over pattern is lacking. Such investigations can provide insights into the role of contact- or load-related proprioceptive feedback during early development of locomotion. However, the detailed characteristics of foot placements and their association with motor patterns are still incompletely documented. We elicited stepping in 33 neonates supported on a table. Unilateral limb kinematics, bilateral plantar pressure distribution and EMG activity from up to 11 ipsilateral leg muscles were recorded. Foot placement characteristics in neonates showed a wide variation. In ~25% of steps, the swinging foot stepped onto the contralateral foot due to generally small step width. In the remaining steps with separate foot placements, the stance phase could start with forefoot (28%), midfoot (47%), or heel (25%) touchdowns. Despite forefoot or heel initial contacts, the kinematic and loading patterns markedly differed relatively to toe-walking or adult-like two-peaked vertical force profile. Furthermore, while the general stepping parameters (cycle duration, step length, range of motion of proximal joints) were similar, the initial foot contact was consistently associated with specific center-of-pressure excursion, range of motion in the ankle joint, and the center-of-activity of extensor muscles (being shifted by ~5% of cycle toward the end of stance in the "heel" relative to "forefoot" condition). In sum, we found a variety of footfall patterns in conjunction with associated changes in motor patterns. These findings suggest the potential contribution of load-related proprioceptive feedback and/or the expression of variations in the locomotor program already during early manifestations of stepping on ground in human babies.

Planar covariance of upper and lower limb elevation angles during hand-foot crawling in healthy young adults.

Habitual quadrupeds have been shown to display a planar covariance of segment elevation angle waveforms in the fore and hind limbs during many forms of locomotion. The purpose of the current study was to determine if humans generate similar patterns in the upper and lower limbs during hand-foot crawling. Nine healthy young adults performed hand-foot crawling on a treadmill at speeds of 1, 2, and 3 km/h. A principal component analysis (PCA) was applied to the segment elevation angle waveforms for the upper (upper arm, lower arm, and hand) and lower (thigh, shank, and foot) limbs separately. The planarity of the elevation angle waveforms was determined using the sum of the variance explained by the first two PCs and the orientation of the covariance plane was quantified using the direction cosines of the eigenvector orthogonal to the plane, projected upon each of the segmental semi-axes. Results showed that planarity of segment elevation angles was maintained in the upper and lower limbs (explained variance >97%), although a slight decrease was present in the upper limb when crawling at 3 km/h. The orientation of the covariance plane was highly limb-specific, consistent with animal studies and possibly related to the functional neural control differences between the upper and lower limbs. These results may suggest that the motor patterns stored in the central nervous system for quadrupedal locomotion may be retained through evolution and may still be exploited when humans perform such tasks.

Human cervical spinal cord circuitry activated by tonic input can generate rhythmic arm movements.

The coordination between arms and legs during human locomotion shares many features with that in quadrupeds, yet there is limited evidence for the central pattern generator for the upper limbs in humans. Here we investigated whether different types of tonic stimulation, previously used for eliciting stepping-like leg movements, may evoke nonvoluntary rhythmic arm movements. Twenty healthy subjects participated in this study. The subject was lying on the side, the trunk was fixed, and all four limbs were suspended in a gravity neutral position, allowing unrestricted low-friction limb movements in the horizontal plane. The results showed that peripheral sensory stimulation (continuous muscle vibration) and central tonic activation (postcontraction state of neuronal networks following a long-lasting isometric voluntary effort, Kohnstamm phenomenon) could evoke nonvoluntary rhythmic arm movements in most subjects. In ∼40% of subjects, tonic stimulation elicited nonvoluntary rhythmic arm movements together with rhythmic movements of suspended legs. The fact that not all participants exhibited nonvoluntary limb oscillations may reflect interindividual differences in responsiveness of spinal pattern generation circuitry to its activation. The occurrence and the characteristics of induced movements highlight the rhythmogenesis capacity of cervical neuronal circuitries, complementing the growing body of work on the quadrupedal nature of human gait.

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.

Locomotor patterns in cerebellar ataxia.

Several studies have demonstrated how cerebellar ataxia (CA) affects gait, resulting in deficits in multijoint coordination and stability. Nevertheless, how lesions of cerebellum influence the locomotor muscle pattern generation is still unclear. To better understand the effects of CA on locomotor output, here we investigated the idiosyncratic features of the spatiotemporal structure of leg muscle activity and impairments in the biomechanics of CA gait. To this end, we recorded the electromyographic (EMG) activity of 12 unilateral lower limb muscles and analyzed kinematic and kinetic parameters of 19 ataxic patients and 20 age-matched healthy subjects during overground walking. Neuromuscular control of gait in CA was characterized by a considerable widening of EMG bursts and significant temporal shifts in the center of activity due to overall enhanced muscle activation between late swing and mid-stance. Patients also demonstrated significant changes in the intersegmental coordination, an abnormal transient in the vertical ground reaction force and instability of limb loading at heel strike. The observed abnormalities in EMG patterns and foot loading correlated with the severity of pathology [International Cooperative Ataxia Rating Scale (ICARS), a clinical ataxia scale] and the changes in the biomechanical output. The findings provide new insights into the physiological role of cerebellum in optimizing the duration of muscle activity bursts and the control of appropriate foot loading during locomotion.