Evolutionary and functional adaptations of morphology and postural tone of the spine and trunk are intrinsically shaped by the field of gravity in which humans move. Gravity also significantly impacts the timing and levels of neuromuscular activation, particularly in foot-support interactions. During step-to-step transitions, the centre of mass velocity must be redirected from downwards to upwards. When walking upright, this redirection is initiated by the trailing leg, propelling the body forward and upward before the foot contact of the leading leg, defined as an anticipated transition. In this study, we investigate the neuromechanical adjustments when walking with a bent posture. Twenty adults walked on an instrumented treadmill at 4 km/h under normal (upright) conditions and with varying degrees of anterior trunk flexion (10, 20, 30, and 40°). We recorded lower-limb kinematics, ground reaction forces under each foot, and the electromyography activity of five lower-limb muscles. Our findings indicate that with increasing trunk flexion, there is a lack of these anticipatory step-to-step transitions, and the leading limb performs the redirection after the ground collision. Surprisingly, attenuating distal extensor muscle activity at the end of stance is one of the main impacts of trunk flexion. Our observations may help to understand the physiological mechanisms and biomechanical regulations underlying our tendency toward an upright posture, as well as possible motor control disturbances in some diseases associated with trunk orientation problems.

Background As foot constitutes the base of support for the whole body, the pregnancy-related anthropometric changes can result in adaptive plantar pressure alterations. The present study aimed to investigate how pregnancy affects foot loading pattern in gait, and if it is related to body adjustments to growing foetus that occur in the course of pregnancy. Methods A prospective longitudinal study included 30 women. Three experimental sessions in accordance with the same procedure were carried out in the first, second and third trimesters of pregnancy. First, the anthropometric measures of the body mass and waist circumference were taken. Then walking trials at a self-selected speed along a ~6-m walkway were registered with the FreeMED force platform (Sensor Medica, Italy). Vertical foot pressure was recorded by the force plate located in the middle of the walkway. Findings The correlation of individual foot loading parameters across different trimesters was relatively high. Nevertheless, our results revealed a longitudinal foot arch flattening with the strongest effect in late pregnancy ( P = 0.01). The anthropometric characteristics also influenced the foot loading pattern depending on the phase of pregnancy. In particular, arch flattening correlated with the body mass in all trimesters (r≥0.44, P ≤0.006) while the medial-lateral loading index correlated only in the first (r = 0.45, P = 0.005) and second (r = 0.36, P = 0.03) trimesters. Waist circumference changes significantly influenced dynamic arch flattening but only in the late pregnancy (r≥0.46, P ≤0.004). In the third trimester, a small though significant increase in the right foot angle was observed ( P = 0.01). Interpretation The findings provided the characteristics of the relative foot areas loading throughout pregnancy. Growing abdominal size increases the risk of medial arch flattening, which can result in less stable gait. The observed increase in foot angle in late pregnancy may constitute a strategy to enhance gait stability.

Terrestrial locomotion requires coordinated bilateral activation of limb muscles, with left–right alternation in walking or running, and synchronous activation in hopping or skipping. The neural mechanisms involved in interlimb coordination at birth are well known in different mammalian species, but less so in humans. Here, 46 neonates (of either sex) performed bilateral and unilateral stepping with one leg blocked in different positions. By recording EMG activities of lower-limb muscles, we observed episodes of left–right alternating or synchronous coordination. In most cases, the frequency of EMG oscillations during sequences of consecutive steps was approximately similar between the two sides, but in some cases it was considerably different, with episodes of 2:1 interlimb coordination and episodes of activity deletions on the blocked side. Hip position of the blocked limb significantly affected ipsilateral, but not contralateral, muscle activities. Thus, hip extension backward engaged hip flexor muscle, and hip flexion engaged hip extensors. Moreover, the sudden release of the blocked limb in the posterior position elicited the immediate initiation of the swing phase of the limb, with hip flexion and a burst of an ankle flexor muscle. Extensor muscles showed load responses at midstance. The variable interlimb coordination and its incomplete sensory modulation suggest that the neonatal locomotor networks do not operate in the same manner as in mature locomotion, also because of the limited cortical control at birth. These neonatal mechanisms share many properties with spinal mammalian preparations (i.e., independent pattern generators for each limb, and for flexor and extensor muscles, load, and hip position feedback). SIGNIFICANCE STATEMENT Bilateral coupling and reciprocal activation of flexor and extensor burst generators represent the fundamental mechanisms used by mammalian limbed locomotion. Considerable progress has been made in deciphering the early development of the spinal networks and left–right coordination in different mammals, but less is known about human newborns. We compared bilateral and unilateral stepping in human neonates, where cortical control is still underdeveloped. We found neonatal mechanisms that share many properties with spinal mammalian preparations (i.e., independent pattern generators for each limb, the independent generators for flexor and extensor muscles, load, and hip-position feedback. The variable interlimb coordination and its incomplete sensory modulation suggest that the human neonatal locomotor networks do not operate in the same manner as in mature locomotion.

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.

ABSTRACT 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.

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.

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.

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.

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.

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.

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.

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.

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.

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. (C) 2018 International Federation of Clinical Neurophysiology. Published by Elsevier B.V. All rights reserved.

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.

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.

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

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.