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During running, the mechanical energy of the centre of mass of the body (COM) oscillates throughout the step like a spring-mass system, where part of its mechanical energy is stored during negative phases to be released during the following positive phases. This storage-release of energy improves muscle-tendon efficiency, which is related to lower-limb stiffness. This study explores the effect of sports background on the bouncing mechanism, by examining differences in stiffness and step spatiotemporal parameters between swimmers and football athletes. All athletes performed three consecutive running bouts on an instrumented treadmill at three different speeds (3.9, 4.4 and 5.0 m·s−1). The ground reaction forces were recorded. Vertical stiffness and step spatiotemporal parameters were analysed and compared using a two-way ANOVA. Vertical stiffness of football players was on average 21.0 ± 1.1% higher than swimmers. The modification of step spatiotemporal parameters also suggests a more elastic rebound by increasing the stretch of tendons relative to muscle within muscle–tendon units in football players. Compared to swimmers, they (1) decrease the effective contact time by 9.7 ± 2.4% and (2) decrease the duration of the push by 15.0 ± 6.4%, suggesting that background training adaptations influence spring-mass behaviour during running. [ABSTRACT FROM AUTHOR]

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 foot contact of the leading leg, defined as an anticipated transition. In this study, we investigated the neuromechanical adjustments when walking with a bent posture. Twenty adults walked on an instrumented treadmill at 4 km h-1 under normal (upright) conditions and with varying degrees of anterior trunk flexion (10, 20, 30 and 40 deg). 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 us to understand the physiological mechanisms and biomechanical regulations underlying our tendency towards an upright posture, as well as possible motor control disturbances in some diseases associated with trunk orientation problems., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

In laboratory settings, human locomotion encounters minimal opposition from air resistance. However, moving in nature often requires overcoming airflow. Here, the drag force exerted on the body by different headwind or tailwind speeds (between 0 and 15 m·s -1 ) was measured during walking at 1.5 m·s -1 and running at 4 m·s -1 . To our knowledge, the biomechanical effect of drag in human locomotion has only been evaluated by simulations. Data were collected on eight male subjects using an instrumented treadmill placed in a wind tunnel. From the ground reaction forces, the drag and external work done to overcome wind resistance and to sustain the motion of the center of mass of the body were measured. Drag increased with wind speed: a 15 m·s -1 headwind exerted a drag of ∼60 N in walking and ∼50 N in running. The same tailwind exerted -55 N of drag in both gaits. At this wind speed, the work done to overcome the airflow represented ∼80% of the external work in walking and ∼50% in running. Furthermore, in the presence of fast wind speeds, subjects altered their drag area ( C d A ) by adapting their posture to limit the increase in air friction. Moving in the wind modified the ratio between positive and negative external work performed. The modifications observed when moving with a head- or tailwind have been compared with moving uphill or downhill. The present findings may have implications for optimizing aerodynamic performance in competitive running, whether in sprints or marathons. NEW & NOTEWORTHY This is the first study to assess the biomechanical adaptations to a wide range of wind speeds inside a wind tunnel. Humans increase their mechanical work and alter their drag area ( C d A ) by adapting their posture when walking and running against increasing head and tailwinds. The observed drag force applied to the subject is different between walking and running at similar headwind speeds.

This study aims to investigate the relationship between soft tissue energy dissipation and leg stiffness during running. Eight recreational healthy male runners (age: 22.2 ± 1.0 years; height: 1.84 ± 0.03 m; mass: 73.7 ± 5.7 kg) were asked to run at different speeds and step frequencies. Their soft tissue energy dissipation was estimated by the difference between the total mechanical work of the body, measured as the work done to move the body centre of mass relative to the surroundings plus the work to move the limbs relative to the body centre of mass, and lower-limb joint work. A mass-spring model with an actuator was used to analyse the force-length curve of the bouncing mechanism of running. In this way, the stiffness and damping coefficient were assessed at each speed and step frequency. Pearson's correlations were used to describe the relationship between the deviation from the spring-mass model and soft tissue energy fluctuations. The soft tissue dissipation was found to be significantly influenced by step frequency, with both positive and negative work phases decreasing when step frequency increases. Moreover, deviation from a spring-mass model was positively associated with the amount of soft tissue dissipation ( r > 0.6). The findings emphasize the substantial role of soft tissues in dissipating or returning energy during running, behaving in a damped-elastic manner. Also, we introduce a novel approach for evaluating the elastic rebound of the body during running. The insights gained may have broad implications for assessing running mechanics, with potential applications in various contexts., Competing Interests: We declare we have no competing interests., (© 2024 The Authors.)

When running on a curve, the lower limbs interact with the ground to redirect the trajectory of the centre of mass of the body (CoM). The goal of this paper is to understand how the trajectory of the CoM and the work done to maintain its movements relative to the surroundings (Wcom) are modified as a function of running speed and radius of curvature. Eleven participants ran at different speeds on a straight line and on circular curves with a 6 m and 18 m curvature. The trajectory of the CoM and Wcom were calculated using force-platforms measuring the ground reaction forces and infrared cameras recording the movements of the pelvis. To follow a circular path, runners overcompensate the rotation of their trajectory during contact phases. The deviation from the circular path increases when the radius of curvature decreases and speed increases. Interestingly, an asymmetry between the inner and outer lower limbs emerges as speed increases. The method to evaluate Wcom on a straight-line was adapted using a referential that rotates at heel strike and remains fixed during the whole step cycle. In an 18 m radius curve and at low speeds on a 6 m radius, Wcom changes little compared to a straight-line run. Whereas at 6 m s-1 on a 6 m radius, Wcom increases by ~25%, due to an augmentation in the work to move the CoM laterally. Understanding these adaptations provides valuable insight for sports sciences, aiding in optimizing training and performance in sports with multidirectional movements., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2024 Mesquita et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Purpose: Running at a given speed can be achieved by taking large steps at a low frequency or on the contrary by taking small steps at a high frequency. The consequences of a change in step frequency, at a fixed speed, affects the stiffness of the lower limb differently. In this study, we compared the running mechanics and kinematics at different imposed step frequencies (from 2 step s -1 to 3.6 step s -1 ) to understand the relationship between kinematic and kinetic parameters., Methods: Eight recreational male runners ran on a treadmill at 5 different speeds and 5 different step frequencies. The lower-limb segment motion and the ground reaction forces were recorded. Mechanical powers, general gait parameters, lower-limb movements and coordination were investigated., Results: At low step frequencies, in order to limit the magnitude of the ground reaction force, the vertical stiffness is reduced and thus runners deviate from an elastic rebound. At high step frequencies, the stiffness is increased and the elastic rebound is optimised in its ability to absorb and restore energy during the contact phase., Conclusion: We studied the consequences of a change in step frequency on the bouncing mechanics of running. We showed that the lower limb stiffness and the intersegmental coordination of the lower-limb segments are affected by running step frequency rather than speed. The runner rather adapts their lower limb stiffness to match a step frequency for a given speed than the opposite., (© 2023. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

The mechanical and metabolic responses of walking by obese children are not yet well understood. The objectives of this study were (1) to compare the pendular mechanism (recovery, phase shift by α and β values, and ratio between forward and vertical mechanical work), the maximum possible elastic energy usage and the bilateral coordination during walking between non-obese and obese children, and (2) to verify if the bilateral coordination could contribute to understanding the pendular mechanism and elastic energy usage in these populations. Nine obese (six female, 8.7 ± 0.5 years, 1.38 ± 0.04 m, 44.4 ± 6.3 kg and 24.1 ± 3.50 kg/m 2 ) and eight non-obese (four female, 7.4 ± 0.5 years, 1.31 ± 0.08 m, 26.6 ± 2.1 kg and 16.4 ± 1.40 kg/m 2 ) children were analysed during walking on a treadmill at five speeds: 1, 2, 3, 4 and 5 km/h. The results indicated that although the mechanical energy response of the centre of mass during walking is similar between obese and non-obese children, the obese children showed a lower pendulum-like mechanism and greater elastic energy usage during level walking. Therefore, obese children seem to use more elastic energy during walking compared to non-obese children, which may be related to their apparent higher positive work production during the double support phase. Finally, bilateral coordination presented high values at slow speeds in both groups and requires further attention due to its association with falls. NEW FINDINGS: What is the central question of this study? Are there any differences of the pendular and elastic mechanisms and bilateral coordination during walking between non-obese and obese children? What is the main finding and its importance? To our knowledge, this study is the first to analyse the mechanical energy usage and the bilateral coordination of obese and non-obese children during walking. Obese children had a lower pendular recovery mechanism and used more elastic energy compared to non-obese children. The bilateral coordination was higher at slow speeds in both groups and requires further attention due to its association with falls., (© 2023 The Authors. Experimental Physiology published by John Wiley & Sons Ltd on behalf of The Physiological Society.)

The development and acquisition of mature walking in children is multifactorial, depending among others on foot interaction with the ground, body dynamics and the knowledge of the 'rules' stemming from the gravity field. Indeed, each step the velocity of the centre of mass must be redirected upwards. This redirection may be initiated by the trailing leg, propulsing forward and upward the body before foot contact, or later by the loading limb after the contact with the ground. While it has been suggested that mature walking develops slowly from first independent steps to about 7 years of age, it is still unknown how children acquire the appropriate loading and propulsion forces during the step-to-step transition. To answer that question, twenty-four children (from 3 to 12 years old) and twelve young adults (from 20 to 27 years old) walked on force platforms at different walking speed. The ground reaction forces under each foot were recorded and the vertical velocity of the centre of mass of the body was computed. With decreasing age and increasing velocity (or Froude number), the occurrence of unanticipated transition is higher, related to a different ratio between the vertical support of the front and back leg. The different transition strategy observed in children indicates that body weight transfer from one limb to the other is not fully mature at 12 years old., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

Purpose: Humans are amongst few animals that step first on the heel, and then roll on the ball of the foot and toes. While this heel-to-toe rolling pattern has been shown to render an energetic advantage during walking, the effect of different foot contact strategies, on the neuromuscular control of adult walking gaits has received less attention. We hypothesised that deviating from heel-to-toe rolling pattern affects the energy transduction and weight acceptance and re-propulsive phases in gait along with the modification of spinal motor activity., Methods: Ten subjects walked on a treadmill normally, then placed their feet flat on the ground at each step and finally walked on the balls of the feet., Results: Our results show that when participants deviate from heel-to-toe rolling pattern strategy, the mechanical work increases on average 85% higher (F = 15.5; p < 0.001), mainly linked to a lack of propulsion at late stance. This modification of the mechanical power is related to a differential involvement of lumbar and sacral segment activation. Particularly, the delay between the major bursts of activation is on average 65% smaller, as compared to normal walking (F = 43.2; p < 0.001)., Conclusion: Similar results are observable in walking plantigrade animals, but also at the onset of independent stepping in toddlers, where the heel-to-toe rolling pattern is not yet established. These indications seem to bring arguments to the fact that the rolling of the foot during human locomotion has evolved to optimise gait, following selective pressures from the evolution of bipedal posture., (© 2023. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

With extended life expectancy, the quality of life of elders is a priority. Loss of mobility, increased morbidity and risks of falls have dramatic individual and societal impacts. Here we consider the age-related modifications of gait, from a biomechanical and neurophysiological perspective. Among the many factors of frailty involved (e.g., metabolic, hormonal, immunological), loss of muscle strength and neurodegenerative changes inducing slower muscle contraction may play a key role. We highlight that the impact of the multifactorial age-related changes in the neuromuscular systems results in common features of gait in the immature gait of infants and older adults. Besides, we also consider the reversibility of age-related neuromuscular deterioration by, on the one hand, exercise training, and the other hand, novel techniques such as direct spinal stimulation (tsDCS)., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. The handling editor LP-T declared a past co-authorship with the author AD., (Copyright © 2023 Nùñez-Lisboa, Valero-Breton and Dewolf.)

Switching locomotion direction is a common task in daily life, and it has been studied extensively in healthy people. Little is known, however, about the locomotor adjustments involved in changing locomotion direction from forward (FW) to sideways (SW) in children with cerebral palsy (CP). The importance of testing the ability of children with CP in this task lies in the assessment of flexible, adaptable adjustments of locomotion as a function of the environmental context. On the one hand, the ability of a child to cope with novel task requirements may provide prognostic cues as to the chances of modifying the gait adaptively. On the other hand, challenging the child with the novel task may represent a useful rehabilitation tool to improve the locomotor performance. SW is an asymmetrical locomotor task and requires a differential control of right and left limb muscles. Here, we report the results of a cross-sectional study comparing FW and SW in 27 children with CP (17 diplegic, 10 hemiplegic, 2-10 years) and 18 age-matched typically developing (TD) children. We analyzed gait kinematics, joint moments, EMG activity of 12 pairs of bilateral muscles, and muscle modules evaluated by factorization of EMG signals. Task performance in several children with CP differed drastically from that of TD children. Only 2/3 of children with CP met the primary outcome, i.e. they succeeded to step sideways, and they often demonstrated attempts to step forward. They tended to rotate their trunk FW, cross one leg over the other, flex the knee and hip. Moreover, in contrast to TD children, children with CP often exhibited similar motor modules for FW and SW. Overall, the results reflect developmental deficits in the control of gait, bilateral coordination and adjustment of basic motor modules in children with CP. We suggest that the sideways (along with the backward) style of locomotion represents a novel rehabilitation protocol that challenges the child to cope with novel contextual requirements., (© 2023. The Author(s).)

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., (Copyright © 2022 the authors.)

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., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2021. Published by The Company of Biologists Ltd.)

Locomotor movements are accommodated to various surface conditions by means of specific locomotor adjustments. This study examined underlying age-related differences in neuromuscular control during level walking and on a positive or negative slope, and during stepping upstairs and downstairs. Ten elderly and eight young adults walked on a treadmill at two different speeds and at three different inclinations (0°, +6°, and -6°). They were also asked to ascend and descend stairs at self-selected speeds. Full body kinematics and surface electromyography of 12 lower-limb muscles were recorded. We compared the intersegmental coordination, muscle activity, and corresponding modifications of spinal motoneuronal output in young and older adults. Despite great similarity between the neuromuscular control of young and older adults, our findings highlight subtle age-related differences in all conditions, potentially reflecting systematic age-related adjustments of the neuromuscular control of locomotion across various support surfaces. The main distinctive feature of walking in older adults is a significantly wider and earlier activation of muscles innervated by the sacral segments. These changes in neuromuscular control are reflected in a reduction or lack of propulsion observed at the end of stance in older adults at different slopes, with the result of a delay in the timing of redirection of the centre-of-mass velocity and of an unanticipated step-to-step transition strategy., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Dewolf, Sylos-Labini, Cappellini, Zhvansky, Willems, Ivanenko and Lacquaniti.)

New Findings: What is the topic of this review? This narrative review explores past and recent findings on the mechanical determinants of energy cost during human locomotion, obtained by using a mechanical approach based on König's theorem (Fenn's approach). What advances does it highlight? Developments in analytical methods and their applications allow a better understanding of the mechanical-bioenergetic interaction. Recent advances include the determination of 'frictional' internal work; the association between tendon work and apparent efficiency; a better understanding of the role of energy recovery and internal work in pathological gait (amputees, stroke and obesity); and a comprehensive analysis of human locomotion in (simulated) low gravity conditions., Abstract: During locomotion, muscles use metabolic energy to produce mechanical work (in a more or less efficient way), and energetics and mechanics can be considered as two sides of the same coin, the latter being investigated to understand the former. A mechanical approach based on König's theorem (Fenn's approach) has proved to be a useful tool to elucidate the determinants of the energy cost of locomotion (e.g., the pendulum-like model of walking and the bouncing model of running) and has resulted in many advances in this field. During the past 60 years, this approach has been refined and applied to explore the determinants of energy cost and efficiency in a variety of conditions (e.g., low gravity, unsteady speed). This narrative review aims to summarize current knowledge of the role that mechanical work has played in our understanding of energy cost to date, and to underline how recent developments in analytical methods and their applications in specific locomotion modalities (on a gradient, at low gravity and in unsteady conditions) and in pathological gaits (asymmetric gait pathologies, obese subjects and in the elderly) could continue to push this understanding further. The recent in vivo quantification of new aspects that should be included in the assessment of mechanical work (e.g., frictional internal work and elastic contribution) deserves future research that would improve our knowledge of the mechanical-bioenergetic interaction during human locomotion, as well as in sport science and space exploration., (© 2021 The Authors. Experimental Physiology © 2021 The Physiological Society.)

Previous studies found significant modification in spatiotemporal parameters of backward walking in healthy older adults, but the age-related changes in the neuromuscular control have been considered to a lesser extent. The present study compared the intersegmental coordination, muscle activity and corresponding modifications of spinal montoneuronal output during both forward and backward walking in young and older adults. Ten older and ten young adults walked forward and backward on a treadmill at different speeds. Gait kinematics and EMG activity of 14 unilateral lower-limb muscles were recorded. As compared to young adults, the older ones used shorter steps, a more in-phase shank and foot motion, and the activity profiles of muscles innervated from the sacral segments were significantly wider in each walking condition. These findings highlight age-related changes in the neuromuscular control of both forward and backward walking. A striking feature of backward walking was the differential organization of the spinal output as compared to forward gait. In addition, the resulting spatiotemporal map patterns also characterized age-related changes of gait. Finally, modifications of the intersegmental coordination with aging were greater during backward walking. On the whole, the assessment of backward walk in addition to routine forward walk may help identifying or unmasking neuromuscular adjustments of gait to aging., Competing Interests: The authors have declared that no competing interests exist.

This mini-review focuses on the emergence of locomotor-related movements in early infancy. In particular, we consider multiples precursor behaviors of locomotion as a manifestation of the development of the neuronal networks and their link in the establishment of precocious locomotor skills. Despite the large variability of motor behavior observed in human babies, as in animals, afferent information is already processed to shape the behavior to specific situations and environments. Specifically, we argue that the closed-loop interaction between the neural output and the physical dynamics of the mechanical system should be considered to explore the complexity and flexibility of pattern generation in human and animal neonates., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Dewolf, Sylos Labini, Ivanenko and Lacquaniti.)

The first years of life represent an important phase of maturation of the central nervous system, processing of sensory information, posture control and acquisition of the locomotor function. Cerebral palsy (CP) is the most common group of motor disorders in childhood attributed to disturbances in the fetal or infant brain, frequently resulting in impaired gait. Here we will consider various findings about functional maturation of the locomotor output in early infancy, and how much the dysfunction of gait in children with CP can be related to spinal neuronal networks vs. supraspinal dysfunction. A better knowledge about pattern generation circuitries in infancy may improve our understanding of developmental motor disorders, highlighting the necessity for regulating the functional properties of abnormally developed neuronal locomotor networks as a target for early sensorimotor rehabilitation. Various clinical approaches and advances in biotechnology are also considered that might promote acquisition of the locomotor function in infants at risk for locomotor delays., (Copyright © 2020 Cappellini, Sylos-Labini, Dewolf, Solopova, Morelli, Lacquaniti and Ivanenko.)

Purpose: Intra-limb and muscular coordination during gait are the result of the organisation of the neuromuscular system, which have been widely studied on a flat terrain. Environmental factors, such as the inclination of the terrain, is a challenge for the postural control system to maintain balance. Therefore, we hypothesised that the central nervous system flexibly modifies its control strategies during locomotion on slopes., Methods: Ten subjects walked on an inclined treadmill at different slopes (from - 9° to + 9°) and speeds (from 0.56 to 2.22 m s -1 ). Intra-limb coordination was investigated via the Continuous Relative Phase, whereas muscular coordination was investigated by decomposing the coordinated muscle activation profiles into Basic Activation Patterns., Results: A greater stride to stride variability of kinematics was observed during walking on slopes, as compared to walking on the level. On positive slopes, the stride period and width present a greater variability without modification of the time-pattern of the muscular activation and of the variability of intersegmental coordination. On negative slopes, the stride width is larger, the variability of the stride period and of the inter-segmental coordination is greater and the basic activation patterns become broader, especially at slow speeds., Conclusion: Our findings suggest that the control strategy of downhill walking corresponds to a more conservative gait pattern, which could be adopted to lower the risk of falling at the cost of a greater energy consumption. In uphill walking, where metabolic demands are high, the strategy adopted may be planned to minimise energy expenditure.

Purpose: Much like running on a slope, running against/with a horizontal traction force which either hinders/aids the forward motion of the runner creates a shift in the positive and negative muscular work, which in turn modifies the bouncing mechanism of running. The purpose of the study is to (1) investigate the energy changes of the centre of mass and the storage/release of energy throughout the step during running associated with speed and increasing hindering and aiding traction forces; and (2) compare these changes to those observed when running on a slope., Methods: Ground reaction forces were measured on eight subjects running on an instrumented treadmill against different traction forces at different speeds., Results: As compared to unperturbed running, running against/with a traction force increases/decreases positive external work by ~ 20-70% and decreases/increases negative work by ~ 40-60%, depending on speed and traction force. The external power to maintain forward motion against a traction is contained by increasing the pushing time and step frequency. When running with an aiding force, the external power during the brake is limited by increasing braking time. Furthermore, the aerial time is increased to reduce the power required to reset the limbs each step., Conclusion: Our results show that the bouncing mechanism of running against/with a hindering/aiding traction force is equivalent to that of running on a positive/negative slope.

How does gait-specific pattern generation evolve in early infancy? The idea that neural and biomechanical mechanisms underlying mature walking and running differ to some extent and involve distinct spinal and supraspinal neural circuits is supported by various studies. Here we consider the issue of human gaits from the developmental point of view, from neonate stepping to adult mature gaits. While differentiating features of the walk and run are clearly distinct in adults, the gradual and progressive developmental bifurcation between the different gaits suggests considerable sharing of circuitry. Gaits development and their biomechanical determinants also depend on maturation of the musculoskeletal system. This review outlines the possible overlap in the neural and biomechanical control of walking and running in infancy, supporting the idea that gaits may be built starting from common, likely phylogenetically conserved elements. Bridging connections between movement mechanics and neural control of locomotion could have profound clinical implications for technological solutions to understand better locomotor development and to diagnose early motor deficits. We also consider the neuromuscular maturation time frame of gaits resulting from active practice of locomotion, underlying plasticity of development., (Copyright © 2020 Dewolf, Sylos-Labini, Cappellini, Lacquaniti and Ivanenko.)

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.

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.

Background: Ageing brings profound changes in walking gait. For example, older adults reduce the modification of pelvic and trunk kinematics with walking speed. However, the modification of the coordination between lower-limb segments with age has never been investigated across various controlled speeds., Research Question: Is the effect of speed on the intersegmental coordination different between elderly and young adults?, Methods: Nineteen senior and eight young adults walked on a treadmill at speeds ranging from 0.56 to 1.94 m s -1 . The motion of the lower-limb segments in the sagittal plane was recorded by cinematography. When the angles of the thigh, shank and foot during a stride are plotted one versus the other, they describe loops constraint on a plane. The coordination between lower-limb segments was thus evaluated by performing a principal component analysis between the thigh, shank and foot elevation angles. The effect of speed and age on the intersegmental coordination was examined using a two-level linear mixed model ANOVA., Results: In both age groups the orientation of the plane changes with speed, due to a more in-phase shank and foot motion. However, the effect of speed on the covariation plane is lessened with age., Significance: Our results demonstrate that there is an age-related specific adjustment of the intersegmental coordination to speed. In particular, older adults restrict their repertoire of angular segment motion. These differences in coordination are mainly related to different foot-shank coordination., (Copyright © 2019. Published by Elsevier B.V.)

When running, energy is lost during stance to redirect the center of mass of the body (COM) from downwards to upwards. The present study uses a collision-based approach to analyze how these energy losses change with slope and speed. Therefore, we evaluate separately the average collision angle, i.e. the angle of deviation from perpendicular relationship between the force and velocity vectors, during the absorptive and generative part of stance. Our results show that on the level, the collision angle of the absorptive phase is smaller than the collision angle of the generative phase, suggesting that the collision is generative to overcome energy losses by soft tissues. When running uphill, the collision becomes more and more generative as slope increases because the average upward vertical velocity of the COM becomes greater than on the level. When running downhill at a constant speed, the collision angle decreases during the generative phase and increases during the absorptive phase because the average downward vertical velocity of the COM becomes greater. As a result, the difference between the collision angles of the generative and absorptive phases observed on the level disappears on a shallow negative slope of ∼-6°, where the collision becomes 'pseudo-elastic' and collisional energy losses are minimized. At this 'optimal' slope, the metabolic energy consumption is minimal. On steeper negative slopes, the collision angle during the absorptive phase becomes greater than during the generative phase and the collision is absorptive. At all slopes, the collision becomes more generative when speed increases., (Copyright © 2018 Elsevier Ltd. 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.

When ascending (descending) a slope, positive (negative) work must be performed to overcome changes in gravitational potential energy at the center of body mass (COM). This modifies the pendulum-like behavior of walking. The aim of this study is to analyze how energy exchange and mechanical work done vary within a step across slopes and speeds. Ten subjects walked on an instrumented treadmill at different slopes (from -9° to 9°), and speeds (between 0.56 and 2.22 m s-1). From the ground reaction forces, we evaluated energy of the COM, recovery (i.e. the potential-kinetic energy transduction) and pendular energy savings (i.e. the theoretical reduction in work due to this recovered energy) throughout the step. When walking uphill as compared to level, pendular energy savings increase during the first part of stance (when the COM is lifted) and decreases during the second part. Conversely in downhill walking, pendular energy savings decrease during the first part of stance and increase during the second part (when the COM is lowered). In uphill and downhill walking, the main phase of external work occurs around double support. Uphill, the positive work phase is extended during the beginning of single support to raise the body. Downhill, the negative work phase starts before double support, slowing the downward velocity of the body. Changes of the pendulum-like behavior as a function of slope can be illustrated by tilting the 'classical compass model' backwards (uphill) or forwards (downhill).

When running on the level, muscles perform as much positive as negative external work. On a slope, the external positive and negative work performed are not equal. The present study analysed how the ratio between positive and negative work modifies the bouncing mechanism of running. Our goals are to: (1) identify the changes in motion of the centre of mass of the body associated with the slope of the terrain and the speed of progression, (2) study the effect of these changes on the storage and release of elastic energy during contact and (3) propose a model that predicts the change in the bouncing mechanism with slope and speed. Therefore, the ground reaction forces were measured on 10 subjects running on an instrumented treadmill at different slopes (from -9 to +9 deg) and different speeds (between 2.2 and 5.6 m s(-1)). The movements of the centre of mass of the body and its external mechanical energy were then evaluated. Our results suggest that the increase in the muscular power is contained (1) on a positive slope, by decreasing the step period and the downward movements of the body, and by increasing the duration of the push, and (2) on a negative slope, by increasing the step period and the duration of the brake, and by decreasing the upward movement of the body. Finally, the spring-mass model of running was adapted to take into account the energy added or dissipated each step on a slope., (© 2016. Published by The Company of Biologists Ltd.)