Changes in intra- and interlimb reflexes from hindlimb cutaneous afferents after staggered thoracic lateral hemisections during locomotion in cats.

When the foot dorsum contacts an obstacle during locomotion, cutaneous afferents signal central circuits to coordinate muscle activity in the four limbs. Spinal cord injury disrupts these interactions, impairing balance and interlimb coordination. We evoked cutaneous reflexes by electrically stimulating left and right superficial peroneal nerves before and after two thoracic lateral hemisections placed on opposite sides of the cord at 9- to 13-week interval in seven adult cats (4 males and 3 females). We recorded reflex responses in ten hindlimb and five forelimb muscles bilaterally. After the first (right T5-T6) and second (left T10-T11) hemisections, coordination of the fore- and hindlimbs was altered and/or became less consistent. After the second hemisection, cats required balance assistance to perform quadrupedal locomotion. Short-latency reflex responses in homonymous and crossed hindlimb muscles largely remained unaffected after staggered hemisections. However, mid- and long-latency homonymous and crossed responses in both hindlimbs occurred less frequently after staggered hemisections. In forelimb muscles, homolateral and diagonal mid- and long-latency response occurrence significantly decreased after the first and second hemisections. In all four limbs, however, when present, short-, mid- and long-latency responses maintained their phase-dependent modulation. We also observed reduced durations of short-latency inhibitory homonymous responses in left hindlimb extensors early after the first hemisection and delayed short-latency responses in the right ipsilesional hindlimb after the first hemisection. Therefore, changes in cutaneous reflex responses correlated with impaired balance/stability and interlimb coordination during locomotion after spinal cord injury. Restoring reflex transmission could be used as a biomarker to facilitate locomotor recovery. KEY POINTS: Cutaneous afferent inputs coordinate muscle activity in the four limbs during locomotion when the foot dorsum contacts an obstacle. Thoracic spinal cord injury disrupts communication between spinal locomotor centres located at cervical and lumbar levels, impairing balance and limb coordination. We investigated cutaneous reflexes during quadrupedal locomotion by electrically stimulating the superficial peroneal nerve bilaterally, before and after staggered lateral thoracic hemisections of the spinal cord in cats. We showed a loss/reduction of mid- and long-latency responses in all four limbs after staggered hemisections, which correlated with altered coordination of the fore- and hindlimbs and impaired balance. Targeting cutaneous reflex pathways projecting to the four limbs could help develop therapeutic approaches aimed at restoring transmission in ascending and descending spinal pathways.

Forelimb movements contribute to hindlimb cutaneous reflexes during locomotion in cats.

During quadrupedal locomotion, interactions between spinal and supraspinal circuits and somatosensory feedback coordinate forelimb and hindlimb movements. How this is achieved is not clear. To determine whether forelimb movements modulate hindlimb cutaneous reflexes involved in responding to an external perturbation, we stimulated the superficial peroneal nerve in six intact cats during quadrupedal locomotion and during hindlimb-only locomotion (with forelimbs standing on stationary platform) and in two cats with a low spinal transection (T12-T13) during hindlimb-only locomotion. We compared cutaneous reflexes evoked in six ipsilateral and four contralateral hindlimb muscles. Results showed similar occurrence and phase-dependent modulation of short-latency inhibitory and excitatory responses during quadrupedal and hindlimb-only locomotion in intact cats. However, the depth of modulation was reduced in the ipsilateral semitendinosus during hindlimb-only locomotion. Additionally, longer-latency responses occurred less frequently in extensor muscles bilaterally during hindlimb-only locomotion, whereas short-latency inhibitory and longer-latency excitatory responses occurred more frequently in the ipsilateral and contralateral sartorius anterior, respectively. After spinal transection, short-latency inhibitory and excitatory responses were similar to both intact conditions, whereas mid- or longer-latency excitatory responses were reduced or abolished. Our results in intact cats and the comparison with spinal-transected cats suggest that the absence of forelimb movements suppresses inputs from supraspinal structures and/or cervical cord that normally contribute to longer-latency reflex responses in hindlimb extensor muscles.NEW & NOTEWORTHY During quadrupedal locomotion, the coordination of forelimb and hindlimb movements involves central circuits and somatosensory feedback. To demonstrate how forelimb movement affects hindlimb cutaneous reflexes during locomotion, we stimulated the superficial peroneal nerve in intact cats during quadrupedal and hindlimb-only locomotion as well as in spinal-transected cats during hindlimb-only locomotion. We show that forelimb movement influences the modulation of hindlimb cutaneous reflexes, particularly the occurrence of long-latency reflex responses.

Abnormal interlimb coordination of motor developmental delay during infant crawling based on kinematic synergy analysis.

BACKGROUND: Previous studies have reported that abnormal interlimb coordination is a typical characteristic of motor developmental delay (MDD) during human movement, which can be visually manifested as abnormal motor postures. Clinically, the scale assessments are usually used to evaluate interlimb coordination, but they rely heavily on the subjective judgements of therapists and lack quantitative analysis. In addition, although abnormal interlimb coordination of MDD have been studied, it is still unclear how this abnormality is manifested in physiology-related kinematic features. OBJECTIVES: This study aimed to evaluate how abnormal interlimb coordination of MDD during infant crawling was manifested in the stability of joints and limbs, activation levels of synergies and intrasubject consistency from the kinematic synergies of tangential velocities of joints perspective. METHODS: Tangential velocities of bilateral shoulder, elbow, wrist, hip, knee and ankle over time were computed from recorded three-dimensional joint trajectories in 40 infants with MDD [16 infants at risk of developmental delay, 11 infants at high risk of developmental delay, 13 infants with confirmed developmental delay (CDD group)] and 20 typically developing infants during hands-and-knees crawling. Kinematic synergies and corresponding activation coefficients were derived from those joint velocities using the non-negative matrix factorization algorithm. The variability accounted for yielded by those synergies and activation coefficients, and the synergy weightings in those synergies were used to measure the stability of joints and limbs. To quantify the activation levels of those synergies, the full width at half maximum and center of activity of activation coefficients were calculated. In addition, the intrasubject consistency was measured by the cosine similarity of those synergies and activation coefficients. RESULTS: Interlimb coordination patterns during infant crawling were the combinations of four types of single-limb movements, which represent the dominance of each of the four limbs. MDD mainly reduced the stability of joints and limbs, and induced the abnormal activation levels of those synergies. Meanwhile, MDD generally reduced the intrasubject consistency, especially in CDD group. CONCLUSIONS: These features have the potential for quantitatively evaluating abnormal interlimb coordination in assisting the clinical diagnosis and motor rehabilitation of MDD.

Left-Right Locomotor Coordination in Human Neonates.

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.

A secondary motor area contributing to interlimb coordination during visually guided locomotion in the cat.

We investigated the contribution of cytoarchitectonic cortical area 4δc, in the caudal bank of the cruciate sulcus of the cat, to the control of visually guided locomotion. To do so, we recorded the activity of 114 neurons in 4δc while cats walked on a treadmill and stepped over an obstacle that advanced toward them. A total of 84/114 (74%) cells were task-related and 68/84 (81%) of these cells showed significant modulation of their discharge frequency when the contralateral limbs were the first to step over the obstacle. These latter cells included a substantial proportion (27/68 40%) that discharged between the passage of the contralateral forelimb and the contralateral hindlimb over the obstacle, suggesting a contribution of this area to interlimb coordination. We further compared the discharge in area 4δc with the activity patterns of cells in the rostral division of the same cytoarchitectonic area (4δr), which has been suggested to be a separate functional region. Despite some differences in the patterns of activity in the 2 subdivisions, we suggest that activity in each is compatible with a contribution to interlimb coordination and that they should be considered as a single functional area that contributes to both forelimb-forelimb and forelimb-hindlimb coordination.

Modulation of the gait pattern during split-belt locomotion after lateral spinal cord hemisection in adult cats.

Most previous studies investigated the recovery of locomotion in animals and people with incomplete spinal cord injury (SCI) during relatively simple tasks (e.g., walking in a straight line on a horizontal surface or a treadmill). We know less about the recovery of locomotion after incomplete SCI in left-right asymmetric conditions, such as turning or stepping along circular trajectories. To investigate this, we collected kinematic and electromyography data during split-belt locomotion at different left-right speed differences before and after a right thoracic lateral spinal cord hemisection in nine adult cats. After hemisection, although cats still performed split-belt locomotion, we observed several changes in the gait pattern compared with the intact state at early (1-2 wk) and late (7-8 wk) time points. Cats with larger lesions showed new coordination patterns between the fore- and hindlimbs, with the forelimbs taking more steps. Despite this change in fore-hind coordination, cats maintained consistent phasing between the fore- and hindlimbs. Adjustments in cycle and phase (stance and swing) durations between the slow and fast sides allowed animals to maintain 1:1 left-right coordination. Periods of triple support involving the right (ipsilesional) hindlimb decreased in favor of quad support and triple support involving the other limbs. Step and stride lengths decreased with concurrent changes in the right fore- and hindlimbs, possibly to avoid interference. The above adjustments in the gait pattern allowed cats to retain the ability to locomote in asymmetric conditions after incomplete SCI. We discuss potential plastic neuromechanical mechanisms involved in locomotor recovery in these conditions.NEW & NOTEWORTHY Everyday locomotion often involves left-right asymmetries, when turning, walking along circular paths, stepping on uneven terrains, etc. To show how incomplete spinal cord injury affects locomotor control in asymmetric conditions, we collected data before and after a thoracic lateral spinal hemisection on a split-belt treadmill with one side stepping faster than the other. We show that adjustments in kinematics and muscle activity allowed cats to retain the ability to perform asymmetric locomotion after hemisection.

Simple decentralized control mechanism that enables limb adjustment for adaptive quadruped running.

Quadrupeds exhibit versatile and adaptive running by exploiting the flying phase during the stride cycle. Various interlimb coordination mechanisms focusing on mechanical loads during the stance phase have been proposed to understand the underlying control mechanism, and various gait patterns have been reproduced. However, the essential control mechanism required to achieve both steady running patterns and non-steady behaviours, such as jumping and landing, remains unclear. Therefore, we focus on the vertical motions of the body parts and propose a new decentralized interlimb coordination mechanism. The simulation results demonstrate that the robot can generate efficient and various running patterns in response to the morphology of the body. Furthermore, the proposed model allows the robot to smoothly change its behaviour between steady running and non-steady landing depending on the situation. These results suggest that the steady and non-steady behaviours in quadruped adaptive running may share a common simple control mechanism based on the mechanical loads and vertical velocities of the body parts.

A Fully Wireless Wearable Motion Tracking System with 3D Human Model for Gait Analysis.

This paper presents a wearable motion tracking system with recording and playback features. This system has been designed for gait analysis and interlimb coordination studies. It can be implemented to help reduce fall risk and to retrain gait in a rehabilitation setting. Our system consists of ten custom wearable straps, a receiver, and a central computer. Comparing with similar existing solutions, the proposed system is affordable and convenient, which can be used in both indoor and outdoor settings. In the experiment, the system calculates five gait parameters and has the potential to identify deviant gait patterns. The system can track upper body parameters such as arm swing, which has potential in the study of pathological gaits and the coordination of the limbs.

Changing coupling between the arms and legs with slow walking speeds alters regulation of somatosensory feedback.

Arm swing movement is coordinated with movement of the legs during walking, where the frequency of coordination depends on walking speed. At typical speeds, arm and leg movements, respectively, are frequency locked in a 1:1 ratio but at slow speeds this changes to a 2:1 ratio. It is unknown if the changes in interlimb ratio that accompany slow walking speeds alters regulation of somatosensory feedback. To probe the neural interactions between the arms and legs, somatosensory linkages in the form of interlimb cutaneous reflexes were examined. It was hypothesized that different interlimb frequencies and walking speeds would result in changes in the modulation of cutaneous reflexes between the arms and legs. To test this hypothesis, participants walked in four combinations of walking speed (typical, slow) and interlimb coordination (1:1, and 2:1), while cutaneous reflexes and background muscle activity were evaluated with stimulation applied to the superficial peroneal nerve at the ankle and superficial radial nerve at the wrist. Results show main effects of interlimb coordination and walking speed on cutaneous reflex modulation, effects are largest in the swing phase, and a directional coupling was observed, where changes in the frequency of arm movements had a greater effect on muscle activity in the legs compared to the reverse. Task-dependent modulation was also revealed from stimulation at local and remote sources. Understanding the underlying neural mechanisms for the organization of rhythmic arm movement, and its coordination with the legs in healthy participants, can give insight into pathological walking, and will facilitate the development of effective strategies for the rehabilitation of walking.

Intralimb and Interlimb Cutaneous Reflexes during Locomotion in the Intact Cat.

When the foot contacts an obstacle during locomotion, cutaneous inputs activate spinal circuits to ensure dynamic balance and forward progression. In quadrupeds, this requires coordinated reflex responses between the four limbs. Here, we investigated the patterns and phasic modulation of cutaneous reflexes in forelimb and hindlimb muscles evoked by inputs from all four limbs. Five female cats were implanted to record muscle activity and to stimulate the superficial peroneal and superficial radial nerves during locomotion. Stimulating these nerves evoked short-, mid-, and longer-latency excitatory and/or inhibitory responses in all four limbs that were phase-dependent. The largest responses were generally observed during the peak activity of the muscle. Cutaneous reflexes during mid-swing were consistent with flexion of the homonymous limb and accompanied by modification of the stance phases of the other three limbs, by coactivating flexors and extensors and/or by delaying push-off. Cutaneous reflexes during mid-stance were consistent with stabilizing the homonymous limb by delaying and then facilitating its push-off and modifying the support phases of the homolateral and diagonal limbs, characterized by coactivating flexors and extensors, reinforcing extensor activity and/or delaying push-off. The shortest latencies of homolateral and diagonal responses were consistent with fast-conducting disynaptic or trisynaptic pathways. Descending homolateral and diagonal pathways from the forelimbs to the hindlimbs had a higher probability of eliciting responses compared with ascending pathways from the hindlimbs to the forelimbs. Thus, in quadrupeds, intralimb and interlimb reflexes activated by cutaneous inputs ensure dynamic coordination of the four limbs, producing a whole-body response.SIGNIFICANCE STATEMENT The skin contains receptors that, when activated, send inputs to spinal circuits, signaling a perturbation. Rapid responses, or reflexes, in muscles of the contacted limb and opposite homologous limb help maintain balance and forward progression. Here, we investigated reflexes during quadrupedal locomotion in the cat by electrically stimulating cutaneous nerves in each of the four limbs. Functionally, responses appear to modify the trajectory or stabilize the movement of the stimulated limb while modifying the support phase of the other limbs. Reflexes between limbs are mediated by fast-conducting pathways that involve excitatory and inhibitory circuits controlling each limb. The comparatively stronger descending pathways from cervical to lumbar circuits controlling the forelimbs and hindlimbs, respectively, could serve a protective function.

Error-related modulations of the sensorimotor post-movement and foreperiod beta-band activities arise from distinct neural substrates and do not reflect efferent signal processing.

While beta activity has been extensively studied in relation to voluntary movement, its role in sensorimotor adaptation remains largely uncertain. Recently, it has been shown that the post-movement beta rebound as well as beta activity during movement-preparation are modulated by movement errors. However, there are critical functional differences between pre- and post-movement beta activities. Here, we addressed two related open questions. Do the pre- and post-movement error-related modulations arise from distinct neural substrates? Do these modulations relate to efferent signals shaping muscle-activation patterns or do they reflect integration of sensory information, intervening upstream of the motor output? For this purpose, first we exploited independent component analysis (ICA) which revealed a double dissociation suggesting that distinct neural substrates are recruited in error-related beta-power modulations observed before and after movement. Second, we compared error-related beta oscillation responses observed in two bimanual reaching tasks involving similar movements but different interlimb coordination, and in which the same mechanical perturbations induced different behavioral adaptive responses. While the task difference was not reflected in the post-movement beta rebound, the pre-movement beta activity was differently modulated according to the interlimb coordination. Critically, we show an uncoupling between the behavioral and the electrophysiological responses during the movement preparation phase, which demonstrates that the error-related modulation of the foreperiod beta activity does not reflect changes in the motor output from primary motor cortex. It seems instead to relate to higher level processing of sensory afferents, essential for sensorimotor adaptation.

Age-related deficits in bilateral motor synergies and force coordination.

BACKGROUND: Ageing may cause impairments in executing bilateral movement control. This study investigated age-related changes in interlimb force coordination across multiple trials by quantifying bilateral motor synergies based on the uncontrolled manifold hypothesis. Participants completed the trials with and without visual feedback. METHODS: Twenty healthy individuals (10 older adults and 10 young adults) performed 12 isometric force control trials for the two vision conditions at 5% of maximal voluntary contraction. All dependent variables were analyzed in two-way mixed model (Group × Vision Condition; 2 × 2) ANOVAs with repeated measures on the last factor. RESULTS: The analyses revealed that older adults had greater mean force produced by two hands in both vision conditions (i.e., yes and no visual feedback). Across both vision conditions, the older adult group showed greater asymmetrical force variability (i.e., standard deviation of non-dominant hand > standard deviation of dominant hand) and revealed more positive correlation coefficients between forces produced by two hands as compared with the young adult group. Finally, an index of bilateral motor synergies was significantly greater in young adults than older adults when visual feedback was available. CONCLUSION: The current findings indicate that deficits in interlimb force coordination across multiple trials appeared in older adults.

Effect of cervicolumbar coupling on spinal reflexes during cycling after incomplete spinal cord injury.

Spinal networks in the cervical and lumbar cord are actively coupled during locomotion to coordinate arm and leg activity. The goals of this project were to investigate the intersegmental cervicolumbar connectivity during cycling after incomplete spinal cord injury (iSCI) and to assess the effect of rehabilitation training on improving reflex modulation mediated by cervicolumbar pathways. Two studies were conducted. In the first, 22 neurologically intact (NI) people and 10 people with chronic iSCI were recruited. The change in H-reflex amplitude in flexor carpi radialis (FCR) during leg cycling and H-reflex amplitude in soleus (SOL) during arm cycling were investigated. In the second study, two groups of participants with chronic iSCI underwent 12 wk of cycling training: one performed combined arm and leg cycling (A&L) and the other legs only cycling (Leg). The effect of training paradigm on the amplitude of the SOL H-reflex was assessed. Significant reduction in the amplitude of both FCR and SOL H-reflexes during dynamic cycling of the opposite limbs was found in NI participants but not in participants with iSCI. Nonetheless, there was a significant reduction in the SOL H-reflex during dynamic arm cycling in iSCI participants after training. Substantial improvements in SOL H-reflex properties were found in the A&L group after training. The results demonstrate that cervicolumbar modulation during rhythmic movements is disrupted in people with chronic iSCI; however, this modulation is restored after cycling training. Furthermore, involvement of the arms simultaneously with the legs during training may better regulate the leg spinal reflexes. NEW & NOTEWORTHY This work systematically demonstrates the disruptive effect of incomplete spinal cord injury on cervicolumbar coupling during rhythmic locomotor movements. It also shows that the impaired cervicolumbar coupling could be significantly restored after cycling training. Actively engaging the arms in rehabilitation paradigms for the improvement of walking substantially regulates the excitability of the lumbar spinal networks. The resulting regulation may be better than that obtained by interventions that focus on training of the legs only.

Effect of salient points in movements on the constraints in bimanual coordination.

The relative-salience hypothesis has been proposed as a possible explanation for the stability of bimanual coordination. This explanation proceeds from a psychological viewpoint and is based on the following tenets: (1) cyclic joint motions involving two movements are conceived of as a unified event, (2) if a "single" point in each movement is seen as the most salient, the salient points of the two movements prefer to go together, and (3) in other cases, a unified event will be constrained by movement direction. In this investigation, we examined whether the relative-salience hypothesis could predict the type of constraint (i.e., action coupling vs movement direction) for various bimanual coordination movements. Participants performed six different joint movements in synchrony with metronome beats. Both index finger flexion/extension and forearm pronation/supination had a "single" salient point (JMsingleSP), the others had "two" salient points (JMtwoSP). Then, we applied the relative-salience hypothesis to four bimanual coordinations. The coupling of simultaneous forearm pronation was more stable than alternate pronation. Similarly, the coupling of finger flexion and forearm pronation was more stable than that of finger flexion and forearm supination. For the coordination of radial flexion/ulnar flexion and index finger flexion/extension as well as forearm pronation/supination and radial flexion/ulnar flexion, symmetric movements were more stable than asymmetric movements. The results indicated that the stability of bimanual coordination was predominantly constrained by coupling of salient points when using two JMsingleSP and it was predominantly constrained by movement direction when coordinating JMsingleSP and JMtwoSP. Thus, the relative-salience hypothesis was supported.

Delayed muscle onset soreness in the gastrocnemius muscle attenuates the spinal contribution to interlimb communication.

PURPOSE: Delayed onset muscle soreness (DOMS) has been shown to induce changes in muscle activity during walking. The aim of this study was to elucidate whether DOMS also affects interlimb communication during walking by investigating its effect on short-latency crossed responses (SLCRs). METHODS: SLCRs were elicited in two recording sessions by electrically stimulating the tibial nerve of the ipsilateral leg, and quantified in the contralateral gastrocnemius muscle. The second recording session occurred 24-36 h after the participants (n = 11) performed eccentric exercises with the ipsilateral calf. RESULTS: DOMS caused a decreased magnitude of the spinally mediated component of the SLCR in the contralateral gastrocnemius medialis. CONCLUSIONS: The results of the current study provide insight on the relationship between pain and motor control. Muscle pain affects the spinal pathway mediating interlimb communication, which might result in a reduced ability to maintain dynamical stability during walking.

Lack of adaptation during prolonged split-belt locomotion in the intact and spinal cat.

KEY POINTS: During split-belt locomotion in humans where one leg steps faster than the other, the symmetry of step lengths and double support periods of the slow and fast legs is gradually restored. When returning to tied-belt locomotion, there is an after-effect, with a reversal in the asymmetry observed in the early split-belt period, indicating that the new pattern was stored within the central nervous system. In this study, we investigated if intact and spinal-transected cats show a similar pattern of adaptation to split-belt locomotion by measuring kinematic variables and electromyography before, during and after 10 min of split-belt locomotion. The results show that cats do not adapt to prolonged split-belt locomotion. Our results suggest an important physiological difference in how cats and humans respond to prolonged asymmetric locomotion. ABSTRACT: In humans, gait adapts to prolonged walking on a split-belt treadmill, where one leg steps faster than the other, by gradually restoring the symmetry of interlimb kinematic variables, such as double support periods and step lengths, and by reducing muscle activity (EMG, electromyography). The adaptation is also characterized by reversing the asymmetry of interlimb variables observed during the early split-belt period when returning to tied-belt locomotion, termed an after-effect. To determine if cats adapt to prolonged split-belt locomotion and to assess if spinal locomotor circuits participate in the adaptation, we measured interlimb variables and EMG in intact and spinal-transected cats before, during and after 10 min of split-belt locomotion. In spinal cats, only the hindlimbs performed stepping with the forelimbs stationary. In intact and spinal cats, step lengths and double support periods were, on average, symmetric, during tied-belt locomotion. They became asymmetric during split-belt locomotion and remained asymmetric throughout the split-belt period. Upon returning to tied-belt locomotion, symmetry was immediately restored. In intact cats, the mean EMG amplitude of hindlimb extensors increased during split-belt locomotion and remained increased throughout the split-belt period, whereas in spinal cats, EMG amplitude did not change. Therefore, the results indicate that the locomotor pattern of cats does not adapt to prolonged split-belt locomotion, suggesting an important physiological difference in the control of locomotion between cats and humans. We propose that restoring left-right symmetry is not required to maintain balance during prolonged asymmetric locomotion in the cat, a quadruped, as opposed to human bipedal locomotion.

Modulation of corticospinal input to the legs by arm and leg cycling in people with incomplete spinal cord injury.

The spinal cervico-lumbar interaction during rhythmic movements in humans has recently been studied; however, the role of arm movements in modulating the corticospinal drive to the legs is not well understood. The goals of this study were to investigate the effect of active rhythmic arm movements on the corticospinal drive to the legs (study 1) and assess the effect of simultaneous arm and leg training on the corticospinal pathway after incomplete spinal cord injury (iSCI) (study 2). In study 1, neurologically intact (NI) participants or participants with iSCI performed combinations of stationary and rhythmic cycling of the arms and legs while motor evoked potentials (MEPs) were recorded from the vastus lateralis (VL) muscle. In the NI group, arm cycling alone could facilitate the VL MEP amplitude, suggesting that dynamic arm movements strongly modulate the corticospinal pathway to the legs. No significant difference in VL MEP between conditions was found in participants with iSCI. In study 2, participants with iSCI underwent 12 wk of electrical stimulation-assisted cycling training: one group performed simultaneous arm and leg (A&L) cycling and the other legs-only cycling. MEPs in the tibialis anterior (TA) muscle were compared before and after training. After training, only the A&L group had a significantly larger TA MEP, suggesting increased excitability in the corticospinal pathway. The findings demonstrate the importance of arm movements in modulating the corticospinal drive to the legs and suggest that active engagement of the arms in lower limb rehabilitation may produce better neural regulation and restoration of function.NEW & NOTEWORTHY This study aimed to demonstrate the importance of arm movements in modulating the corticospinal drive to the legs. It provides direct evidence in humans that active movement of the arms could facilitate corticospinal transmission to the legs and, for the first time, shows that facilitation is absent after spinal cord injury. Active engagement of the arms in lower limb rehabilitation increased the excitability of the corticospinal pathway and may produce more effective improvement in leg function.

The neural control of interlimb coordination during mammalian locomotion.

Neuronal networks within the spinal cord directly control rhythmic movements of the arms/forelimbs and legs/hindlimbs during locomotion in mammals. For an effective locomotion, these networks must be flexibly coordinated to allow for various gait patterns and independent use of the arms/forelimbs. This coordination can be accomplished by mechanisms intrinsic to the spinal cord, somatosensory feedback from the limbs, and various supraspinal pathways. Incomplete spinal cord injury disrupts some of the pathways and structures involved in interlimb coordination, often leading to a disruption in the coordination between the arms/forelimbs and legs/hindlimbs in animal models and in humans. However, experimental spinal lesions in animal models to uncover the mechanisms coordinating the limbs have limitations due to compensatory mechanisms and strategies, redundant systems of control, and plasticity within remaining circuits. The purpose of this review is to provide a general overview and critical discussion of experimental studies that have investigated the neural mechanisms involved in coordinating the arms/forelimbs and legs/hindlimbs during mammalian locomotion.

Contralateral conditioning to the soleus H-reflex as a function of age and physical activity.

Coordination between ipsilateral and contralateral muscles of the lower limbs has a critical role in movement control. However, the roles that aging and physical activity have in maintaining bilateral coordination are understudied. The aim of this study was to examine the roles of physical activity and age on pathways between the soleus and tibialis anterior muscles. Fourteen young and 14 older subjects (7 active, 7 sedentary) participated. A Hoffmann (H)-reflex was elicited in the ipsilateral soleus following a conditioning stimulus to the contralateral common peroneal nerve at differing intervals (25-300 ms). Significant H-reflex facilitation from the control value was observed for the sedentary group at the 50-ms (28.7 %), 75-ms (24.5 %), and 150-ms (34.0 %) intervals when compared with the physically active group. There were also significant differences between the young and older groups. Results demonstrated differences in soleus H-reflex excitability as a result of contralateral conditioning and highlighted the influence of age and physical activity in maintaining these neural pathways.

Soleus Hoffmann reflex amplitudes are specifically modulated by cutaneous inputs from the arms and opposite leg during walking but not standing.

Electrical stimulation of cutaneous nerves innervating heteronymous limbs (the arms or contralateral leg) modifies the excitability of soleus Hoffmann (H-) reflexes. The differences in the sensitivities of the H-reflex pathway to cutaneous afferents from different limbs and their modulation during the performance of motor tasks (i.e., standing and walking) are not fully understood. In the present study, we investigated changes in soleus H-reflex amplitudes induced by electrical stimulation of peripheral nerves. Selected targets for conditioning stimulation included the superficial peroneal nerve, which innervates the foot dorsum in the contralateral ankle (cSP), and the superficial radial nerve, which innervates the dorsum of the hand in the ipsilateral (iSR) or contralateral wrist (cSR). Stimulation and subsequent reflex assessment took place during the standing and early-stance phase of treadmill walking in ten healthy subjects. Cutaneous stimulation produced long-latency inhibition (conditioning-test interval of ~100 ms) of the H-reflex during the early-stance phase of walking, and the inhibition was stronger following cSP stimulation compared with iSR or cSR stimulation. In contrast, although similar conditioning stimulation significantly facilitated the H-reflex during standing, this effect remained constant irrespective of the different conditioning sites. These findings suggest that cutaneous inputs from the arms and contralateral leg had reversible effects on the H-reflex amplitudes, including inhibitions with different sensitivities during the early-stance phase of walking and facilitation during standing. Furthermore, the differential sensitivities of the H-reflex modulations were expressed only during walking when the locations of the afferent inputs were functionally relevant.