Search

BACKGROUND: Increased knee osteoarthritis risk in patients with unilateral lower extremity limb loss is attributed to increased intact limb loading. Modulating powered ankle prosthesis push-off power may be an effective way to modulate intact limb loading. We examined how changes in the parameter settings of a commercial prosthetic ankle affect power delivery during push-off and the resulting collisional work experienced by the intact limb in persons with unilateral lower extremity limb loss. METHODS: Five subjects with unilateral transtibial amputation were fitted with a commercially available powered ankle prosthesis (Ottobock Empower). Subjects walked on a treadmill in seven conditions, where ankle power delivery settings were adjusted using methods accessible to clinicians. Kinetics and kinematics data were collected. RESULTS: Standard adjustment of parameter settings within the prosthetic foot did not alter timing of peak prosthesis power or intact limb collisional work, but did have a significant effect on the magnitude of positive prosthesis ankle work. Increased prosthesis work did not decrease intact limb collisional work as predicted. CONCLUSIONS: Altering the parameter settings on a commercial powered ankle prosthesis affected the magnitude, but not the timing, of power delivered. Increased prosthesis push-off power did not decrease intact limb loading.

Despite the prevalence of rat models to study human disease and injury, existing methods for quantifying behavior through skeletal movements are problematic owing to skin movement inaccuracies associated with optical video analysis, or require invasive implanted markers or time-consuming manual rotoscoping for X-ray video approaches. We examined the use of a machine learning tool, DeepLabCut, to perform automated, markerless tracking in bi-planar X-ray videos of locomoting rats. Models were trained on 590 pairs of video frames to identify 19 unique skeletal landmarks of the pelvic limb. Accuracy, precision and time savings were assessed. Machine-identified landmarks deviated from manually labeled counterparts by 2.4±0.2 mm (n=1710 landmarks). DeepLabCut decreased analysis time by over three orders of magnitude (1627×) compared with manual labeling. Distribution of these models may enable the processing of a large volume of accurate X-ray kinematics locomotion data in a fraction of the time without requiring surgically implanted markers.

Humans can inherently adapt their gait pattern in a way that minimizes the metabolic cost of transport, or walking economy, within a few steps, which is faster than any known direct physiological sensor of metabolic energy. Instead, walking economy may be indirectly sensed through mechanoreceptors that correlate with the metabolic cost per step to make such gait adaptations. We tested whether velocity feedback from tibialis anterior (TA) muscle fascicles during the collisional phase of walking could potentially act to indirectly sense walking economy. As participants walked within a range of steady-state speeds and step frequencies, we observed that TA fascicles lengthen on almost every step. Moreover, the average peak fascicle velocity experienced during lengthening reflected the metabolic cost of transport of the given walking condition. We observed that the peak TA muscle activation occurred earlier than could be explained by a short latency reflex response. The activation of the TA muscle just prior to heel strike may serve as a prediction of the magnitude of the ground collision and its associated energy loss. In this scenario, any unexpected length change experienced by the TA fascicle would serve as an error signal to the nervous system and provide additional information about energy lost per step. Our work helps provide a biomechanical framework to understand the possible neural mechanisms underlying the rapid optimization of walking economy.

Background Manual tuning of robotic lower limb prostheses can be time consuming for both the patient and the clinician and requires in-person visits to a clinic. An automated process for the tuning parameters of a robotic lower limb prosthesis could result in a substantial savings in healthcare resources. A critical challenge to an automated parameter tuning algorithm is the quantification of a person’s gait quality. There is not good agreement in the literature of an objective outcome measure that can rapidly assess gait quality in lower limb amputees. As a first step, we investigated the ability of four common gait quality metrics to detect differences in gait quality: Prosthetic Observational Gait Score (POGS), Gait Deviation Index (GDI), Lateral Sway, and Impulse Asymmetry. Methods We systematically applied four unilateral lower limb joint constraint conditions (baseline/no constraint, ankle constraint, knee constraint, and knee + ankle constraint) to nine able-bodied participants walking at three different speeds (0.7, 0.85 and 1.0 m/s). We calculated and compared the resulting GDI, POGS, Lateral Sway and Impulse Asymmetry scores across all conditions. We performed a 2-way ANOVA statistical analysis to compare sensitivity of the metrics to the various conditions with significance defined by an alpha-level = 0.05. Results The Lateral Sway metric distinguished three joint constraint conditions and two of the speed conditions. Both GDI and POGS were able to distinguish four out of six possible constraint-speed conditions, while Impulse Asymmetry was only able to detect differences between three of the six constraint-speed conditions. Conclusions No single gait quality metric could distinguish every condition. Accordingly, a single metric of gait quality may be inadequate for tuning a prosthesis and therefore multiple metrics and sensors may provide the best results for tuning a prosthesis to the most natural gait pattern for an individual. Compared to the more complex gait measures, Lateral Sway performed well as a simple metric that might easily be operationalized into a real-time parameter tuning controller.

STUDY DESIGN A mechanical testing protocol was used to compare the material properties of commercially available foams with that of a newly designed granular jamming orthosis prototypes. BACKGROUND Foot orthoses have an inherent limitation of predetermined mechanical material properties coupled with a fixed orthotic interface shape that cannot be readily changed. OBJECTIVES To develop and test a novel orthotic insole design concept that incorporates principles of granular jamming. METHODS Granular media were used in combination with vacuum pressure to create a variable stiffness granular foot orthosis. Four types of granular media (rice, poppy seeds, micropolystyrene, and polystyrene beads) were tested in different prototype configurations varying in volume fill and particulate size. Stress-strain curves were obtained from uniaxial compression tests to characterize granular foot orthosis prototypes in comparison with commercial orthotic foams. RESULTS Increasing vacuum pressure increased prototype stiffness for most configurations. A single granular jamming orthosis could exhibit energy absorption values that spanned the entire commercial foam performance range, and in some cases extended far beyond the upper values of the tested foams. CONCLUSION The results suggest that granular jamming principles can provide clinicians the capability for rapid selection of mechanical properties over a wide range of orthosis stiffnesses. Importantly, patients could don the orthosis because the clinician makes real-time assessments and adjustments in the clinic.

Category: Ankle Arthritis Introduction/Purpose: Total ankle replacement (TAR) is common tool used by the foot and ankle specialist to treat end stage ankle arthritis. Current data about ankle motion following TAR is derived from gait analysis utilizing external markers. Utilizing Xray Reconstruction of Moving Morphology (XROMM), which combines 3-D mapping technology with biplanar fluoroscopy in vivo to visualize true skeletal motion, we can evaluate true motion of TAR implants. Current TAR replacement systems are either mobile bearing or fixed bearing. We hypothesized that subjects implanted with a fixed bearing prosthesis would exhibit less tibiotalar rotation and translation than subjects implanted with a mobile bearing prosthesis. Methods: Six subjects with total ankle replacement at least one-year post implantation gave informed consent before participating (IRB #H16496). Three subjects with a mobile bearing prosthesis with an average age 63.3+-11.1 yrs were compared to three matched subjects with a fixed bearing prosthesis with an average age of 64.7+-1.5 yrs. Utilizing 3D slicer software, lower body CT scans for each subject were evaluated to create 3D models of the foot and ankle bones and implant components. All subjects walked for several trials at a self-selected pace along a walkway while their foot and ankle motions were captured by a high-speed biplanar fluoroscopic x-ray motion analysis (XMA) system. The 3D models were combined with the x-ray images within a 3D animation platform and rotoscoped to resolve accurate kinematic motions at the tibiotalar joint during stance phase of gait. We examined for differences between the two groups using a two-sample t-test (pResults: Subjects with a mobile-bearing prosthesis demonstrated mean ROM’s of 7.4+-1.1°, 5.3+-2.3° and 7.1+-4.3° for dorsiflexion/plantarflexion, inversion/eversion, and internal/external rotation, respectively. Subjects with a fixed bearing ankle prosthesis did not exhibit significantly different mean ROM’s for dorsiflexion/plantarflexion (9.1+-4.0°, p=0.35), inversion/eversion (4.4+-2.1°, p=0.42), and internal/external rotation (9.0+-3.4°, p=0.35), respectively. Subjects with a fixed bearing prosthesis displayed significantly more translation along the anteroposterior (3.6+-1.2mm, pConclusion: Our preliminary results indicate that mobile and fixed bearing prosthesis provides similar angular motion at the tibiotalar joint, however, the fixed bearing prosthesis exhibits greater translational motion during walking. Further, there is the same amount of internal and external translation with both component designs. The implications of this work on success or failure of current implant designs is beyond the scope of this study but this work will provide the basis for future studies to help determine optimal future total ankle replacement designs.

Locomotor adaptation is commonly studied using split-belt treadmill walking, in which each foot is placed on a belt moving at a different speed. As subjects adapt to split-belt walking, they reduce metabolic power, but the biomechanical mechanism behind this improved efficiency is unknown. Analyzing mechanical work performed by the legs and joints during split-belt adaptation could reveal this mechanism. Because ankle work in the step-to-step transition is more efficient than hip work, we hypothesized that control subjects would reduce hip work on the fast belt and increase ankle work during the step-to-step transition as they adapted. We further hypothesized that subjects with unilateral, trans-tibial amputation would instead increase propulsive work from their intact leg on the slow belt. Control subjects reduced hip work and shifted more ankle work to the step-to-step transition, supporting our hypothesis. Contrary to our second hypothesis, intact leg work, ankle work and hip work in amputees were unchanged during adaptation. Furthermore, all subjects increased collisional energy loss on the fast belt, but did not increase propulsive work. This was possible because subjects moved further backward during fast leg single support in late adaptation than in early adaptation, compensating by reducing backward movement in slow leg single support. In summary, subjects used two strategies to improve mechanical efficiency in split-belt walking adaptation: a CoM displacement strategy that allows for less forward propulsion on the fast belt; and, an ankle timing strategy that allows efficient ankle work in the step-to-step transition to increase while reducing inefficient hip work.

Summary Objective This study examined the effects of reduced and elevated weight bearing on post-traumatic osteoarthritis (PTOA) development, locomotor joint kinematics, and degree of voluntary activity in rats following medial meniscal transection (MMT). Design Twenty-one adult rats were subjected to MMT surgery of the left hindlimb and then assigned to one of three groups: (1) regular (i.e., no intervention), (2) hindlimb immobilization, or (3) treadmill running. Sham surgery was performed in four additional rats. Voluntary wheel run time/distance was measured, and 3D hindlimb kinematics were quantified during treadmill locomotion using biplanar radiography. Rats were euthanized 8 weeks after MMT or sham surgery, and the microstructure of the tibial cartilage and subchondral bone was quantified using contrast enhanced micro-CT. Results All three MMT groups showed signs of PTOA (full-thickness lesions and/or increased cartilage volume) compared to the sham group, however the regular and treadmill-running groups had greater osteophyte formation than the immobilization group. For the immobilization group, increased volume was only observed in the anterior region of the cartilage. The treadmill-running group demonstrated a greater knee varus angle at mid-stance than the sham group, while the immobilization group demonstrated greater reduction in voluntary running than all the other groups at 2 weeks post-surgery. Conclusions Elevated weight-bearing via treadmill running at a slow/moderate speed did not accelerate PTOA in MMT rats when compared to regular weight-bearing. Reduced weight-bearing via immobilization may attenuate overall PTOA but still resulted in regional cartilage degeneration. Overall, there were minimal differences in hindlimb kinematics and voluntary running between MMT and sham rats.

For the controller of wearable lower-limb assistive devices, quantitative understanding of human locomotion serves as the basis for human motion intent recognition and joint-level motion control. Traditionally, the required gait data are obtained in gait research laboratories, utilizing marker-based optical motion capture systems. Despite the high accuracy of measurement, marker-based systems are largely limited to laboratory environments, making it nearly impossible to collect the desired gait data in real-world daily-living scenarios. To address this problem, the authors propose a novel exoskeleton-based gait data collection system, which provides the capability of conducting independent measurement of lower limb movement without the need for stationary instrumentation. The basis of the system is a lightweight exoskeleton with articulated knee and ankle joints. To minimize the interference to a wearer&rsquo, s natural lower-limb movement, a unique two-degrees-of-freedom joint design is incorporated, integrating a primary degree of freedom for joint motion measurement with a passive degree of freedom to allow natural joint movement and improve the comfort of use. In addition to the joint-embedded goniometers, the exoskeleton also features multiple positions for the mounting of inertia measurement units (IMUs) as well as foot-plate-embedded force sensing resistors to measure the foot plantar pressure. All sensor signals are routed to a microcontroller for data logging and storage. To validate the exoskeleton-provided joint angle measurement, a comparison study on three healthy participants was conducted, which involves locomotion experiments in various modes, including overground walking, treadmill walking, and sit-to-stand and stand-to-sit transitions. Joint angle trajectories measured with an eight-camera motion capture system served as the benchmark for comparison. Experimental results indicate that the exoskeleton-measured joint angle trajectories closely match those obtained through the optical motion capture system in all modes of locomotion (correlation coefficients of 0.97 and 0.96 for knee and ankle measurements, respectively), clearly demonstrating the accuracy and reliability of the proposed gait measurement system.

Propulsive force production (indicative of intrinsic force-length-velocity characteristics of the plantar flexor muscles) has been shown to be a major determinant of the human walk-to-run transition. The purpose of this work was to determine the gait transition speed of persons with unilateral, transtibial amputation donning a passive-elastic prosthesis and assess whether a mechanical limit of their intact side plantar flexor muscles is a major determinant of their walk-to-run transition. We determined each individual’s gait transition speed (GTS) via an incremental protocol and assessed kinetics and kinematics during walking at speeds 50, 60, 70, 80, 90, 100, 120, and 130% of that gait transition speed (100%:GTS). Unilateral, transtibial amputees transitioned between gaits at significantly slower absolute speeds than matched able-bodied controls (1.73±0.13 and 2.09±0.05m/s respectively, p 120%: 0.23±0.05BW, p

Clinically, measuring gait kinematics and ground reaction force (GRF) is useful to determine the effectiveness of treatment. However, it is inconvenient and expensive to maintain a laboratory-grade gait analysis system in most clinics. The purpose of this study was to validate a Wii Balance Board, Kinovea motion-tracking software, and a video camera as a portable, low-cost system, and overground gait analysis system. We validated this low-cost system against a multicamera Vicon system and research-grade force platform (Advanced Mechanical Technology, Inc). After validation trials with known weights and angles, 5 subjects walked across an instrumented walkway for multiple times (n = 8/subject). We collected vertical GRF and segment angles. Average GRF data from the 2 systems were similar, with peak GRF errors below 3.5%BW. However, variability in the balance board’s sampling rate led to large GRF errors early and late in stance, when the GRF changed rapidly. The thigh, shank, and foot angle measurements were similar between the single and multicamera, but the pelvis angle was far less accurate. The proposed system has the potential to provide accurate segment angles and peak GRF at low cost but does not match the accuracy of the multicamera system and force platform, in part because of the Wii Balance Board’s variable sampling rate.

Bone development, maintenance, and regeneration are remarkably sensitive to mechanical cues. Consequently, mechanical stimulation has long been sought as a putative target to promote endogenous healing after fracture. Given the transient nature of bone repair, tissue-level mechanical cues evolve rapidly over time after injury and are challenging to measure noninvasively. The objective of this work was to develop and characterize an implantable strain sensor for noninvasive monitoring of axial strain across a rodent femoral defect during functional activity. Herein, we present the design, characterization, and in vivo demonstration of the device’s capabilities for quantitatively interrogating physiological dynamic strains during bone regeneration. Ex vivo experimental characterization of the device showed that it possessed promising sensitivity, signal resolution, and electromechanical stability for in vivo applications. The digital telemetry minimized power consumption, enabling extended intermittent data collection. Devices were implanted in a rat 6 mm femoral segmental defect model, and after three days, data were acquired wirelessly during ambulation and synchronized to corresponding radiographic videos, validating the ability of the sensor to noninvasively measure strain in real-time. Together, these data indicate the sensor is a promising technology to quantify tissue mechanics in a specimen specific manner, facilitating more detailed investigations into the role of the mechanical environment in dynamic bone healing and remodeling processes.

Bone development, maintenance, and regeneration are remarkably sensitive to mechanical cues. Consequently, mechanical stimulation has long been sought as a putative target to promote endogenous healing after fracture. Given the transient nature of bone repair, tissue-level mechanical cues evolve rapidly over time after injury and are challenging to measure non-invasively. The objective of this work was to develop and characterize an implantable strain sensor for non-invasive monitoring of axial strain across a rodent femoral defect during functional activity. Herein, we present the design, characterization, and in vivo demonstration of the device’s capabilities for quantitatively interrogating physiological dynamic strains during bone regeneration. Ex vivo experimental characterization of the device showed that it exceeded the technical requirements for sensitivity, signal resolution, and electromechanical stability. The digital telemetry minimized power consumption, enabling long-term intermittent data collection. Devices were implanted in a rat 6 mm femoral segmental defect model and after three days, data were acquired wirelessly during ambulation and synchronized to corresponding radiographic videos, validating the ability of the sensor to non-invasively measure strain in real-time. Lastly, in vivo strain measurements were utilized in a finite element model to estimate the strain distribution within the defect region. Together, these data indicate the sensor is a promising technology to quantify local tissue mechanics in a specimen specific manner, facilitating more detailed investigations into the role of the mechanical environment in dynamic skeletal healing and remodeling.

This file contains text with supplemental methods for the static analysis. It also has text legends for the supplementary videos and figures.

This file contains images of a disarticulated hip joint from a dissected flamingo carcass showing the relative association of the bony and cartilaginous features of the pelvis and proximal femur (S3). Additional images from a disarticulated flamingo museum specimen showing the prominent features of the knee joint (S4).

Afferent feedback alters muscle activity during locomotion and must be tightly controlled. As primary afferent depolarization-induced presynaptic inhibition (PAD-PSI) regulates afferent signaling, we investigated hindlimb PAD-PSI during locomotion in an in vitro rat spinal cord-hindlimb preparation. We compared the relation of PAD-PSI, measured as dorsal root potentials (DRPs), to observed ipsilateral and contralateral limb endpoint forces. Afferents activated during stance-phase force strongly and proportionately influenced DRP magnitude in the swinging limb. Responses increased with locomotor frequency. Electrical stimulation of contralateral afferents also preferentially evoked DRPs in the opposite limb during swing (flexion). Nerve lesioning, in conjunction with kinematic results, support a prominent contribution from toe Golgi tendon organ afferents. Thus, force-dependent afferent feedback during stance binds interlimb sensorimotor state to a proportional PAD-PSI in the swinging limb, presumably to optimize interlimb coordination. These results complement known actions of ipsilateral afferents on PAD-PSI during locomotion.

Minimizing whole body metabolic cost has been suggested to drive the neural processes of locomotor adaptation. Mechanical work performed by the legs should dictate the major changes in whole body metabolic cost of walking while providing greater insight into temporal and spatial mechanisms of adaptation. We hypothesized changes in mechanical work by the legs during an asymmetric split-belt walking adaptation task could explain previously observed changes in whole body metabolic cost. We predicted subjects would immediately increase mechanical work performed by the legs when first exposed to split-belt walking, followed by a gradual decrease throughout adaptation. Fourteen subjects walked on a dual-belt, instrumented treadmill. Baseline trials were followed by a ten-minute split-belt adaptation condition with one belt running three times faster than the other. A post-adaptation trial with both belts moving at 0.5m/s demonstrated neural adaptation. As predicted, summed mechanical work from both legs initially increased abruptly and gradually decreased over the adaptation period. The initial increase in work was primarily due to increased positive work by the leg on the fast belt during the pendular phase of the gait cycle. Neural adaptation in asymmetric split-belt walking reflected the reduction of pendular phase work in favor of more economical step-to-step transition work. This may represent a generalizable framework for how humans initially and chronically learn new walking patterns.

Human walking is a complex task, and we lack a complete understanding of how the neuromuscular system organizes its numerous muscles and joints to achieve consistent and efficient walking mechanics. Focused control of select influential task-level variables may simplify the higher-level control of steady-state walking and reduce demand on the neuromuscular system. As trailing leg power generation and force application can affect the mechanical efficiency of step-to-step transitions, we investigated how joint torques are organized to control leg force and leg power during human walking. We tested whether timing of trailing leg force control corresponded with timing of peak leg power generation. We also applied a modified uncontrolled manifold analysis to test whether individual or coordinated joint torque strategies most contributed to leg force control. We found that leg force magnitude was adjusted from step to step to maintain consistent leg power generation. Leg force modulation was primarily determined by adjustments in the timing of peak ankle plantar-flexion torque, while knee torque was simultaneously covaried to dampen the effect of ankle torque on leg force. We propose a coordinated joint torque control strategy in which the trailing leg ankle acts as a motor to drive leg power production while trailing leg knee torque acts as a brake to refine leg power production.

Presynaptic inhibition is a powerful mechanism for selectively and dynamically gating sensory inputs entering the spinal cord. We investigated how hindlimb mechanics influence presynaptic inhibition during locomotion using pioneering approaches in an in vitro spinal cord–hindlimb preparation. We recorded lumbar dorsal root potentials to measure primary afferent depolarization-mediated presynaptic inhibition and compared their dependence on hindlimb endpoint forces, motor output, and joint kinematics. We found that stance-phase force on the opposite limb, particularly at toe contact, strongly influenced the magnitude and timing of afferent presynaptic inhibition in the swinging limb. Presynaptic inhibition increased in proportion to opposite limb force, as well as locomotor frequency. This form of presynaptic inhibition binds the sensorimotor states of the two limbs, adjusting sensory inflow to the swing limb based on forces generated by the stance limb. Functionally, it may serve to adjust swing-phase sensory transmission based on locomotor task, speed, and step-to-step environmental perturbations.

The conventional method for filtering force plate data, low-pass filtering, does not always give accurate results when applied to force data from a custom-made, instrumented treadmill. Therefore, this study compares low-pass filtered data to the same data passed through a wavelet filter. We collected data with the treadmill running. However these include motor noise with ground reaction force at two force plates. We found that he proposed wavelet method eliminated motor noise to result in more accurate force plate data than the conventional low-pass filter, particularly at high speed motor operation. In this study we suggested the convolution wavelet (CNW) which was compared to that of a low-pass filter. The CNW showed better performance as compared to band-pass filtering particularly for low signal-to-noise ratios, and a lower computational load.

The voluntary EMG (vEMG) signal from electrically stimulated muscle is very useful for feedback control in functional electrical stimulation. However, the recorded EMG signal from surface electrodes has unwanted stimulation artifact and M-wave as well as vEMG. Here, we propose an event-synchronous adaptive digital filter for the suppression of stimulation artifact and M-wave in this application. The proposed method requires a simple experimental setup that does not require extra hardware connections to obtain the reference signals of adaptive digital filter. For evaluating the efficiency of this proposed method, the filter was tested and compared with a least square (LS) algorithm using previously measured data. We conclude that the cancellation of both primary and residual stimulation artifacts is enhanced with an event-synchronous adaptive digital filter and shows promise for clinical application to rehabilitate paretic limbs. Moreover because this algorithm is far simpler than the LS algorithm, it is portable and ready for real-time application.

Understanding how sensory feedback can be used to facilitate locomotion is vital for designing effective locomotor rehabilitation strategies for spinal cord injury. Here we describe an in vitro spinal cord-hindlimb preparation (SCHP) that can be used to answer clinically relevant questions about the interaction of sensory feedback and spinal locomotor circuitry. The SCHP, composed of the exposed rodent spinal cord with hindlimbs intact, combines the neural accessibility and manipulability of classic in vitro isolated cord preparations with intact sensory feedback and many of the behavioral analysis techniques used in the clinic, such as kinematics, treadmill walking, and electromyography. Using the SCHP, we demonstrate the importance of task-appropriate limb loading and posture for establishing intact rat-like kinematic trajectories and muscle activation patterns. We also show that applying stepping-like movements to the limbs, similar to those applied during physical therapist-assisted or robot-guided bo...

The sophistication of current rodent injury and disease models outpaces that of the most commonly used behavioral assays. The first objective of this study was to measure rat locomotion using high-speed X-ray video to establish an accurate baseline for rat hindlimb kinematics. The second objective was to quantify the kinematics errors due to skin movement artefacts by simultaneously recording and comparing hindlimb kinematics derived from skin markers and from direct visualization of skeletal landmarks. Joint angle calculations from skin-derived kinematics yielded errors as high as 39 degrees in the knee and 31 degrees in the hip around paw contact with respect to the X-ray data. Triangulation of knee position from the ankle and hip skin markers provided closer, albeit still inaccurate, approximations of bone-derived, X-ray kinematics. We found that soft tissue movement errors are the result of multiple factors, the most impressive of which is overall limb posture. Treadmill speed had surprisingly little effect on kinematics errors. These findings illustrate the significance and context of skin movement error in rodent kinematics.

SUMMARY Why is maximal running speed reduced on curved paths? The leading explanation proposes that an increase in lateral ground reaction force necessitates a decrease in peak vertical ground reaction force, assuming that maximum leg extension force is the limiting factor. Yet, no studies have directly measured these forces or tested this critical assumption. We measured maximum sprint velocities and ground reaction forces for five male humans sprinting along a straight track and compared them to sprints along circular tracks of 1, 2, 3, 4 and 6 m radii. Circular track sprint trials were performed either with or without a tether that applied centripetal force to the center of mass. Sprinters generated significantly smaller peak resultant ground reaction forces during normal curve sprinting compared to straight sprinting. This provides direct evidence against the idea that maximum leg extension force is always achieved and is the limiting factor. Use of the tether increased sprint speed, but not to expected values. During curve sprinting, the inside leg consistently generated smaller peak forces compared to the outside leg. Several competing biomechanical constraints placed on the stance leg during curve sprinting likely make the inside leg particularly ineffective at generating the ground reaction forces necessary to attain maximum velocities comparable to straight path sprinting. The ability of quadrupeds to redistribute function across multiple stance legs and decouple these multiple constraints may provide a distinct advantage for turning performance.

During movement, errors are typically corrected only if they hinder performance. Preferential correction of task-relevant deviations is described by the minimal intervention principle but has not been demonstrated in the joints during locomotor adaptation. We studied hopping as a tractable model of locomotor adaptation of the joints within the context of a limb-force-specific task space. Subjects hopped while adapting to shifted visual feedback that induced them to increase peak ground reaction force (GRF). We hypothesized subjects would preferentially reduce task-relevant joint torque deviations over task-irrelevant deviations to increase peak GRF. We employed a modified uncontrolled manifold analysis to quantify task-relevant and task-irrelevant joint torque deviations for each individual hop cycle. As would be expected by the explicit goal of the task, peak GRF errors decreased in early adaptation before reaching steady state during late adaptation. Interestingly, during the early adaptation performance improvement phase, subjects reduced GRF errors by decreasing only the task-relevant joint torque deviations. In contrast, during the late adaption performance maintenance phase, all torque deviations decreased in unison regardless of task relevance. In deadaptation, when the shift in visual feedback was removed, all torque deviations decreased in unison, possibly because performance improvement was too rapid to detect changes in only the task-relevant dimension. We conclude that limb force adaptation in hopping switches from a minimal intervention strategy during performance improvement to a noise reduction strategy during performance maintenance, which may represent a general control strategy for locomotor adaptation of limb force in other bouncing gaits, such as running.

How do arm-swinging apes locomote ef- fectively over a variety of speeds? One way to reduce the metabolic energy cost of locomotion is to transfer energy between reversible mechanical modes. In terrestrial ani- mals, at least two transfer mechanisms have been identi- fied: 1) a pendulum-like mechanism for walking, with exchange between gravitational potential energy and translational kinetic energy, and 2) a spring-like mecha- nism for running, where the elastic strain energy of stretched muscle and tendon is largely returned to reac- celerate the animal. At slower speeds, a brachiator will always have at least one limb in contact with the support, similar to the overlap of foot contact in bipedal walking. At faster speeds, brachiators exhibit an aerial phase, similar to that seen in bipedal running. Are there two distinct brachiation gaits even though the animal appears to sim- ply swing beneath its overhead support? If so, are differ- ent exchange mechanisms employed? Our kinetic analysis of brachiation in a white-handed gibbon (Hylobates lar) indicates that brachiation is indeed comprised of two me- chanically distinct gaits. At slower speeds in "continuous contact" brachiation, the gibbon utilizes a simple pendu- lum-like transfer of mechanical energy within each stride. At faster speeds in "ricochetal" brachiation, translational and rotational kinetic energy are exchanged in a novel "whip-like" transfer. We propose that brachiators utilize the transfer between translational and rotational kinetic energy to control the dynamics of their swing. This ma- neuver may allow muscle action at the shoulder to control the transfer and adjust the ballistic portion of the step to meet the requirements for the next hand contact.Am J Phys Anthropol 115:319 -326, 2001. © 2001 Wiley-Liss, Inc.

We compared the kinetics of brachiation to bipedal walking and running. Gibbons use pectoral limbs in continuous contact with their overhead support at slow speeds, but exhibit aerial phases (or ricochetal brachiation) at faster speeds. This basic interaction between limb and support suggests some analogy to walking and running. We quantified the forces in three axes and torque about the vertical axis generated by a brachiating White-handed gibbon (Hylobates lar) and compared them with bipedal locomotion. Handholds oriented perpendicular to the direction of travel (as in ladder rungs) were spaced 0.80, 1.20, 1.60, 1.72, 1.95, and 2.25 m apart. The gibbon proportionally matched forward velocity to stride length. Handhold reaction forces resembled ground reaction forces of running humans except that the order of horizontal braking and propulsion were reversed. Peak vertical forces in brachiation increased with speed as in bipedal locomotion. In contrast to bipedalism, however, peak horizontal forces changed little with speed. Gait transition occurred within the same relative velocity range as the walk-run transition in bipeds (Froude number = 0.3-0.6). We oriented handholds parallel to the direction of travel (as in a continuous pole) at 0.80 and 1.60 m spacings. In ricochetal brachiation, the gibbon generated greater torque with handholds oriented perpendicular as opposed to parallel to the direction of travel. Handhold orientation did not affect peak forces. The similarities and differences between brachiation and bipedalism offer insight into the ubiquity of mechanical principles guiding all limbed locomotion and the distinctiveness of brachiation as a unique mode of locomotion.

Previous studies have suggested that generating vertical force on the ground to support body weight (BWt) is the major determinant of the metabolic cost of running. Because horizontal forces exerted on the ground are often an order of magnitude smaller than vertical forces, some have reasoned that they have negligible cost. Using applied horizontal forces (AHF; negative is impeding, positive is aiding) equal to −6, −3, 0, +3, +6, +9, +12, and +15% of BWt, we estimated the cost of generating horizontal forces while subjects were running at 3.3 m/s. We measured rates of oxygen consumption (V˙o2) for eight subjects. We then used a force-measuring treadmill to measure ground reaction forces from another eight subjects. With an AHF of −6% BWt,V˙o2increased 30% compared with normal running, presumably because of the extra work involved. With an AHF of +15% BWt, the subjects exerted ∼70% less propulsive impulse and exhibited a 33% reduction inV˙o2. Our data suggest that generating horizontal propulsive forces constitutes more than one-third of the total metabolic cost of normal running.

Locomotion persists across all manner of internal and external perturbations. The objective of this study was to identify locomotor compensation strategies in rodent models of peripheral nerve injury. We found that hip-to-toe limb length and limb angle was preferentially preserved over individual joint angles after permanent denervation of rat ankle extensor muscles. These findings promote further enquiry into the significance of limb-level function for neuromechanical control of legged locomotion.

Humans can robustly locomote over complex terrains even while simultaneously attending to other tasks such as accurate foot placement on the ground. We investigated whether subjects would exploit motor redundancy across the joints of the leg to stabilize overall limb kinematics when presented with a hopping task that constrained foot placement position. Subjects hopped in place on one leg (2.2 Hz) while having to place their foot into one of three target sizes upon landing (0.250, 0.063, 0.010 m(2)). As takeoff and landing angles are critical to this task performance, we hypothesized smaller target sizes would increase the need to stabilize (i.e., make more consistent) the leg orientation through motor equivalent combinations of segment angles. As it was not critical to the targeting task, we hypothesized no changes for leg length stabilization across target size. With smaller target sizes, we saw total segment angle variance increase due to greater signal-dependent noise associated with an increased activation of leg extensor muscles (medial and lateral gastrocnemius, vastus medialis, vastus lateralis and rectus femoris). At smaller target sizes, more segment angle variance was aligned to kinematic deviations with the goal of maintaining leg orientation trajectory. We also observed a decrease in the variance structure for stabilizing leg length at the smallest target conditions. This trade-off effect is explained by the nearly orthogonal relationship between the two goal-equivalent manifolds for leg length vs. leg orientation stabilization. Our results suggest humans increasingly rely on kinematic redundancy in their legs to achieve robust, consistent locomotion when faced with novel conditions that constrain performance requirements. These principles may generalize to other human locomotor gaits and provide important insights into the control of the legs during human walking and running.

Human walking dynamics are typically framed in the context of mechanics and energetics rather than in the context of neuromuscular control. Dynamic walking principles describe one helpful theoretical approach to characterize efficient human walking mechanics over many steps. These principles do not, however, address how such walking is controlled step-by-step despite small perturbations from natural variability. Our purpose was to identify neuromechanical control strategies used to achieve consistent and robust locomotion despite natural step-to-step force variability. We used the uncontrolled manifold concept to test whether human walkers select combinations of leading and trailing leg-forces that generate equivalent net-force trajectories during step-to-step transitions. Subjects selected leading and trailing leg-force combinations that generated consistent vertical net-force during step-to-step transitions. We conclude that vertical net-force is an implicit neuromechanical goal of human walking whose trajectory is stabilized for consistent step-to-step transitions, which agrees with the principles of dynamic walking. In contrast, inter-leg-force combinations modulated anterior-posterior net-force trajectories with each step to maintain constant walking speed, indicating that a consistent anterior-posterior net-force trajectory is not an implicit goal of walking. For a more complete picture of hierarchical locomotor control, we also tested whether each individual leg-force trajectory was stabilized through the selection of leg-force equivalent joint-torque combinations. The observed consistent vertical net-force trajectory was achieved primarily through the selection of joint-torque combinations that modulated trailing leg-force during step-to-step transitions. We conclude that humans achieve robust walking by harnessing inherent motor abundance of the joints and legs to maintain consistent step-by-step walking performance.

The neonatal rodent spinal cord maintained in vitro is a powerful model system to understand the central properties of spinal circuits generating mammalian locomotion. We describe three enabling approaches that incorporate afferent input and attached hindlimbs. (i) Sacral dorsal column stimulation recruits and strengthens ongoing locomotor-like activity, and implementation of a closed positive-feedback paradigm is shown to support its stimulation as an untapped therapeutic site for locomotor modulation. (ii) The spinal cord hindlimbs-restrained preparation allows suction electrode electromyographic recordings from many muscles. Inducible complex motor patterns resemble natural locomotion, and insights into circuit organization are demonstrated during spontaneous motor burst 'deletions', or following sensory stimuli such as tail and paw pinch. (iii) The spinal cord hindlimbs-pendant preparation produces unrestrained hindlimb stepping. It incorporates mechanical limb perturbations, kinematic analyses, ground reaction force monitoring, and the use of treadmills to study spinal circuit operation with movement-related patterns of sensory feedback while providing for stable whole-cell recordings from spinal neurons. Such techniques promise to provide important additional insights into locomotor circuit organization.