Three-dimensional knee joint contact forces during walking in unilateral transtibial amputees

Individuals with unilateral transtibial amputations have greater prevalence of osteoarthritis in the intact knee joint relative to the residual leg and non-amputees, but the cause of this greater prevalence is unclear. The purpose of this study was to compare knee joint contact forces and the muscles contributing to these forces between amputees and non-amputees during walking using forward dynamics simulations. We predicted that the intact knee contact forces would be higher than those of the residual leg and non-amputees. In the axial and mediolateral directions, the intact and non-amputee legs had greater peak tibio-femoral contact forces and impulses relative to the residual leg. The peak axial contact force was greater in the intact leg relative to the non-amputee leg, but the stance phase impulse was greater in the non-amputee leg. The vasti and hamstrings muscles in early stance and gastrocnemius in late stance were the largest contributors to the joint contact forces in the non-amputee and intact legs. Through dynamic coupling, the soleus and gluteus medius also had large contributions, even though they do not span the knee joint. In the residual leg, the prosthesis had large contributions to the joint forces, similar to the soleus in the intact and non-amputee legs. These results identify the muscles that contribute to knee joint contact forces during transtibial amputee walking and suggest that the peak knee contact forces may be more important than the knee contact impulses in explaining the high prevalence of intact leg osteoarthritis.     

Development of an above-knee prosthesis equipped with a microcomputer-controlled knee joint: first test results

The shortcomings of conventional above-knee prostheses are due to their lack of adaptive control. Implementation of a microcomputer controlling the knee joint in a passive way has been suggested to enhance the patient's gait comfort, safety and cosmesis. This approach was used in the design of a new prosthetic system for the above-knee amputee, and tested on one patient. The knee joint of a conventional, modular prosthesis was replaced by a knee joint mechanism, equipped with a controllable brake on the knee joint axis. Sensors and a microcomputer were added, keeping the system self-contained. The modularity of the design permits the use of an alternative, external, PC-based control unit, emulating the self-contained one, and offering extended data monitoring and storage facilities. For both units an operating environment was written, including sensor/actuator interfacing and the implementation of a real-time interrupt, executing the control algorithm. A double finite state approach was used in the design of the control algorithm. On a higher level, the mode identification algorithm reveals the patient's intent. Within a specific mode (lower level), the relevant mode control algorithm looks for the current phase within the gait cycle. Within a particular phase, a specific simple control action with the brake replaces normal knee muscle activity. Test were carried out with one prosthetic patient using a basic control algorithm for level walking, allowing controlled knee flexion during stance phase. The technical feasibility of such a concept is illustrated by the test results, even though only flexion during early stance phase was controlled during the trials. Patient acceptance is not straightforward since knee flexion during stance phase is associated with knee buckling.     

Stance phase control of above-knee prostheses: knee control versus SACH foot design

The mobility of above-knee amputees (A/K) is limited, in part, due to the performance of A/K prostheses during the stance phase. Currently stance phase control of most conventional A/K prostheses can only be achieved through leg alignment and choice of the SACH (Solid Ankle Cushioned Heel) foot. This paper examines the role of the knee controller in relation to a SACH foot during the stance phase of level walking. The three-dimensional gait mechanics were measured under two stance phase conditions. In the first set of trials, the amputee used a prosthesis with a conventional knee controller that allowed the amputee to maintain the knee joint in full extension during the stance phase. In the second set of trials, the prosthetic knee, during stance, echoed the modified kinematics of the amputee's sound (intact) knee that had been recorded during the previous sound stance phase. Analysis and interpretation of the data indicate the following: (1) SACH foot design can strongly influence the walking mechanics independent of the knee controller; (2) knee controller design and SACH foot design are mutually interdependent; and (3) normal kinematics imposed on the prosthetic knee does not necessarily produce normal hip kinematics (e.g. reduce the abnormal rise in the prosthetic side hip trajectory). Future research is necessary to explore and exploit the interdependency of prosthetic knee control and foot design.     

Muscle and prosthesis contributions to amputee walking mechanics: A modeling study

Unilateral, below-knee amputees have altered gait mechanics, which can significantly affect their mobility. Below-knee amputees lose the functional use of the ankle muscles, which are critical during walking to provide body support, forward propulsion, leg-swing initiation and mediolateral balance. Thus, either muscles must compensate or the prosthesis must provide the functional tasks normally provided by the ankle muscles. Three-dimensional (3D) forward dynamics simulations of amputee and non-amputee walking were generated to identify muscle and prosthesis contributions to amputee walking mechanics, including the subtasks of body support, forward propulsion, leg-swing initiation and mediolateral balance. Results showed that the prosthesis provided body support in the absence of the ankle muscles. The prosthesis contributed to braking from early to mid-stance and propulsion in late stance. The prosthesis also functioned like the uniarticular soleus muscle by transferring energy from the residual leg to the trunk to provide trunk propulsion. The residual-leg vasti and rectus femoris reduced their contributions to braking in early stance, which mitigated braking from the prosthesis during this period. The prosthesis did not replace the function of the gastrocnemius, which normally generates energy to the leg to initiate swing. As a result, lower overall energy was delivered to the residual leg. The prosthesis also acted to accelerate the body laterally in the absence of the ankle muscles. These results provide further insight into muscle and prosthesis function in below-knee amputee walking and can help guide rehabilitation methods and device designs to improve amputee mobility.     

The effect of direct measurement versus cadaver estimates of anthropometry in the calculation of joint moments during above-knee prosthetic gait in pediatrics

Joint reaction forces, moments and powers are important in interpreting gait mechanics and compensatory strategies used by patients walking with above-knee prostheses. Segmental anthropometrics, required to calculate joint moments, are often estimated using data from cadaver studies. However, these values may not be accurate for patients following amputation as prostheses are composed of non-biologic material. The purpose of this study was to compare joint moments using anthropometrics calculated from cadaver studies versus direct measurements of the residual limb and prosthesis for children with an above-knee amputation. Gait data were collected for four subjects with above-knee prostheses walking at preferred and fast speeds. Joint moments were computed using anthropometrics from cadaver studies and direct measurements for each subject. The difference between these two methods primarily affected the inertia couple (Ialpha term) and the inertial effect due to gravity, which comprised a greater percentage of the total joint moment during swing as compared to stance. Peak hip and knee flexor and extensor moments during swing were significantly greater when calculated using cadaver data (p<0.05). These differences were greater while walking fast as compared to slow speeds. A significant difference was not found between these two methods for peak hip and knee moments during stance. A significant difference was found for peak ankle joint moments during stance, but the magnitude was not clinically important. These results support the use of direct measurements of anthropometry when examining above-knee prosthetic gait, particularly during swing.     

Mechanical energetic contributions from individual muscles and elastic prosthetic feet during symmetric unilateral transtibial amputee walking: a theoretical study

Energy storage and return (ESAR) foot-ankle prostheses have been developed in an effort to improve gait performance in lower-limb amputees. However, little is known about their effectiveness in providing the body segment mechanical energetics normally provided by the ankle muscles. The objective of this theoretical study was to use muscle-actuated forward dynamics simulations of unilateral transtibial amputee and non-amputee walking to identify the contributions of ESAR prostheses to trunk support, forward propulsion and leg swing initiation and how individual muscles must compensate in order to produce a normal, symmetric gait pattern. The simulation analysis revealed the ESAR prosthesis provided the necessary trunk support, but it could not provide the net trunk forward propulsion normally provided by the plantar flexors and leg swing initiation normally provided by the biarticular gastrocnemius. To compensate, the residual leg gluteus maximus and rectus femoris delivered increased energy to the trunk for forward propulsion in early stance and late stance into pre-swing, respectively, while the residual iliopsoas delivered increased energy to the leg in pre- and early swing to help initiate swing. In the intact leg, the soleus, gluteus maximus and rectus femoris delivered increased energy to the trunk for forward propulsion in the first half of stance, while the iliopsoas increased the leg energy it delivered in pre- and early swing. Thus, the energy stored and released by the ESAR prosthesis combined with these muscle compensations was able to produce a normal, symmetric gait pattern, although various neuromuscular and musculoskeletal constraints may make such a pattern non-optimal.     

Analysis of a new polycentric above-knee prosthesis with a pneumatic swing phase control

The four-bar linkage mechanism used by many designers for the knee joint of an above-knee prosthesis can provide knee rotation suitable for walking only. In Afro-Asian countries people are habituated to the squatting posture in their daily activities. A six-bar linkage knee-ankle mechanism above-knee prosthesis with a pneumatic swing phase control is developed and its analysis shows its suitability for producing a motion closely resembling that of a normal subject during walking and squatting.     

Forces and moments telemetered from two distal femoral replacements during various activities.

Two distal femoral replacements were instrumented to measure axial force, torque and bending moments in the prosthesis shaft. Data are reported up to 2.5 years for the following activities: uni- and bi-lateral standing, walking, stair climbing and descending, treadmill walking, jogging and jumping. In the first subject the greatest averaged peak shaft forces found were: jogging 3.6Bodyweight (BW), stair descending 3.1BW, walking 2.8BW, treadmill walking 2.75BW, and stair ascending 2.8BW. Bending moments about the antero-posterior axis (varus-valgus) and medio-lateral axis (flexion-extension) peaked in the range 8.5-9.8 and 4.7-7.6BWcm respectively, over the follow-up period. Axial torques peaked in the range 0.2-1.3BWcm, outwardly directed. At most follow-up sessions, forces and moments during jogging were generally greater than those for other gait activities. In the second subject forces and moments were generally only 45-70% of those in the first subject, due to inadequate musculature around the knee. The data can be applied to the design and testing of distal femoral replacements and even to total knee replacements, and contributes to the knowledge of forces acting in the distal femur during activity.     

Adaptative control of above knee electro-hydraulic prosthesis.

Conventional designs of an above-knee prosthesis are based on mechanisms with mechanical properties (such as friction, spring and damping coefficients) that remain constant during changing cadence. These designs are unable to replace natural legs due to the lack of active knee joint control. Since the nonlinear and time-varying dynamic coupling between the thigh and the prosthetic limb is high during swing phase, an adaptive control is employed to control the knee joint motion. Two dimensional simulation indicates that the adaptive controller can improve the appearance of gait pattern. It is adaptable to walking speed and can compensate for the variations of hip moment, hip trajectory and toe-off conditions.     

Biomechanics of below-knee amputee gait

Sagittal plane biomechanical and EMG analyses from eight below knee (B/K) amputee trials demonstrate considerably modified motor patterns from the residual muscles at the hip and knee. Five SACH fittings, two Uniaxial and one Gressinger prostheses were analysed. Moments of force and mechanical power were analysed on all eight trials and EMG profiles are reported for three of the amputees fitted with SACH prostheses. The findings can be summarized as follows: 1. All eight trials had similar internal moment of force patterns at the ankle. A dorsiflexor moment commenced at heel contact and continued for the first third of stance. The prostheses generated a plantarflexor moment for the balance of stance which increased in late stance to about 2/3 that seen in normals. 2. The two Uniaxial prostheses showed a 20% recovery of stored energy which was returned at push-off. The recovery by the Gressinger fitting was 30%. 3. For all but the Gressinger prosthesis the knee moment of force was negligible during early stance (when normals have an extensor moment), below normal in late stance and fairly normal during swing. The amputee wearing the Gressinger prosthesis had a normal but slightly reduced pattern of moments of force over the entire stride. 4. All eight trials had hyperactive hip extensors during early and mid-stance which resulted in above-normal energy generation by these concentrically contracting muscles. This compensation makes up for the loss of the major energy generation by the plantarflexors at push-off. 5. The moment of force and power patterns at the hip for all eight trials during late stance and swing were fairly normal.(ABSTRACT TRUNCATED AT 250 WORDS)     

Energetics and passive dynamics of the ankle in downhill walking

This study investigated the energetics of the human ankle during the stance phase of downhill walking with the goal of modeling ankle behavior with a passive spring and damper mechanism. Kinematic and kinetic data were collected on eight male participants while walking down a ramp with inclination varying from 0° to 8°. The ankle joint moment in the sagittal plane was calculated using inverse dynamics. Mechanical energy injected or dissipated at the ankle joint was computed by integrating the power across the duration of the stance phase. The net mechanical energy of the ankle was approximately zero for level walking and monotonically decreased (i.e., became increasingly negative) during downhill walking as the slope decreased. The indication is that the behavior of the ankle is energetically passive during downhill walking, playing a key role in dissipating energy from one step to the next. A passive mechanical model consisting of a pin joint coupled with a revolute spring and damper was fit to the ankle torque and its parameters were estimated for each downhill slope using linear regression. The passive model demonstrated good agreement with actual ankle dynamics as indicated by low root-mean-square error values. These results indicate the stance phase behavior of the human ankle during downhill walking may be effectively duplicated by a passive mechanism with appropriately selected spring and damping characteristics.     

Muscle and external load contribution to knee joint contact loads during normal gait

Large knee adduction moments during gait have been implicated as a mechanical factor related to the progression and severity of tibiofemoral osteoarthritis and it has been proposed that these moments increase the load on the medial compartment of the knee joint. However, this mechanism cannot be validated without taking into account the internal forces and moments generated by the muscles and ligaments, which cannot be easily measured. Previous musculoskeletal models suggest that the medial compartment of the tibiofemoral joint bears the majority of the tibiofemoral load, with the lateral compartment unloaded at times during stance. Yet these models did not utilise explicitly measured muscle activation patterns and measurements from an instrumented prosthesis which do not portray lateral compartment unloading. This paper utilised an EMG-driven model to estimate muscle forces and knee joint contact forces during healthy gait. Results indicate that while the medial compartment does bear the majority of the load during stance, muscles provide sufficient stability to counter the tendency of the external adduction moment to unload the lateral compartment. This stability was predominantly provided by the quadriceps, hamstrings, and gastrocnemii muscles, although the contribution from the tensor fascia latae was also significant. Lateral compartment unloading was not predicted by the EMG-driven model, suggesting that muscle activity patterns provide useful input to estimate muscle and joint contact forces.     

Load transfer mechanics between trans-tibial prosthetic socket and residual limb--dynamic effects

The effects of inertial loads on the interface stresses between trans-tibial residual limb and prosthetic socket were investigated. The motion of the limb and prosthesis was monitored using a Vicon motion analysis system and the ground reaction force was measured by a force platform. Equivalent loads at the knee joint during walking were calculated in two cases with and without consideration of the material inertia. A 3D nonlinear finite element (FE) model based on the actual geometry of residual limb, internal bones and socket liner was developed to study the mechanical interaction between socket and residual limb during walking. To simulate the friction/slip boundary conditions between the skin and liner, automated surface-to-surface contact was used. The prediction results indicated that interface pressure and shear stress had the similar double-peaked waveform shape in stance phase. The average difference in interface stresses between the two cases with and without consideration of inertial forces was 8.4% in stance phase and 20.1% in swing phase. The maximum difference during stance phase is up to 19%. This suggests that it is preferable to consider the material inertia effect in a fully dynamic FE model.     

Design and Testing of a Bionic Dancing Prosthesis

Traditionally, prosthetic leg research has focused on improving mobility for activities of daily living. Artistic expression such as dance, however, is not a common research topic and consequently prosthetic technology for dance has been severely limited for the disabled. This work focuses on investigating the ankle joint kinetics and kinematics during a Latin-American dance to provide unique motor options for disabled individuals beyond those of daily living. The objective of this study was to develop a control system for a bionic ankle prosthesis that outperforms conventional prostheses when dancing the rumba. The biomechanics of the ankle joint of a non-amputee, professional dancer were acquired for the development of the bionic control system. Subsequently, a professional dancer who received a traumatic transtibial amputation in April 2013 tested the bionic dance prosthesis and a conventional, passive prosthesis for comparison. The ability to provide similar torque-angle behavior of the biological ankle was assessed to quantify the biological realism of the prostheses. The bionic dancing prosthesis overlapped with 37 ± 6% of the non-amputee ankle torque and ankle angle data, compared to 26 ± 2% for the conventional, passive prosthesis, a statistically greater overlap (p = 0.01). This study lays the foundation for quantifying unique, expressive activity modes currently unavailable to individuals with disabilities. Future work will focus on an expansion of the methods and types of dance investigated in this work.     

Patterns of mechanical energy change in tetrapod gait: pendula, springs and work

Kinematic and center of mass (CoM) mechanical variables used to define terrestrial gaits are compared for various tetrapod species. Kinematic variables (limb phase, duty factor) provide important timing information regarding the neural control and limb coordination of various gaits. Whereas, mechanical variables (potential and kinetic energy relative phase, %Recovery, %Congruity) provide insight into the underlying mechanisms that minimize muscle work and the metabolic cost of locomotion, and also influence neural control strategies. Two basic mechanisms identified by Cavagna et al. (1977. Am J Physiol 233:R243-R261) are used broadly by various bipedal and quadrupedal species. During walking, animals exchange CoM potential energy (PE) with kinetic energy (KE) via an inverted pendulum mechanism to reduce muscle work. During the stance period of running (including trotting, hopping and galloping) gaits, animals convert PE and KE into elastic strain energy in spring elements of the limbs and trunk and regain this energy later during limb support. The bouncing motion of the body on the support limb(s) is well represented by a simple mass-spring system. Limb spring compliance allows the storage and return of elastic energy to reduce muscle work. These two distinct patterns of CoM mechanical energy exchange are fairly well correlated with kinematic distinctions of limb movement patterns associated with gait change. However, in some cases such correlations can be misleading. When running (or trotting) at low speeds many animals lack an aerial period and have limb duty factors that exceed 0.5. Rather than interpreting this as a change of gait, the underlying mechanics of the body's CoM motion indicate no fundamental change in limb movement pattern or CoM dynamics has occurred. Nevertheless, the idealized, distinctive patterns of CoM energy fluctuation predicted by an inverted pendulum for walking and a bouncing mass spring for running are often not clear cut, especially for less cursorial species. When the kinematic and mechanical patterns of a broader diversity of quadrupeds and bipeds are compared, more complex patterns emerge, indicating that some animals may combine walking and running mechanics at intermediate speeds or at very large size. These models also ignore energy costs that are likely associated with the opposing action of limbs that have overlapping support times during walking. A recent model of terrestrial gait (Ruina et al., 2005. J Theor Biol, in press) that treats limb contact with the ground in terms of collisional energy loss indicates that considerable CoM energy can be conserved simply by matching the path of CoM motion perpendicular to limb ground force. This model, coupled with the earlier ones of pendular exchange during walking and mass-spring elastic energy savings during running, provides compelling argument for the view that the legged locomotion of quadrupeds and other terrestrial animals has generally evolved to minimize muscle work during steady level movement.     

Approximate Perturbation Stance Map of the SLIP Runner and Application to Locomotion Control

This paper presents a novel method of perturbation to obtain the analytic approximate solution to the Spring-Loaded Inverted Pendulum (SLIP) dynamics in stance phase with considering the effect of gravity.This perturbation solution achieves higher accuracy in predicting the apex state variables than the typical existing analytic approximations.Particularly,our solution is validated for non-symmetric trajectory of hopping in a large angle range.Furthermore,the stance controller of the SLIP runner is developed to regulate the apex state based on the approximate apex return map.To compensate the energy variation between the current and desired apex states,a stiffness adjustment of the leg spring in stance phase is presented.The deadbeat controller of the angle of attack is designed to track the regulated apex height and velocity.The simulation demonstrates that the SLIP runner applying the proposed stance controller reveals higher tracking accuracy and more rapidly converges to the regulated apex state.     

Gimbals in the insect leg

We studied the common kinematic features of the coxa and trochanter in cursorial and raptorial legs, which are the short size of the podomers, predominantly monoaxial joints, and the approximate orthogonality of adjacent joint axes. The chain coxa-trochanter with its short elements and serial orthogonality of joint axes resembles the gimbals which combine versatility and tolerance to external perturbations. The geometry of legs was studied in 23 insect species of 12 orders. Insects with monoaxial joints were selected. The joint between the trochanter and the femur (TFJ) is defined either by two vestigial condyles or by a straight anterior hinge. Direction of the joint axes in the two basal podomers was assessed by 3D measurements or by goniometry in two planes. Length of the coxa is <15% (mostly <8%) of the total length of the cursorial leg, that of the trochanter <10%. Angles between the proximal and distal joint axes in the middle coxa range from 124 to 84 degrees (mean 97+/-14 degrees ), in the trochanter (in all legs studied) from 125 to 72 degrees (mean 90+/-13 degrees ). Vectors of the distal axis in the coxa are concentrated about the normal to the plane defined by the proximal axis and the midpoint between the distal condyles. These vectors in the trochanter lie at various angles to the normal; angles are correlated with the direction of the TFJ relative to the femur. Range of reduction about the TFJ is over 60 degrees in the foreleg of Ranatra linearis, Mantispa lobata and the hind leg in Carabus coriaceus (confirming observations of previous authors), 40-60 degrees in the foreleg of Vespa crabro and in the middle one in Ammophila campestris, 10-30 degrees in other studied specimens. The special role of the trochanter in autotomy and in active propulsion in some insect groups is discussed. The majority of insects possess small trochanters and slightly movable TFJs with the joint axis laying in the femur-tibia plane. We pose the hypothesis that the TFJ damps external forces, the vectors of which lie off the femur-tibia plane, the reductor muscle acting as a spring. Thus the TFJ contributes to dynamic stability of legged locomotion.     

Multi-functionality of the cat medical gastrocnemius during locomotion

The functional role of biarticular muscles was investigated based on direct force measurement in the cat medial gastrocnemius (MG) and analysis of hindlimb kinematics and kinetics for the stance phase of level, uphill, and downhill walking. Four primary functional roles of biarticular muscles have been proposed in the past. These functional roles have typically been discussed independently of each other, and biarticular muscles have rarely been assigned more than one functional roles for different phases of the work cycle. The purpose of this study was to elucidate the functional role of the biarticular cat MG during locomotion. It was found that MG forces were primarily associated with the moment requirements at the ankle for most of the stance phase, but also helped to satisfy the moments at the knee in the initial phase of stance. In the second half of stance, MG transferred mechanical energy from the knee to the ankle from the knee to the ankle, while simultaneously producing a substantial amount of mechanical work. Based on these results, we hypothesize that MG's primary function is that of an ankle extensor. However, because of the coupling of the ankle extensor moment with a knee flexor moment in the initial, and a knee extensor moment in the final phase of stance, MG satisfies two joint moments in early stance, and transfers mechanical energy from the knee to the ankle in late stance. We conclude that cat MG has multiple functional roles during the stance phase of locomotion, and speculate that such multi-functionality also exists in other bi- and multi-articular muscles.     

A Phase-Dependent Hypothesis for Locomotor Functions of Human Foot Complex

The human foot is a very complex structure comprising numerous bones, muscles, ligaments and synovial joints. As the only component in contact with the ground, the foot complex delivers a variety of biomechanical functions during human locomotion, e.g. body support and propulsion, stability maintenance and impact absorption. These need the human foot to be rigid and damped to transmit ground reaction forces to the upper body and maintain body stability, and also to be compliant and resilient to moderate risky impacts and save energy. How does the human foot achieve these apparent conflicting functions？ In this study, we propose a phase-dependent hypothesis for the overall locomotor functions of the human foot complex based on in-vivo measurements of human natural gait and simulation results of a mathematical foot model. We propse that foot functions are highly dependent on gait phase, which is a major characteristics of human locomotion. In early stance just after heel strike, the foot mainly works as a shock absorber by moderating high impacts using the viscouselastic heel pad in both vertical and horizontal directions. In mid-stance phase （-80% of stance phase）, the foot complex can be considered as a springy rocker, reserving external mechanical work using the foot arch whilst moving ground contact point forward along a curved path to maintain body stability. In late stance after heel off, the foot complex mainly serves as a force modulator like a gear box, modulating effective mechanical advantages of ankle plantiflexor muscles using metatarsal-phalangeal joints. A sound under- standing of how diverse functions are implemented in a simple foot segment during human locomotion might be useful to gain insight into the overall foot locomotor functions and hence to facilitate clinical diagnosis, rehabilitation product design and humanoid robot development.     

Three-dimensional kinematics of capuchin monkey bipedalism

Capuchin monkeys are known to use bipedalism when transporting food items and tools. The bipedal gait of two capuchin monkeys in the laboratory was studied with three-dimensional kinematics. Capuchins progress bipedally with a bent-hip, bent-knee gait. The knee collapses into flexion during stance and the hip drops in height. The knee is also highly flexed during swing to allow the foot which is plantarflexed to clear the ground. The forefoot makes first contact at touchdown. Stride frequency is high, and stride length and limb excursion low. Hind limb retraction is limited, presumably to reduce the pitch moment of the forward-leaning trunk. Unlike human bipedalism, the bipedal gait of capuchins is not a vaulting gait, and energy recovery from pendulum-like exchanges is unlikely. It extends into speeds at which humans and other animals run, but without a human-like gait transition. In this respect it resembles avian bipedal gaits. It remains to be tested whether energy is recovered through cyclic elastic storage and release as in bipedal birds at higher speeds. Capuchin bipedalism has many features in common with the facultative bipedalism of other primates which is further evidence for restrictions on a fully upright striding gait in primates that transition to bipedalism. It differs from the facultative bipedalism of other primates in the lack of an extended double-support phase and short aerial phases at higher speeds that make it a run by kinematic definition. This demonstrates that facultative bipedalism of quadrupedal primates need not necessarily be a walking gait.