Center of mass mechanics of chimpanzee bipedal walking

Center of mass (CoM) oscillations were documented for 81 bipedal walking strides of three chimpanzees. Full‐stride ground reaction forces were recorded as well as kinematic data to synchronize force to gait events and to determine speed. Despite being a bent‐hip, bent‐knee (BHBK) gait, chimpanzee walking uses pendulum‐like motion with vertical oscillations of the CoM that are similar in pattern and relative magnitude to those of humans. Maximum height is achieved during single support and minimum height during double support. The mediolateral oscillations of the CoM are more pronounced relative to stature than in human walking when compared at the same Froude speed. Despite the pendular nature of chimpanzee bipedalism, energy recoveries from exchanges of kinetic and potential energies are low on average and highly variable. This variability is probably related to the poor phasic coordination of energy fluctuations in these facultatively bipedal animals. The work on the CoM per unit mass and distance (mechanical cost of transport) is higher than that in humans, but lower than that in bipedally walking monkeys and gibbons. The pronounced side sway is not passive, but constitutes 10% of the total work of lifting and accelerating the CoM. CoM oscillations of bipedally walking chimpanzees are distinctly different from those of BHBK gait of humans with a flat trajectory, but this is often described as “chimpanzee‐like” walking. Human BHBK gait is a poor model for chimpanzee bipedal walking and offers limited insights for reconstructing early hominin gait evolution. Am J Phys Anthropol 156:422–433, 2015. © 2014 Wiley Periodicals, Inc.     

Biomechanical analysis of primate bipedal walking by computer simulation

A computer simulation technique was applied to make clear the mechanical characteristics of primate bipedal walking. A primate body and the walking mechanism were modeled mathematically with a set of dynamic equations. Using a digital computer, the following were calculated from these equations by substituting measured displacements and morphological data of each segment of the primate: the acceleration, joint angle, center of gravity, foot force, joint moment, muscular force, transmitted force at the joint, electric activity of the muscle, generated power by the leg and energy expenditure in walking.The model was evaluated by comparing some of the calculated results with the experimental results such as foot force and electromyographic data, and improved in order to obtain the agreement between them.The level bipedal walking of man, chimpanzee and Japanese monkey and several types of synthesized walking were analyzed from the viewpoint of biomechanics.It is concluded that the bipedal walking of chimpanzee is nearer to that of man than to that of the Japanese monkey because of its propulsive mechanism, but it requires large muscular force for supporting the body weight.     

Compliant bipedal model with the center of pressure excursion associated with oscillatory behavior of the center of mass reproduces the human gait dynamics

Although the compliant bipedal model could reproduce qualitative ground reaction force (GRF) of human walking, the model with a fixed pivot showed overestimations in stance leg rotation and the ratio of horizontal to vertical GRF. The human walking data showed a continuous forward progression of the center of pressure (CoP) during the stance phase and the suspension of the CoP near the forefoot before the onset of step transition. To better describe human gait dynamics with a minimal expense of model complexity, we proposed a compliant bipedal model with the accelerated pivot which associated the CoP excursion with the oscillatory behavior of the center of mass (CoM) with the existing simulation parameter and leg stiffness. Owing to the pivot acceleration defined to emulate human CoP profile, the arrival of the CoP at the limit of the stance foot over the single stance duration initiated the step-to-step transition. The proposed model showed an improved match of walking data. As the forward motion of CoM during single stance was partly accounted by forward pivot translation, the previously overestimated rotation of the stance leg was reduced and the corresponding horizontal GRF became closer to human data. The walking solutions of the model ranged over higher speed ranges (~1.7 m/s) than those of the fixed pivoted compliant bipedal model (~1.5 m/s) and exhibited other gait parameters, such as touchdown angle, step length and step frequency, comparable to the experimental observations. The good matches between the model and experimental GRF data imply that the continuous pivot acceleration associated with CoM oscillatory behavior could serve as a useful framework of bipedal model.     

Forward dynamic simulation of bipedal walking in the Japanese macaque: investigation of causal relationships among limb kinematics, speed, and energetics of bipedal locomotion in a nonhuman primate

Japanese macaques that have been trained for monkey performances exhibit a remarkable ability to walk bipedally. In this study, we dynamically reconstructed bipedal walking of the Japanese macaque to investigate causal relationships among limb kinematics, speed, and energetics, with a view to understanding the mechanisms underlying the evolution of human bipedalism. We constructed a two-dimensional macaque musculoskeletal model consisting of nine rigid links and eight principal muscles. To generate locomotion, we used a trajectory-tracking control law, the reference trajectories of which were obtained experimentally. Using this framework, we evaluated the effects of changes in cycle duration and gait kinematics on locomotor efficiency. The energetic cost of locomotion was estimated based on the calculation of mechanical energy generated by muscles. Our results demonstrated that the mass-specific metabolic cost of transport decreased as speed increased in bipedal walking of the Japanese macaque. Furthermore, the cost of transport in bipedal walking was reduced when vertical displacement of the hip joint was virtually modified in the simulation to be more humanlike. Human vertical fluctuations in the body's center of mass actually contributed to energy savings via an inverted pendulum mechanism.     

A bipedal compliant walking model generates periodic gait cycles with realistic swing dynamics

A simple spring mechanics model can capture the dynamics of the center of mass (CoM) during human walking, which is coordinated by multiple joints. This simple spring model, however, only describes the CoM during the stance phase, and the mechanics involved in the bipedality of the human gait are limited. In this study, a bipedal spring walking model was proposed to demonstrate the dynamics of bipedal walking, including swing dynamics followed by the step-to-step transition. The model consists of two springs with different stiffnesses and rest lengths representing the stance leg and swing leg. One end of each spring has a foot mass, and the other end is attached to the body mass. To induce a forward swing that matches the gait phase, a torsional hip joint spring was introduced at each leg. To reflect the active knee flexion for foot clearance, the rest length of the swing leg was set shorter than that of the stance leg, generating a discrete elastic restoring force. The number of model parameters was reduced by introducing dependencies among stiffness parameters. The proposed model generates periodic gaits with dynamics-driven step-to-step transitions and realistic swing dynamics. While preserving the mimicry of the CoM and ground reaction force (GRF) data at various gait speeds, the proposed model emulated the kinematics of the swing leg. This result implies that the dynamics of human walking generated by the actuations of multiple body segments is describable by a simple spring mechanics.     

Bioenergetics and the origin of hominid bipedalism

Compared to most quadrupedal mammals, humans are energetically inefficient when running at high speeds. This fact can be taken to mean that human bipedalism evolved for reasons other than to reduce relative energy cost during locomotion. Recalculation of the energy expended during human walking at normal speeds shows that (1) human bipedalism is at least as efficient as typical mammalian quadrupedalism and (2) human gait is much more efficient than bipedal or quadrupedal locomotion in the chimpanzee. We conclude that bipedalism bestowed an energetic advantage on the Miocene hominoid ancestors of the Hominidae.     

Resonance-based oscillations could describe human gait mechanics under various loading conditions

The oscillatory behavior of the center of mass (CoM) and the corresponding ground reaction force (GRF) of human gait for various gait speeds can be accurately described in terms of resonance using a spring–mass bipedal model. Resonance is a mechanical phenomenon that reflects the maximum responsiveness and energetic efficiency of a system. To use resonance to describe human gait, we need to investigate whether resonant mechanics is a common property under multiple walking conditions. Body mass and leg stiffness are determinants of resonance; thus, in this study, we investigated the following questions: (1) whether the estimated leg stiffness increased with inertia, (2) whether a resonance-based CoM oscillation could be sustained during a change in the stiffness, and (3) whether these relationships were consistently observed for different walking speeds. Seven healthy young subjects participated in over-ground walking trials at three different gait speeds with and without a 25-kg backpack. We measured the GRFs and the joint kinematics using three force platforms and a motion capture system. The leg stiffness was incorporated using a stiffness parameter in a compliant bipedal model that best fitted the empirical GRF data. The results showed that the leg stiffness increased with the load such that the resonance-based oscillatory behavior of the CoM was maintained for a given gait speed. The results imply that the resonance-based oscillation of the CoM is a consistent gait property and that resonant mechanics may be useful for modeling human gait.     

Chimpanzee bipedal locomotion in the Gombe National Park,East Africa

An adult male chimpanzee in the natural habitat has been observed to walk predominantly bipedally after a total forelimb paralysis in 1966. The major differences from previously described bipedal chimpanzee gait are (1) one third of the femoral extension is posterior to the hip joint in propulsion, (2) excursion of the swinging foot is close to midline, due to adduction of the lower hindlimb in swing and propulsive phases, (3) depressed pelvic tilt is on the side of the swinging limb, (4) thoracic vertebrae rotate and are vertical and erect, and (5) there is only a moderate lateral sway of the midline. This locomotory complex is interpreted as individual variability and suggests an evolutionary model for the origin of hominid bipedal locomotion.     

Kinematics of bipedalism in Propithecus verreauxi

Bipedalism is rare in primates and has evolved in two distantly related groups: hominoids and indrids. Although copious data are available on the mechanics of bipedal locomotion in hominoids and vertical clinging and leaping (VCL) in indrids, no research has addressed the unique mode of bipedal locomotion exhibited by select indrid primates. Propithecus verreauxi is a highly specialized indrid vertical clinger and leaper that uses a peculiar form of bipedalism on the ground. The objectives of this study were to describe the bipedal gait of Propithecus , to assess the influence of VCL specializations on the kinematic patterns and propulsion mechanisms used by Propithecus during bipedalism, and to compare Propithecus bipedalism with the bipedal gaits of other primates capable of using bipedalism. Video was collected of five adult P. verreauxi moving bipedally in a seminatural setting at the Duke University Primate Center. Duty factor, footfall patterns, joint angles and center of mass movement were quantified in the sagittal plane for 73 steps. Propithecus uses a bipedal gallop, a gait unique to Propithecus . The kinematic similarities (e.g. large hip and knee angular excursions and preparatory countermovements) between bipedal galloping and VCL lead us to suggest that Propithecus takes advantage of specializations for VCL to conserve energy during bipedal galloping. Propithecus also walks bipedally at slower speeds. When Propithecus walks, it utilizes a relatively compliant gait similar to that of other primate facultative bipeds ( Pan , Hylobates ). During bipedal walking, energy conservation may be sacrificed for increased balance and reduced joint loads.     

Do highly trained monkeys walk like humans? A kinematic study of bipedal locomotion in bipedally trained Japanese macaques

In this study, we examined the kinematics of bipedal walking in macaque monkeys that have been highly trained to stand and walk bipedally, and compared them to the kinematics of bipedal walking in ordinary macaques. The results revealed that the trained macaques walked with longer and less frequent strides than ordinary subjects. In addition, they appear to have used inverted pendulum mechanics during bipedal walking, which resulted in an efficient exchange of potential and kinetic energy. These gait characteristics resulted from the relatively more extended hindlimb joints of the trained macaques. By contrast, the body of the ordinary macaques translated downward during the single-limb stance phase due to more flexed hindlimb joints. This resulted in almost in-phase fluctuations of potential and kinetic energy, which indicated that energy transformation was less efficient in the ordinary macaques. The findings provide two insights into the early stage of the evolution of human bipedalism. First, the finding that training considerably improved bipedal walking a posteriori may explain why the very first bipeds that might not yet have been morphologically adapted to bipedal walking continued to walk bipedally. The evolutionary transition from quadrupedalism to bipedalism might not be as difficult as has been envisioned. In addition, the finding that macaques, which are phylogenetically distant from humans and in which bipedal walking is unlike human walking, could develop humanlike gait characteristics with training, provides strong support for the commonly held but unproven idea that the characteristics of the human gait are advantageous to human bipedalism.     

Quantitative evaluation of the major determinants of human gait

Accurate knowledge of the isolated contributions of joint movements to the three-dimensional displacement of the center of mass (COM) is fundamental for understanding the kinematics of normal walking and for improving the treatment of gait disabilities. Saunders et al. (1953) identified six kinematic mechanisms to explain the efficient progression of the whole-body COM in the sagittal, transverse, and coronal planes. These mechanisms, referred to as the major determinants of gait, were pelvic rotation, pelvic list, stance knee flexion, foot and knee mechanisms, and hip adduction. The aim of the present study was to quantitatively assess the contribution of each major gait determinant to the anteroposterior, vertical, and mediolateral displacements of the COM over one gait cycle. The contribution of each gait determinant was found by applying the concept of an ‘influence coefficient’, wherein the partial derivative of the COM displacement with respect to a prescribed determinant was calculated. The analysis was based on three-dimensional measurements of joint angular displacements obtained from 23 healthy young adults walking at slow, normal and fast speeds. We found that hip flexion, stance knee flexion, and ankle-foot interaction (comprised of ankle plantarflexion, toe flexion and the displacement of the center of pressure) are the major determinants of the displacements of the COM in the sagittal plane, while hip adduction and pelvic list contribute most significantly to the mediolateral displacement of the COM in the coronal plane. Pelvic rotation and pelvic list contribute little to the vertical displacement of the COM at all walking speeds. Pelvic tilt, hip rotation, subtalar inversion, and back extension, abduction and rotation make negligible contributions to the displacements of the COM in all three anatomical planes.     

Form in the context of function: Fundamentals of an energy effective striding walk,the role of the plantigrade foot and its expected size

What morphological and functional factors allow for the unique and characteristic upright striding walk of the hominin lineage? Predictive models of locomotion that arise from considering mechanisms of energy loss indicate that collision-like losses at the transition between stance limbs are important determinants of bipedal gait. Theoretical predictions argue that these collisional losses can be reduced by having “functional extra legs” which are physically the heel and the toe part of a single anatomical foot. The ideal spacing for these “functional legs” are up to a quarter of a stride length, depending on the model employed. We evaluate the foot in the context of the dynamics of a bipedal system and compare predictions of optimal foot size against empirical data from modern humans, the Laetoli footprint trackways, and chimpanzees walking bipedally. The dynamics-based modeling approach provides substantial insight into how, and why, walking works as it does, even though current models are too simple to make predictions at a level adequate to anticipate specific morphology except at the most general level.     

Minimizing center of mass vertical movement increases metabolic cost in walking.

A human walker vaults up and over each stance limb like an inverted pendulum. This similarity suggests that the vertical motion of a walker's center of mass reduces metabolic cost by providing a mechanism for pendulum-like mechanical energy exchange. Alternatively, some researchers have hypothesized that minimizing vertical movements of the center of mass during walking minimizes the metabolic cost, and this view remains prevalent in clinical gait analysis. We examined the relationship between vertical movement and metabolic cost by having human subjects walk normally and with minimal center of mass vertical movement ("flat-trajectory walking"). In flat-trajectory walking, subjects reduced center of mass vertical displacement by an average of 69% (P = 0.0001) but consumed approximately twice as much metabolic energy over a range of speeds (0.7-1.8 m/s) (P = 0.0001). In flat-trajectory walking, passive pendulum-like mechanical energy exchange provided only a small portion of the energy required to accelerate the center of mass because gravitational potential energy fluctuated minimally. Thus, despite the smaller vertical movements in flat-trajectory walking, the net external mechanical work needed to move the center of mass was similar in both types of walking (P = 0.73). Subjects walked with more flexed stance limbs in flat-trajectory walking (P < 0.001), and the resultant increase in stance limb force generation likely helped cause the doubling in metabolic cost compared with normal walking. Regardless of the cause, these findings clearly demonstrate that human walkers consume substantially more metabolic energy when they minimize vertical motion.     

Human Locomotion

The development of bipedal plantigrade progression is a purely human, and apparently learned, accomplishment. Experimental findings confirm the hypothesis that the human body will integrate the motion of various segments of the body and control the activity of muscles to minimize energy expenditure.Movements which are integrated for this purpose include vertical displacement of the body, horizontal rotation of the pelvis, mediolateral pelvic tilt, flexion of the knee, plantar flexion of the ankle and foot, lateral displacement of the torso and rotation of the shoulder girdle.Raising and lowering the body results in gains and losses of potential energy, and acceleration and deceleration result in gains and losses of kinetic energy. The motions are so co-ordinated that a transfer of energy back and forth from kinetic to potential occurs during walking, which tends to minimize total energy expenditure as well as muscle work.     

Voluntary bipedal walking of infant chimpanzees

The voluntary bipedal walking of infant chimpanzees was studied by the analysis of foot force and by motion analysis. The infants were trained to locomote on a level platform without any restrictions on the locomotor pattern. The voluntary bipedal walking was compared with the other types of locomotion at the same age and with the trained bipedal walking performed by other chimpanzees, including adult chimpanzees. The characteristics of voluntary bipedal walking in the infant until one year of age were: (1) high-speed walking with short cycle duration; (2) short stance phase duration; (3) small braking component of the preceding leg and large acceleration of the following leg; (4) one downward peak in the vertical component; and (5) a relatively small transverse component. Bipedal walking usually continued for less than one second and ended in quadrupedal locomotion. During walking, the preceding foot touched the floor, heel first, as in the case of older chimpanzees and humans. At this age, bipedal walking was similar to high-speed locomotion. The voluntary bipedal walking of the two-year-old and frour-yearold chimpanzees was characterized as follows: (1) slower speed than during quadrupedal locomotion, (2) relatively long periods and distances; (3) well balanced accelerating and braking components; and (4) a vertical component showing two downward peaks and a trough in between during numerous trials. The last characteristic means that the body center of gravity is higher in the single stance phase, just as in the bipedal walkinbg of the adult chimpanzees and humans. The bipedal walking of infant chimpanzees was discussed in comparison with the walking of humans, including infants.     

Control of lateral weight transfer is associated with walking speed in individuals post-stroke

Restoring functional gait speed is an important goal for rehabilitation post-stroke. During walking, transferring of one’s body weight between the limbs and maintaining balance stability are necessary for independent functional gait. Although it is documented that individuals post-stroke commonly have difficulties with performing weight transfer onto their paretic limbs, it remains to be determined if these deficits contributed to slower walking speeds. The primary purpose of this study was to compare the weight transfer characteristics between slow and fast post-stroke ambulators. Participants (N = 36) with chronic post-stroke hemiparesis walked at their comfortable and maximal walking speeds on a treadmill. Participants were stratified into 2 groups based on their comfortable walking speeds (≥0.8 m/s or <0.8 m/s). Minimum body center of mass (COM) to center of pressure (COP) distance, weight transfer timing, step width, lateral foot placement relative to the COM, hip moment, peak vertical and anterior ground reaction forces, and changes in walking speed were analyzed. Results showed that slow walkers walked with a delayed and deficient weight transfer to the paretic limb, lower hip abductor moment, and more lateral paretic limb foot placement relative to the COM compared to fast walkers. In addition, propulsive force and walking speed capacity was related to lateral weight transfer ability. These findings demonstrated that deficits in lateral weight transfer and stability could potentially be one of the limiting factors underlying comfortable walking speeds and a determinant of chronic stroke survivors’ ability to increase walking speed.     

Maximum walking speed and lower limb length in hominids

In 1984, Helene (Am. J. Physics 52:656) and Alexander (Am. Scientist 72:348–354) presented equations which purported to explain how lower limb length limited maximum walking speed in humans. The equations were based on a simplified model of human walking in which the center of mass (CoM) “vaults” over the supporting leg. Increasing walking speed by increasing stride frequency or stride length would increase the upward acceleration of the CoM in the first half of stance phase, to the point that it would be greater than the downward pull of gravity, and the individual would become airborne. This constitutes running by most definitions. While these models ignored various mechanical factors, such as knee flexion during midstance, that reduce the vertical movement of the CoM, the general idea is plausible inasmuch as the CoM of the body does oscillate vertically with each step. One hypothesis tested here is whether it is indeed the interaction between the pull of gravity and the individual's own upward acceleration that determines at what speed (or cadence) he changes from walking to running. Another hypothesis considered is that increased lower limb length (L) was selected for in early hominids, because of the locomotor advantages of longer lower limbs. Results indicate, however, that while L was clearly related to maximum possible walking speed, it was not an important factor in determining maximum “comfortable” walking speed. These and other results from the recent literature suggest that increased lower limb length provided no selective advantage in locomotion, and other explanations should be sought. © 1996 Wiley-Liss, Inc.     

The gait of children with and without cerebral palsy: work, energy, and angular momentum

This paper describes a method to characterize gait pathologies like cerebral palsy using work, energy, and angular momentum. For a group of 24 children, 16 with spastic diplegic cerebral palsy and 8 typically developed, kinematic data were collected at the subjects self selected comfortable walking speed. From the kinematics, the work-internal, external, and whole body; energy-rotational and relative linear; and the angular momentum were calculated. Our findings suggest that internal work represents 53% and 40% respectively of the whole body work in gait for typically developed children and children with cerebral palsy. Analysis of the angular momentum of the whole body, and other subgroupings of body segments, revealed a relationship between increased angular momentum and increased internal work. This relationship allows one to use angular momentum to assist in determining the kinetics and kinematics of gait which contribute to increased internal work. Thus offering insight to interventions which can be applied to increase the efficiency of bipedal locomotion, by reducing internal work which has no direct contribution to center of mass motion, in both normal and pathologic populations.