A simple model of bipedal walking predicts the preferred speed-step length relationship

We used a simple model of passive dynamic walking, with the addition of active powering on level ground, to study the preferred relationship between speed and step length in humans. We tested several hypothetical metabolic costs, with one component proportional to the mechanical work associated with pushing off with the stance leg at toe-off, and another component associated with several possible costs of forcing oscillations of the swing leg. For this second component, a cost based on the amount of force needed to oscillate the leg divided by the time duration of that force predicts the preferred speed-step length relationship much better than other costs, such as the amount of mechanical work done in swinging the leg. The cost of force/time models the need to recruit fast muscle fibers for large forces at short durations. The actual mechanical work performed by muscles on the swing leg appears to be of relatively less importance, although it appears to be minimized by the use of short bursts of muscle activity in near-isometric conditions. The combined minimization of toe-off mechanical work and force divided by time predicts the preferred speed-step length relationship.     

Limb work and joint work minimization reveal an energetic benefit to the elbows-back,knees-forward limb design in parasagittal quadrupeds

Quadrupedal animal locomotion is energetically costly. We explore two forms of mechanical work that may be relevant in imposing these physiological demands. Limb work, due to the forces and velocities between the stance foot and the centre of mass, could theoretically be zero given vertical limb forces and horizontal centre of mass path. To prevent pitching, skewed vertical force profiles would then be required, with forelimb forces high in late stance and hindlimb forces high in early stance. By contrast, joint work—the positive mechanical work performed by the limb joints—would be reduced with forces directed through the hip or shoulder joints. Measured quadruped kinetics show features consistent with compromised reduction of both forms of work, suggesting some degree of, but not perfect, inter-joint energy transfer. The elbows-back, knees-forward design reduces the joint work demand of a low limb-work, skewed, vertical force profile. This geometry allows periods of high force to be supported when the distal segment is near vertical, imposing low moments about the elbow or knee, while the shoulder or hip avoids high joint power despite high moments because the proximal segment barely rotates—translation over this period is due to rotation of the distal segment.     

Consequences of forward translation of the point of force application for the mechanics of running

In running humans, the point of force application between the foot and the ground moves forwards during the stance phase. Our aim was to determine the mechanical consequences of this 'point of force translation' (POFT). We modified the planar spring-mass model of locomotion to incorporate POFT, and then compared spring-mass simulations with and without POFT. We found that, if leg stiffness is adjusted appropriately, it is possible to maintain very similar values of peak vertical ground reaction force (GRF), stance time, contact length and vertical centre of mass displacement, whether or not POFT occurs. The leg stiffness required to achieve this increased as the distance of POFT increased. Peak horizontal GRF and mechanical work per step were lower when POFT occurred. The results indicate that the lack of POFT in the traditional spring-mass model should not prevent it from providing good predictions of peak vertical GRF, stance time, contact length and vertical centre of mass displacement in running humans, if an appropriate spring stiffness is used. However, the model can be expected to overestimate peak horizontal GRF and mechanical work per step. When POFT occurs, the spring stiffness in the traditional spring-mass model is not equivalent to leg stiffness. Therefore, caution should be exercised when using spring stiffness to understand how the musculoskeletal system adapts to different running conditions. This can explain the contradictory results in the literature regarding the effect of running speed on leg stiffness.     

Why not walk faster?

Bipedal walking following inverted pendulum mechanics is constrained by two requirements: sufficient kinetic energy for the vault over midstance and sufficient gravity to provide the centripetal acceleration required for the arc of the body about the stance foot. While the acceleration condition identifies a maximum walking speed at a Froude number of 1, empirical observation indicates favoured walk-run transition speeds at a Froude number around 0.5 for birds, humans and humans under manipulated gravity conditions. In this study, I demonstrate that the risk of 'take-off' is greatest at the extremes of stance. This is because before and after kinetic energy is converted to potential, velocities (and so required centripetal accelerations) are highest, while concurrently the component of gravity acting in line with the leg is least. Limitations to the range of walking velocity and stride angle are explored. At walking speeds approaching a Froude number of 1, take-off is only avoidable with very small steps. With realistic limitations on swing-leg frequency, a novel explanation for the walk-run transition at a Froude number of 0.5 is shown.     

Gait transitions in simulated reduced gravity

Gravity has a strong effect on gait and the speed of gait transitions. A gait has been defined as a pattern of locomotion that changes discontinuously at the transition to another gait. On Earth, during gradual speed changes, humans exhibit a sudden discontinuous switch from walking to running at a specific speed. To study the effects of altered gravity on both the stance and swing legs, we developed a novel unloading exoskeleton that allows a person to step in simulated reduced gravity by tilting the body relative to the vertical. Using different simulation techniques, we confirmed that at lower gravity levels the transition speed is slower (in accordance with the previously reported Froude number ～0.5). Surprisingly, however, we found that at lower levels of simulated gravity the transition between walking and running was generally gradual, without any noticeable abrupt change in gait parameters. This was associated with a significant prolongation of the swing phase, whose duration became virtually equal to that of stance in the vicinity of the walk-run transition speed, and with a gradual shift from inverted-pendulum gait (walking) to bouncing gait (running).     

Correlation of muscle function and bone strain in the hindlimb of the river cooter turtle (Pseudemys concinna)

During terrestrial locomotion, limb muscles must generate mechanical work and stabilize joints against the ground reaction force. These demands can require high force production that imposes substantial loads on limb bones. To better understand how muscle contractile function influences patterns of bone loading in terrestrial locomotion, and refine force platform equilibrium models used to estimate limb bone safety factors, we correlated in vivo recordings of femoral strain with muscle activation and strain in a major propulsive hindlimb muscle, flexor tibialis internus (FTI), of a species with a published model of hindlimb force production (river cooter turtles, Pseudemys concinna). Electromyography (EMG) recordings indicate FTI activity prior to footfall that continues through approximately 50% of the stance phase. Large EMG bursts occur just after footfall when the muscle has reached its maximum length and is beginning to actively shorten, concurrent with increasing compressive strain on the anterior femur. The FTI muscle shortens through 35% of stance, with mean fascicle shortening strains reaching 14.0 ± 5.4% resting length (L₀). At the time of peak compressive strains on the femur, the muscle fascicles remain active, but fascicles typically lengthen until mid‐stance as the knee extends. Influenced by the activity of the dorsal knee extensor femorotibialis, the FTI muscle continues to passively lengthen simultaneously with knee extension and a shift to tensile axial strain on the anterior femur at approximately 40% of stance. The near coincidence in timing of peak compressive bone strain and peak muscle shortening (5.4 ± 4.1% stance) indicates a close correlation between the action of the hip extensor/knee flexor, FTI, and femoral loading in the cooter hindlimb. In the context of equilibrium models of limb bone loading, these results may help explain differences in safety factor estimates observed between previous force platform and in vivo strain analyses in cooters. J. Morphol. 274:1060–1069, 2013. © 2013 Wiley Periodicals, Inc.     

External forces and torques generated by the brachiating white-handed gibbon (Hylobates lar)

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.     

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.     

Adaptive changes in running kinematics as a function of head stability demands and their effect on shock transmission

This study aimed to identify adaptive changes in running kinematics and impact shock transmission as a function of head stability requirements. Fifteen strides from twelve recreational runners were collected during preferred speed treadmill running. Head stability demands were manipulated through real-time visual feedback that required head-gaze orientation to maintain within boxes of different sizes, ranging from 21° to 3° of visual angle with 3° decrements. The main outcome measures were tibial and head peak accelerations in the time and frequency domains (impact and active phases), shock transmission from tibia to head, stride parameters, and sagittal plane joint kinematics. Increasing head stability requirements resulted in decreases in the amplitude and integrated power of head acceleration during the active phase of stance. During the impact portion of stance tibial and head acceleration and shock transmission remained similar across visual conditions. In response to increased head stability requirements, participants increased stride frequency approximately 8% above preferred, as well as hip flexion angle at impact; stance time and knee and ankle joint angles at impact did not change. Changes in lower limb joint configurations (smaller hip extension and ankle plantar-flexion and greater knee flexion) occurred at toe-off and likely contributed to reducing the vertical displacement of the center of mass with increased head stability demands. These adaptive changes in the lower limb enabled runners to increase the time that voluntary control is allowed without embedding additional impact loadings, and therefore active control of the head orientation was facilitated in response to different visual task constraints.     

Asymmetric interlimb role-sharing in mechanical power during human sideways locomotion

Sideways movement at a wide variety of speeds is required in daily life and sports. The purpose of this study was to identify the characteristics of asymmetry in power output between lower limbs during sideways gait patterns. Seven healthy men performed steady-state sideways locomotion at various speeds. The mechanical external power of each limb was calculated and decomposed to the lateral and vertical components by the center of mass velocity and ground reaction force. We acquired data from 126 steps of sideways walking at 0.44–1.21 m/s, and from 41 steps of sideways galloping at 1.04–3.00 m/s. The results showed asymmetric power production between the limbs during sideways locomotion. During sideways walking, the trailing limb predominantly produced positive external power and the leading limb produced predominantly negative external power, and these amplitudes increased with step speed. In contrast, during sideways galloping, negative and subsequent positive power production was observed in both limbs. These differences in asymmetric interlimb role-sharing were mainly due to the vertical component. During sideways galloping, the trailing limb absorbs vertical power produced by the leading limb due to the longer flight time. This characteristic of vertical power production in the trailing limb may explain the presence of a double-support phase, which is not observed during forward running, even at high speeds. Our results will help to elucidate the asymmetric movements of the limbs in lateral directions at various speeds.     

Muscle mechanical advantage of human walking and running: implications for energy cost.

Muscular forces generated during locomotion depend on an animal's speed, gait, and size and underlie the energy demand to power locomotion. Changes in limb posture affect muscle forces by altering the mechanical advantage of the ground reaction force (R) and therefore the effective mechanical advantage (EMA = r/R, where r is the muscle mechanical advantage) for muscle force production. We used inverse dynamics based on force plate and kinematic recordings of humans as they walked and ran at steady speeds to examine how changes in muscle EMA affect muscle force-generating requirements at these gaits. We found a 68% decrease in knee extensor EMA when humans changed gait from a walk to a run compared with an 18% increase in hip extensor EMA and a 23% increase in ankle extensor EMA. Whereas the knee joint was extended (154-176 degrees) during much of the support phase of walking, its flexed position (134-164 degrees) during running resulted in a 5.2-fold increase in quadriceps impulse (time-integrated force during stance) needed to support body weight on the ground. This increase was associated with a 4.9-fold increase in the ground reaction force moment about the knee. In contrast, extensor impulse decreased 37% (P < 0.05) at the hip and did not change at the ankle when subjects switched from a walk to a run. We conclude that the decrease in limb mechanical advantage (mean limb extensor EMA) and increase in knee extensor impulse during running likely contribute to the higher metabolic cost of transport in running than in walking. The low mechanical advantage in running humans may also explain previous observations of a greater metabolic cost of transport for running humans compared with trotting and galloping quadrupeds of similar size.     

Ability of the planar spring-mass model to predict mechanical parameters in running humans

The planar spring-mass model is a simple mathematical model of bouncing gaits, such as running, trotting and hopping. Although this model has been widely used in the study of locomotion, its accuracy in predicting locomotor mechanics has not been systematically quantified. We determined the percent error of the model in predicting 10 locomotor parameters in running humans by comparing the model predictions to experimental data from humans running in normal gravity and simulated reduced gravity. We tested the hypotheses that the model would overestimate horizontal impulse and the change in mechanical energy of the centre of mass (COM) during stance. The model provided good predictions of stance time, vertical impulse, contact length, duty factor, relative stride length and relative peak force. All predictions of these parameters were within 20% of measured values and at least 90% of predictions of each parameter were within 10% of measured values (median absolute errors: <7%). This suggests that the model incorporates all features of running humans that have a significant influence upon these six parameters. As simulated gravity level decreased, the magnitude of the errors in predicting each of these parameters either decreased or stayed constant, indicating that this is a good model of running in simulated reduced gravity. As hypothesised, horizontal impulse and change in mechanical energy of the COM during stance were overestimated (median absolute errors: 43.6% and 26.2%, respectively). Aerial time and peak vertical COM displacement during stance were also systematically overestimated (median absolute errors: 17.7% and 22.9%, respectively). Care should be taken to ensure that the model is used only to investigate parameters which it can predict accurately. It would be useful to extend this analysis to other species and gaits.     

Mechanical and metabolic determinants of the preferred step width in human walking

We studied the selection of preferred step width in human walking by measuring mechanical and metabolic costs as a function of experimentally manipulated step width (0.00-0.45L, as a fraction of leg length L). We estimated mechanical costs from individual limb external mechanical work and metabolic costs using open circuit respirometry. The mechanical and metabolic costs both increased substantially (54 and 45%, respectively) for widths greater than the preferred value (0.15-0.45L) and with step width squared (R(2) = 0.91 and 0.83, respectively). As predicted by a three-dimensional model of walking mechanics, the increases in these costs appear to be a result of the mechanical work required for redirecting the centre of mass velocity during the transition between single stance phases (step-to-step transition costs). The metabolic cost for steps narrower than preferred (0.10-0.00L) increased by 8%, which was probably as a result of the added cost of moving the swing leg laterally in order to avoid the stance leg (lateral limb swing cost). Trade-offs between the step-to-step transition and lateral limb swing costs resulted in a minimum metabolic cost at a step width of 0.12L, which is not significantly different from foot width (0.11L) or the preferred step width (0.13L). Humans appear to prefer a step width that minimizes metabolic cost.     

Grip and limb force limits to turning performance in competition horses

Manoeuverability is a key requirement for successful terrestrial locomotion, especially on variable terrain, and is a deciding factor in predator-prey interaction. Compared with straight-line running, bend running requires additional leg force to generate centripetal acceleration. In humans, this results in a reduction in maximum speed during bend running and a published model assuming maximum limb force as a constraint accurately predicts how much a sprinter must slow down on a bend given his maximum straight-line speed. In contrast, greyhounds do not slow down or change stride parameters during bend running, which suggests that their limbs can apply the additional force for this manoeuvre. We collected horizontal speed and angular velocity of heading of horses while they turned in different scenarios during competitive polo and horse racing. The data were used to evaluate the limits of turning performance. During high-speed turns of large radius horizontal speed was lower on the bend, as would be predicted from a model assuming a limb force limit to running speed. During small radius turns the angular velocity of heading decreased with increasing speed in a manner consistent with the coefficient of friction of the hoof-surface interaction setting the limit to centripetal force to avoid slipping.     

The relationship between joint kinetic factors and the walk-run gait transition speed during human locomotion

The primary purpose of this project was to examine whether lower extremity joint kinetic factors are related to the walk-run gait transition during human locomotion. Following determination of the preferred transition speed (PTS), each of the 16 subjects walked down a 25-m runway, and over a floor-mounted force platform at five speeds (70, 80, 90, 100, and 110% of the PTS), and ran over the force platform at three speeds (80, 100, and 120% of the PTS) while being videotaped (240 Hz) from the right sagittal plane. Two-dimensional kinematic data were synchronized with ground reaction force data (960 Hz). After smoothing, ankle and knee joint moments and powers were calculated using standard inverse dynamics calculations. The maximum dorsiflexor moment was the only variable tested that increased as walking speed increased and then decreased when gait changed to a run at the PTS, meeting the criteria set to indicate that this variable influences the walk-run gait transition during human locomotion. This supports previous research suggesting that an important factor in changing gaits at the PTS is the prevention of undue stress in the dorsiflexor muscles.     

Effect of surgery to implant motion and force sensors on vertical ground reaction forces in the ovine model

Activities of daily living (ADLs) generate complex, multidirectional forces in the anterior cruciate ligament (ACL). While calibration problems preclude direct measurement in patients, ACL forces can conceivably be measured in animals after technical challenges are overcome. For example, motion and force sensors can be implanted in the animal but investigators must determine the extent to which these sensors and surgery affect normal gait. Our objectives in this study were to determine (1) if surgically implanting knee motion sensors and an ACL force sensor significantly alter normal ovine gait and (2) how increasing gait speed and grade on a treadmill affect ovine gait before and after surgery. Ten skeletally mature, female sheep were used to test four hypotheses: (1) surgical implantation of sensors would significantly decrease average and peak vertical ground reaction forces (VGRFs) in the operated limb, (2) surgical implantation would significantly decrease single limb stance duration for the operated limb, (3) increasing treadmill speed would increase VGRFs pre- and post operatively, and (4) increasing treadmill grade would increase the hind limb VGRFs pre- and post operatively. An instrumented treadmill with two force plates was used to record fore and hind limb VGRFs during four combinations of two speeds (1.0 m/s and 1.3 m/s) and two grades (0 deg and 6 deg). Sensor implantation decreased average and peak VGRFs less than 10% and 20%, respectively, across all combinations of speed and grade. Sensor implantation significantly decreased the single limb stance duration in the operated hind limb during inclined walking at 1.3 m/s but had no effect on single limb stance duration in the operated limb during other activities. Increasing treadmill speed increased hind limb peak (but not average) VGRFs before surgery and peak VGRF only in the unoperated hind limb during level walking after surgery. Increasing treadmill grade (at 1 m/s) significantly increased hind limb average and peak VGRFs before surgery but increasing treadmill grade post op did not significantly affect any response measure. Since VGRF values exceeded 80% of presurgery levels, we conclude that animal gait post op is near normal. Thus, we can assume normal gait when conducting experiments following sensor implantation. Ultimately, we seek to measure ACL forces for ADLs to provide design criteria and evaluation benchmarks for traditional and tissue engineered ACL repairs and reconstructions.     

Changes in relative work of the lower extremity joints and distal foot with walking speed

The modulation of walking speed results in adaptations to the lower limbs which can be quantified using mechanical work. A 6 degree-of-freedom (DOF) power analysis, which includes additional translations as compared to the 3 DOF (all rotational) approach, is a comprehensive approach for quantifying lower limb work during gait. The purpose of this study was to quantify the speed-related 6 DOF joint and distal foot work adaptations of all the lower extremity limb constituents (hip, knee, ankle, and distal foot) in healthy individuals. Relative constituent 6 DOF work, the amount of constituent work relative to absolute limb work, was calculated during the stance and swing phases of gait. Eight unimpaired adults walked on an instrumented split-belt treadmill at slow, moderate, and typical walking speeds (0.4, 0.6, and 0.8 statures/s, respectively). Using motion capture and force data, 6 DOF powers were calculated for each constituent. Contrary to previously published results, 6 DOF positive relative ankle work and negative relative distal foot work increased significantly with increased speed during stance phase (p < 0.05). Similar to previous rotational DOF results in the sagittal plane, negative relative ankle work decreased significantly with increased speed during stance phase (p < 0.05). Scientifically, these findings provide new insight into how healthy individuals adapt to increased walking speed and suggest limitations of the rotational DOF approach for quantifying limb work. Clinically, the data presented here for unimpaired limbs can be used to compare with speed-matched data from limbs with impairments.     

The Cost of Leg Forces in Bipedal Locomotion: A Simple Optimization Study

Simple optimization models show that bipedal locomotion may largely be governed by the mechanical work performed by the legs, minimization of which can automatically discover walking and running gaits. Work minimization can reproduce broad aspects of human ground reaction forces, such as a double-peaked profile for walking and a single peak for running, but the predicted peaks are unrealistically high and impulsive compared to the much smoother forces produced by humans. The smoothness might be explained better by a cost for the force rather than work produced by the legs, but it is unclear what features of force might be most relevant. We therefore tested a generalized force cost that can penalize force amplitude or its n-th time derivative, raised to the p-th power (or p-norm), across a variety of combinations for n and p. A simple model shows that this generalized force cost only produces smoother, human-like forces if it penalizes the rate rather than amplitude of force production, and only in combination with a work cost. Such a combined objective reproduces the characteristic profiles of human walking (R² = 0.96) and running (R² = 0.92), more so than minimization of either work or force amplitude alone (R² = −0.79 and R² = 0.22, respectively, for walking). Humans might find it preferable to avoid rapid force production, which may be mechanically and physiologically costly.     

Mechanical output of the cat soleus during treadmill locomotion: in vivo vs in situ characteristics

To study the mechanical output of skeletal muscle, four adult cats were trained to run on a treadmill and then implanted under sterile conditions and anesthesia with a force transducer on the soleus tendon and EMG electrodes in the muscle belly. After a two-week recovery period, five consecutive step cycles were filmed at treadmill speeds of 0.8, 1.3 and 2.2 m s-1. Locomotion data in vivo included individual muscle force, length and velocity changes and EMG during each step cycle. Data for an average step cycle at each speed were compared to the force-velocity properties obtained on the same muscle under maximal nerve stimulation and isotonic loading conditions in situ. Results indicate that the force and power generated at a given velocity of shortening during late stance in vivo were greater at the higher speeds of locomotion than the force and power generated at the same shortening velocity in situ. Strain energy stored in the muscle-tendon unit during the yield phase in early stance is felt to be a major contributor to the muscle's enhanced mechanical output during muscle shortening in late stance.     

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.