Kinematic analysis of bipedal locomotion of a Japanese macaque that lost its forearms due to congenital malformation

Spontaneously acquired bipedal locomotion of an untrained Japanese monkey (Macaca fuscata) is measured and compared with the elaborated bipedal locomotion of highly trained monkeys to assess the natural ability of a quadrupedal primate to walk bipedally. The subject acquired bipedalism by himself because of the loss of his forearms and hands due to congenital malformation. Two other subjects are performing monkeys that have been extensively trained for bipedal posture and locomotion. We videotaped their bipedal locomotion with two cameras in a lateral view and calculated joint angles (hip, knee, and ankle) and inertial angle of the trunk from the digitized joint positions. The results show that all joints are relatively more flexed in the untrained monkey. Moreover, it is noted that the ankle is less plantar flexed and the knee is more flexed in mid-to-late stance phase in the untrained monkey, suggesting that the trunk is not lifted up to store potential energy. In the trained monkeys, the joints are extended to bring the trunk as high as possible in the stance phase, and then stored potential energy is exchanged for kinetic energy to move forward. The efficient inverted pendulum mechanism seems to be absent in the untrained monkeys locomotion, implying that acquisition of such efficient bipedal locomotion is not a spontaneous ability for a Japanese monkey. Rather, it is probably a special skill that can only be acquired through artificial training for an inherently quadrupedal primate.This revised version was published online in April 2005 with corrections to the cover date of the issue.     

Dynamic plantar pressure distribution during terrestrial locomotion of bonobos (Pan paniscus)

We collected high-resolution plantar pressure distributions of seven bonobos during terrestrial bipedal and quadrupedal locomotion (N = 146). Functional foot length, degree of hallux abduction, and total contact time were determined, and plots, showing pressure as a function of time for six different foot regions, were generated. We also studied five adult humans for comparison (N = 13). Both locomotion types of the bonobo show a large variation in plantar pressure distributions, which could be due to the interference of instantaneous behavior with locomotion and differences in walking speed and body dimensions. The heel and the lateral midfoot typically touch down simultaneously at initial ground contact in bipedal and quadrupedal walking of bonobos, in contrast with the typical heel-strike of human bipedalism. The center of pressure follows a curved course during quadrupedalism, as a consequence of the medial weight transfer during mid-stance. Bipedal locomotion of bonobos is characterized by a more plantar positioning of the feet and by a shorter contact time than during quadrupedal walking, according to a smaller stride and step length at a higher frequency. We observed a varus position of the foot with an abducted hallux, which likely possesses an important sustaining and stabilizing function during terrestrial locomotion.     

Speed modulation in hylobatid bipedalism: a kinematic analysis

Gibbons are highly arboreal apes, and it is expected that their bipedal locomotion will show some particularities related to the arboreal environment. Previous research has shown that, during hylobatid bipedalism, unsupported phases are rare and stride frequencies are relatively low. This study confirms previous findings, and we suggest that low stride frequencies and the absence of unsupported phases are ways to reduce disadvantageous branch oscillations during arboreal travel. Despite these restrictions, gibbons are able to locomote at a wide range of speeds, implying that they likely exploit other mechanisms to modulate their locomotor speed. To investigate this possibility, we collected video images of a large number of spontaneous bipedal bouts of four untrained white-handed gibbons by using an instrumented walkway with four synchronized cameras. These video images were digitized to obtain a quantification of the 3D kinematics of hylobatid bipedalism. We defined a large number of spatiotemporal and kinematic gait variables, and the relationship between these gait variables and (dimensionless) speed was statistically tested. It was found that gibbons mainly increase stride length to increase their locomotor speed; the main speed-modulating mechanisms are hip and ankle excursion and coupled knee and ankle extension at toe-off. Although aerial phases are rare, gibbons generally adopt a bipedal bouncing gait at most speeds and a clear-cut gait transition, as seen in human locomotion, is absent. Comparison with human and bonobo bipedalism showed that the variability of the 3D joint angles of the hind limb are comparable during human and gibbon bipedalism, and much lower than during bonobo bipedalism. The low variability found in gibbons might be related to constraints imposed by the arboreal environment. These arboreal constraints clearly affect the bipedal gait characteristics of gibbons, but do not constrain the ability to adopt a bipedal bouncing gait during terrestrial locomotion.     

Spatio-temporal gait characteristics of the hind-limb cycles during voluntary bipedal and quadrupedal walking in bonobos (Pan paniscus)

Spatio-temporal gait characteristics (step and stride length, stride frequency, duty factor) were determined for the hind-limb cycles of nine bonobos (Pan paniscus) walking quadrupedally and bipedally at a range of speeds. The data were recalculated to dimensionless quantities according to the principle of dynamic similarity. Lower leg length was used as the reference length. Interindividual variability in speed modulation strategy of bonobos appears to be low. Compared to quadrupedal walking, bipedal bonobos use smaller steps to attain a given speed (differences increase with speed), resulting in shorter strides at a higher frequency. In the context of the ("hybrid") dynamic pattern approach to locomotion (Latach, 1998) we argue that, despite these absolute differences, intended walking speed is the basic control variable which elicits both quadrupedal and bipedal walking kinematics in a similar way. Differences in the initial status of the dynamic system may be responsible for the differences in step length between both gaits. Comparison with data deduced from the literature shows that the effects of walking speed on stride length and frequency are similar in bonobos, common chimpanzees, and humans. This suggests that (at least) within extant homininae, spatio-temporal gait characteristics are highly comparable, and this in spite of obvious differences in mass distribution and bipedal posture.     

Kinematics and ontogeny of locomotion in monkeys and human babies

Early ontogenetic stages are often assumed to reflect or to be similar to past phylogenetic stages within the evolution of man. Therefore, as a first step, the quadrupedal crawling locomotion of human children was analysed and compared to the quadrupedal walk of Macaca fascicularis. The movements of the human child were not only more irregular, they differed from the walk of the monkey mainly through extraordinarily short swing phases, and also through strong scoliotic movements of the spine. There is a compulsory synchronisation in the hip and knee joint movements of the human crawling baby. We conclude that human crawling may be a behavioural recapitulation of a quadrupedal evolutionary stage. However, with reference to kinematics, man is not only characterised by his unique, habitually bipedal, upright gait but also by a second, equally unique locomotion, namely crawling, which he assumes for a short phase during his first year of life.--The walking movements of the limbs in toddling infants were mainly characterised by i) rather stiff, abducted arms, which were moved mostly by spine torsions (similar to those of bipedally walking Gorilla) and not as a suspensory pendulum. However, they rather work as levers for the elastic torsion pendulum of the spine. ii) They are also characterised by frequently lacking the minor knee flexion, which occurs at about the heel strike within each stride of the adult human. Besides many other details of the results, foot movements differed from adult ones mainly in that the whole plantar surface was placed flat on the ground within a few milliseconds.     

Bipedal versus Quadrupedal Hind Limb and Foot Kinematics in a Captive Sample of <Emphasis Type="Italic">Papio anubis:</Emphasis> Setup and Preliminary Results

Setups that integrate both kinematics and morpho-functional investigations of a single sample constitute recent developments in the study of nonhuman primate bipedalisms. We introduce the integrated setup built at the Primatology Station of the French National Centre for Scientific Research (CNRS), which allows analysis of both bipedal and quadrupedal locomotion in a population of 55–60 captive olive baboons. As a first comparison, we present the hind limb kinematics of both locomotor modalities in 10 individuals, focusing on the stance phase. The main results are: 1) differences in bipedal and quadrupedal kinematics at the hip, knee, and foot levels; 2) a variety of foot contacts to the ground, mainly of semiplantigrade type, but also of plantigrade type; 3) equal variations between bipedal and quadrupedal foot angles; 4) the kinematics of the foot joints act in coordinated and stereotyped manners, but are triggered differently according to whether the support is bipedal or quadrupedal. Although very occasionally realized, the bipedal walk of olive baboon appears to be a habitual and nonerratic locomotor modality.     

The impact of adding trunk motion to the interpretation of the role of joint moments during normal walking

Biomechanical model assumptions affect the interpretation of the role of the muscle or joint moments to the segmental power estimated by induced acceleration analysis (IAA). We evaluated the effect of modeling the pelvis and trunk segments as two separate segments (8 SM) versus as a single segment (7 SM) on the segmental power, support of the body, knee and hip extension acceleration produced by the joint moments during the stance phase of normal walking. Significant differences were observed in the contribution of the stance hip abductor and extensor moments to support, ipsilateral knee and hip acceleration, and ipsilateral thigh and upper body power. The primary finding was that the role of the stance hip moment in generating ipsilateral thigh and upper body power differed based on degrees of freedom in the model. Secondarily, the magnitude of contributions also differed. For example, the hip abductor and extensor moments showed greater contribution to support, hip and knee acceleration in the 8 SM. IAA and segment power analysis are sensitive to the degrees of freedom between the pelvis and trunk. There is currently no gold standard by which to evaluate the accuracy of IAA predictions. However, modeling the pelvis and trunk as separate segments is closer to the anatomical architecture of the body. An 8 SM appears to be more appropriate for estimating the role of joint moments, particularly to motion of more proximal segments during normal walking.     

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.     

Ground-reaction-force profiles of bipedal walking in bipedally trained Japanese monkeys

Ground-reaction-force (GRF) profiles of bipedal locomotion in bipedally trained Japanese macaques (performing monkeys) were analyzed in order to clarify the dynamic characteristics of their locomotion. Five trained and two ordinary monkeys participated in the experiment. They walked on a wooden walkway at a self-selected speed, and three components of the GRF vector were measured using a force platform. Our measurements reveal that trained monkeys exhibited vertical-GRF profiles that were single-peaked, similar to those of ordinary monkeys; they did not generate the double-peaked force curve that is seen in humans, despite their extensive training. However, in the trained monkeys, the peak appeared relatively earlier in the stance phase, and overall shape was more triangular than that of the more parabolic profile generated by ordinary monkeys. Comparisons of vertical fluctuation of the center of body mass calculated from the measured profiles suggest that this was larger in the trained monkeys, indicating that storage and release of potential energy actually took place in their bipedal walking. This energetic advantage seems limited, however, because efficient exchange of potential and kinetic energy during walking were not completely out of phase as in human walking. We suggest that anatomically restricted range of hip-joint motion impedes the inherently quadrupedal monkeys from generating humanlike bipedal locomotion, and that morphological rearrangement of the hip joint was an essential precondition for protohominids to acquire humanlike bipedalism.     

Walking with increased ankle pushoff decreases hip muscle moments

In a simple bipedal walking model, an impulsive push along the trailing limb (similar to ankle plantar flexion) or a torque at the hip can power level walking. This suggests a tradeoff between ankle and hip muscle requirements during human gait. People with anterior hip pain may benefit from walking with increased ankle pushoff if it reduces hip muscle forces. The purpose of our study was to determine if simple instructions to alter ankle pushoff can modify gait dynamics and if resulting changes in ankle pushoff have an effect on hip muscle requirements during gait. We hypothesized that changes in ankle kinetics would be inversely related to hip muscle kinetics. Ten healthy subjects walked on a custom split-belt force-measuring treadmill at 1.25m/s. We recorded ground reaction forces and lower extremity kinematic data to calculate joint angles and internal muscle moments, powers and angular impulses. Subjects walked under three conditions: natural pushoff, decreased pushoff and increased pushoff. For the decreased pushoff condition, subjects were instructed to push less with their feet as they walked. Conversely, for the increased pushoff condition, subjects were instructed to push more with their feet. As predicted, walking with increased ankle pushoff resulted in lower peak hip flexion moment, power and angular impulse as well as lower peak hip extension moment and angular impulse (p<0.05). Our results emphasize the interchange between hip and ankle kinetics in human walking and suggest that increased ankle pushoff during gait may help to compensate for hip muscle weakness or injury and reduce hip joint forces.     

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.     

A mathematical model of hind-limb control in cats when walking backward

The “walking backward” mode was achieved within a single model of cat hind-limb locomotion with the balance maintenance only due to a change in the controlling actions (in addition to the “forward walking” mode). The skeletal part of the model contains the spine, pelvis, and two limbs consisting of the thigh, shin, and foot. The hip joint and spine mount in the thoracic region have three degrees of freedom; the knee and ankle joints have one degree of freedom. The pelvis is rigidly connected to the spine. Control is performed by model muscles (flexors and extensors of the thigh, shin, and foot). The muscle activation is performed by the effects that are typical for motoneurons that control the muscles. The feet in the support phase touch the treadmill, which moves at a constant speed. The model qualitatively reproduces multiple characteristics of feline movements during forward and backward walking (supporting its validity).     

The efficacy of a video-based marker-less tracking system for gait analysis

Most clinical gait analyses are conducted using motion capture systems which track retro-reflective markers that are placed on key landmarks of the participants. An alternative to a three-dimensional (3D) motion capture, marker-based, optical camera system may be a marker-less video-based tracking system. The aim of our study was to investigate the efficacy of the use of a marker-less tracking system in the calculation of 3D joint angles for possible use in clinical gait analysis. Ten participants walked and jogged on a treadmill and their kinematic data were captured with a marker and marker-less tracking system simultaneously. The hip, knee and ankle angles in the frontal, sagittal and transverse planes were computed. Root Mean Square differences (RMS_diff) between corresponding angles for each participant’s support phase were calculated and averaged to derive the mean within-subject RMS_diff. These within-subject means were averaged to obtain the mean between-subject RMS_diff for the relevant joint angles in the two gait conditions (walking and jogging). The RMS_diff between the two tracking systems was less than 1° for all rotations of the three joint angles of the hip and knee. However, there were slightly larger differences in the ankle joint angles. The results of this study suggest a potential application in gait analysis in clinical settings where observations of anatomical motions may provide meaningful feedback.     

The relationship between speed, contact time and peak plantar pressure in terrestrial walking of bonobos

The aim of this paper is twofold. Firstly, we investigate whether contact times, as recorded by pedobarographic systems during quadrupedal and bipedal walking of bonobos, can be used to reliably calculate actual velocities, by applying formulae based on lateral-view video recordings. Secondly, we investigate the effect of speed on peak plantar pressures during bipedal and quadrupedal walking of the bonobo. Data were obtained from 4 individuals from a group of bonobos at the Animal Park Planckendael. From our study, we can conclude that both walking speeds calculated from contact times and lower leg length or simply from recorded contact times are good estimators for walking speed, when direct observation of the latter is impossible. Further, it was found that effects of speed on peak plantar pressures and vertical forces are absent or at least subtle in comparison to a large variation in pressure patterns. In bonobos, the same pressure patterns are used at all walking speeds, and, in consequence, we do not expect major changes in foot function.     

Limb morphology,bipedal gait,and the energetics of hominid locomotion

How viable is the argument that increased locomotor efficiency was an important agent in the origin of hominid bipedalism? This study reviews data from the literature on the cost of human bipedal walking and running and compares it to data on quadrupedal mammals including several non-human primate species. Literature data comparing the cost of bipedal and quadrupedal locomotion in trained capuchin monkeys and chimpanzees are also considered. It is concluded that increased energetic efficiency would not have accrued to early bipeds. Presumably, however, selection for improved efficiency in the bipedal stance would have occurred once the transition was made. Would such a process have included selection for increased limb length? Data on the cost of locomotion vs. limb length reveal no significant relationship between these variables in 21 species of mammals or in human walking or running. © 1996 Wiley-Liss, Inc.     

The relative contributions of sagittal,frontal, and transverse joint works to self-paced incline and decline slope walking

Positive and negative work are generated at the lower limb joints in order to locomote over various terrains. Joint work quantifies the changes in energy that are necessary to adapt gait to environmental demands. The aim of this study was to quantify 3D joint work at the hip, knee, and ankle during slope walking. Work was calculated for ten males (23.9 ± 1.1 years) walking at a self-selected speed on inclines and declines (−20, −12, −6, 0, 6, 12, 20 degrees). Sagittal positive work significantly increased at the hip, knee, and ankle for incline walking (for example, hip positive work increased 153%, 280%, and 453% for 6, 12, and 20 degrees, respectively; knee and ankle positive work also increased) (p ≤ 0.05), in order to raise and propel the body forward. Sagittal negative work increased significantly at the hip, knee and ankle for decline walking (for example, knee negative work increased 193%, 355%, and 496% for −6, −12, and −20 degrees, respectively; hip and ankle negative work also increased) (p ≤ 0.05), in order to control body descent. These substantial changes in work will be especially challenging for people with compromised strength due to age and disease. Furthermore, changes in work were not limited to the sagittal plane: 46% of the total hip joint work occurred in the frontal and transverse planes for six degree decline walking. Thus, decline walking placed greater demands on the hip ab/adductors and rotators, and this may be related to the greater risk of falls observed for descent versus ascent.