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.     

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.     

Dynamics of primate bipedal walking as viewed from the force of foot

Bipedal walking of the six species of anthropoid primates including man were examined by means of the force plate technique. Though each species has a particular pattern of bipedal walking, we can classify two types of patterns in these primates as far as the foot force is concerned. The first type includes the man, chimpanzee, and spider monkey and the second type contains the Japanese monkey, hamadryas baboon, and gibbon. It was emphasized that the similarity of man to the chimpanzee and spider monkey in bipedal walking has some evolutionary significance.     

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.     

Back muscle function during bipedal walking in chimpanzee and gibbon: implications for the evolution of human locomotion

The evolution of erect posture and locomotion continues to be a major focus of interest among paleoanthropologists and functional morphologists. To date, virtually all of our knowledge about the functional role of the back muscles in the evolution of bipedalism is based on human experimental data. In order to broaden our evolutionary perspective on the vertebral region, we have undertaken an electromyographic (EMG) analysis of three deep back muscles (multifidus, longissimus thoracis, iliocostalis lumborum) in the chimpanzee (Pan troglodytes) and gibbon (Hylobates lar) during bipedal walking. The recruitment patterns of these three muscles seen in the chimpanzee closely parallel those observed in the gibbon. The activity patterns of multifidus and longissimus are more similar to each other than either is to iliocostalis. Iliocostalis recruitment is clearly related to contact by the contralateral limb during bipedal walking in both species. It is suggested that in both the chimpanzee and gibbon, multifidus controls trunk movement primarily in the sagittal plane, iliocostalis responds to and adjusts movement in the frontal plane, while longissimus contributes to both of these functions. In many respects, the activity patterns shared by the chimpanzee and gibbon are quite consistent with recent human experimental data. This suggests a basic similarity in the mechanical constraints placed on the back during bipedalism among these three hominoids. Thus, the acquisition of habitual bipedalism in humans probably involved not so much a major change in back muscle action or function, but rather an improvement in the mechanical advantages and architecture of these muscles.     

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.     

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.     

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.     

The pelvis in the bipedalism of primates

The pelvis in the “Normalstellung” tends in man to tilt backwards. In primates, the pelvis shows a marked tendency to tilt into quadruped posture. In both the bodyweight intensifies those tendencies. Consequently, assuming erect posture and maintaining bipedal balance encounter, in primates, constant resistance from pelvis and bodyweight. In gibbons and ponginae loss of the tail, lengthening of the sacrum and broadening of the ilium combined with a pronounced development of the sacro-tuberous ligament furnish the gluteal muscles with a more powerful reflectory moment on pelvis and body. Moreover, the development of a short head of the femoral bicipital muscle, made the hip-joint to a greater degree independent of the knee-joint. Thus, maintenance of bipedal balance is better developed in gibbons and ponginae than in monkeys. In man broadening of the sacrum altered the position of the acetabulum. Together with ensuing changes of the ilium and the ischium this brought about in the pelvis a balance relationship so that the bodyweight not only causes no resistance to standing upright but even facilitates it. The essence of bipedalism, however, is not only the capacity to rise upright and to stand, but also to walk, that is, to keep the body balanced in the frontal plane on one leg. It is the ventralwards broadening of the ilium, and the sidewards bend of its middle segment, that allow, the strongest part of, the gluteus medius (and minimus) to produce a powerful abducent moment on the pelvis. The lack of a corresponding osteomuscular structure in monkeys as well as in gibbon and ponginae compels these primates to move faster and run straddle-legged, in a more or less forward-bending posture. Lastly it is found that in gibbons and ponginae, not only is the ilium markedly broadened laterally, but also that its sacral part is turned completely sidewards. This indicates that bipedalism in man has developed along quite other lines than in gibbons and ponginae.     

Estimating the complete ground reaction forces with pressure insoles in walking

This study presented a method to estimate the complete ground reaction forces from pressure insoles in walking. Five male subjects performed 10 walking trials in a laboratory. The complete ground reaction forces were collected during a right foot stride by a force plate at 1000Hz. Simultaneous plantar pressure data were collected at 100Hz by a pressure insole system with 99 sensors covering the whole plantar area. Stepwise linear regressions were performed to individually reconstruct the complete ground reaction forces in three directions from the 99 individual pressure data until redundancy among the predictors occurred. An additional linear regression was performed to reconstruct the vertical ground reaction force by the sum of the value of the 99 pressure sensors. Five other subjects performed the same walking test for validation. Estimated ground reaction forces in three directions were calculated with the developed regression models, and were compared with the real data recorded from force plate. Accuracy was represented by the correlation coefficient and the root mean square error. Results showed very good correlation in anterior-posterior (0.928) and vertical (0.989) directions, and reasonable correlation in medial-lateral direction (0.719). The root mean square error was about 12%, 5% and 28% of the peak recorded value. Future studies should aim to generalize the methods or to establish specific methods to other subjects, patients, motions, footwear and floor conditions. The method gives an extra option to study an estimation of the complete ground reaction forces in any environment without the constraints from the number and location of force plates.     

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.     

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.     

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.     

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.     

Biomechanical analysis of vertical climbing in the spider monkey and the Japanese macaque

Climbing is one of the most important components of primate locomotor modes. We previously reported that the kinesiological characteristics of vertical climbing by the spider monkey and Japanese macaque are clearly different, based on their kinetics and kinematics. In this study, a more detailed analysis using inverse dynamics was conducted to estimate the biomechanical characteristics of vertical climbing in the spider monkey and Japanese macaque. One of the main findings was the difference in forelimb use by the two species. The results of a joint moment analysis and estimates of muscular force indicate that the spider monkey uses its forelimbs to keep the body close to the substrate, rather than to generate propulsion. The forelimb of the Japanese macaque, on the other hand, likely contributes more to propulsion. This supports the idea that "forelimb-hindlimb differentiation" is promoted in the spider monkey. The estimated muscular force also suggests that the spider monkey type of climbing could develop the hindlimb extensor muscles, which are important in bipedal posture and walking. As a result, we conclude that the spider monkey type of climbing could be functionally preadaptive for human bipedalism. This type of climbing would develop the hip and knee extensor muscles, and result in more extended lower limb joints, a more erect trunk posture, and more functionally differentiated fore- and hindlimbs, all of which are important characteristics of human bipedalism.     

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.     

Optimum walking techniques for idealized animals

The vertical component of the force exerted by a foot on the ground, in the course of a step, may rise to a single maximum and decline again (as in human running) or may show two distinct maxima (as in human walking). A foot may remain on the ground for a large or small fraction of the duration of a stride. Mathematical models are used to investigate the effects of these differences of technique on the energy cost of locomotion. The optimum technique for a biped at a given speed is different from the optimum for a hypothetical many-legged animal. The optima for quadrupedal walking are likely to lie between these extremes.
The walking techniques adopted by men at different speeds are close to the optima indicated by the bipedal model. The two maxima of the force exerted by a foot are higher, and have a lower minimum between them, at higher speeds of walking. The techniques adopted by a sheep are close to the optima indicated by the many-legged model but dogs use techniques rather closer to the optima for bipeds.
The limitations of the models are discussed.     

Hand preferences in captive gorillas,orang-utans and gibbons

Hand preference was assessed in 12 gorillas (Gorilla gorilla gorilla), 13 orang-utans (Pongo pygmaeus abelii), and 9 gibbons (Hylobates lar) by using a floor retrieval task and a mesh retrieval task. Hand preference was also assessed in 8 gorillas and 8 orang-utans by using a task involving the unfastening of a hasp. A bipedal requirement during testing (mesh retrieval task) facilitated detection of hand preferences. A significant left-hand preference was found for the gibbons with 6 of 6 gibbons preferring their left hand on the mesh retrieval task. Similarly, a significant right-hand preference was found for the gorillas with 10 of 12 gorillas preferring their right hand on the mesh retrieval task. The data for the orang-utan suggest a bimodal distribution on all tasks. Since the gibbon and gorilla in the wild engage in bipedal locomotion more frequently than the orangutan, one possible interpretation for these results correlates the degree of bipedal behavior of a species in its natural environment with its readiness to exhibit a unilateral population-level hand preference.