Human Bipedalism Research Articles

The Origins of Human Bipedalism

The Origins of Human Bipedalism

Response to Comment on "Origin of Human Bipedalism As an Adaptation for Locomotion on Flexible Branches"

Begun et al . purport to present technical concerns regarding our case for an arboreal origin for terrestrial bipedalism in early hominins, but merely reiterate their knuckle-walking hypothesis, which lacks support from the fossil record and is highly unparsimonious. The technical concerns are refuted by published studies cited in our study and thus do not affect our original conclusions.

Reappraisal of the comparative cost of human locomotion using gait-specific allometric analyses

The alleged high net energy cost of running and low net energy cost of walking in humans have played an important role in the interpretation of the evolution of human bipedalism and the biomechanical determinants of the metabolic cost of locomotion. This study re-explores how the net metabolic energy cost of running and walking (J kg(-1) m(-1)) in humans compares to that of animals of similar mass using new allometric analyses of previously published data. Firstly, this study shows that the use of the slope of the regression between the rate of energy expenditure and speed to calculate the net energy cost of locomotion overestimates the net cost of human running. Also, the net energy cost of human running is only 17% higher than that predicted based on their mass. This value is not exceptional given that over a quarter of the previously examined mammals and birds have a net energy cost of running that is 17% or more above their allometrically predicted value. Using a new allometric equation for the net energy cost of walking, this study also shows that human walking is 20% less expensive than predicted for their mass. Of the animals used to generate this equation, 25% have a relatively lower net cost of walking compared with their allometrically predicted value. This new walking allometric analysis also indicates that the scaling of the net energy cost of locomotion with body mass is gait dependent. In conclusion, the net costs of running and walking in humans are moderately different from those predicted from allometry and are not remarkable for an animal of its size.

Chimpanzee locomotor energetics and the origin of human bipedalism

Bipedal walking is evident in the earliest hominins [Zollikofer CPE, Ponce de Leon MS, Lieberman DE, Guy F, Pilbeam D, et al. (2005) Nature 434:755-759], but why our unique two-legged gait evolved remains unknown. Here, we analyze walking energetics and biomechanics for adult chimpanzees and humans to investigate the long-standing hypothesis that bipedalism reduced the energy cost of walking compared with our ape-like ancestors [Rodman PS, McHenry HM (1980) Am J Phys Anthropol 52:103-106]. Consistent with previous work on juvenile chimpanzees [Taylor CR, Rowntree VJ (1973) Science 179:186-187], we find that bipedal and quadrupedal walking costs are not significantly different in our sample of adult chimpanzees. However, a more detailed analysis reveals significant differences in bipedal and quadrupedal cost in most individuals, which are masked when subjects are examined as a group. Furthermore, human walking is approximately 75% less costly than both quadrupedal and bipedal walking in chimpanzees. Variation in cost between bipedal and quadrupedal walking, as well as between chimpanzees and humans, is well explained by biomechanical differences in anatomy and gait, with the decreased cost of human walking attributable to our more extended hip and a longer hindlimb. Analyses of these features in early fossil hominins, coupled with analyses of bipedal walking in chimpanzees, indicate that bipedalism in early, ape-like hominins could indeed have been less costly than quadrupedal knucklewalking.

Limb length and locomotor biomechanics in the genus Homo: An experimental study

The striking variation in limb proportions within the genus Homo during the Pleistocene has important implications for understanding biomechanics in the later evolution of human bipedalism, because longer limbs and limb segments may increase bending moments about bones and joints. This research tested the hypothesis that long lower limbs and tibiae bring about increases in A-P bending forces on the lower limb during the stance phase of human walking. High-speed 3-D video data, force plates, and motion analysis software were used to analyze the walking gait of 27 modern human subjects. Limb length, as well as absolute and relative tibia length, were tested for associations with a number of kinetic and kinematic variables. Results show that individuals with longer limbs do incur greater bending moments along the lower limb during the first half of stance phase. During the second half of stance, individuals moderate bending moments through a complex of compensatory mechanisms, including keeping the knee in a more extended position. Neither absolute nor relative tibia length had any effect on the kinetic or kinematic variables tested. If these patterns apply to fossil Homo, groups with relatively long limbs (e.g. H. ergaster or early H. sapiens) may have experienced elevated bending forces along the lower limb during walking compared to those with relatively shorter limbs (e.g. the Neandertals). These increased forces could have led to greater reinforcement of joints and diaphyses. These results must be considered when formulating explanations for variation in limb morphology among Pleistocene hominins.

Origin of Human Bipedalism As an Adaptation for Locomotion on Flexible Branches

Human bipedalism is commonly thought to have evolved from a quadrupedal terrestrial precursor, yet some recent paleontological evidence suggests that adaptations for bipedalism arose in an arboreal context. However, the adaptive benefit of arboreal bipedalism has been unknown. Here we show that it allows the most arboreal great ape, the orangutan, to access supports too flexible to be negotiated otherwise. Orangutans react to branch flexibility like humans running on springy tracks, by increasing knee and hip extension, whereas all other primatesdothe reverse. Human bipedalism is thus less an innovation than an exploitation of a locomotor behavior retained from the common great ape ancestor.

Walking on Trees

Observations of modern orangutans suggest that human bipedalism may have evolved in the trees rather than on the ground.

Resolving Head Rotation for Human Bipedalism

Alignment of the body to the gravitational vertical is considered to be the key to human bipedalism. However, changes to the semicircular canals during human evolution suggest that the sense of head rotation that they provide is important for modern human bipedal locomotion. When walking, the canals signal a mix of head rotations associated with path turns, balance perturbations, and other body movements. It is uncertain how the brain uses this information. Here, we show dual roles for the semicircular canals in balance control and navigation control. We electrically evoke a head-fixed virtual rotation signal from semicircular canal nerves as subjects walk in the dark with their head held in different orientations. Depending on head orientation, we can either steer walking by "remote control" or produce balance disturbances. This shows that the brain resolves the canal signal according to head posture into Earth-referenced orthogonal components and uses rotations in vertical planes to control balance and rotations in the horizontal plane to navigate. Because the semicircular canals are concerned with movement rather than detecting vertical alignment, this result shows the importance of movement control and agility rather than precise vertical alignment of the body for human bipedalism.

Locomotor versatility in the white-handed gibbon ( Hylobates lar): A spatiotemporal analysis of the bipedal, tripedal, and quadrupedal gaits

This study gives a qualitative and quantitative description of the different terrestrial locomotor modes of a group of white-handed gibbons ( Hylobates lar) from the Wild Animal Park Planckendael, Belgium. The gibbons were filmed during voluntary locomotion on a grassy and smooth substrate and on a pole. These video images allowed us to define seven different gait types, based on spatial and temporal footfall patterns. Consequent digitization of the video images ( n = 254) yielded duty factors, stride lengths, and stride frequencies of the fore- and hind limbs during locomotion at a wide range of speeds. These spatiotemporal gait characteristics were regressed against velocity, and the regression lines of the different gait types were compared. In addition, gibbon bipedalism was compared with bonobo ( Pan paniscus) and human bipedalism. Gibbons appear to be very versatile animals, using a bipedal, tripedal, or quadrupedal gait during terrestrial travel with an overlapping speed range. The spatiotemporal characteristics of these gaits are largely similar, although they have clearly distinct footfall patterns. Bipedal walking on the pole is slightly different from terrestrial bipedalism, but differences between substrate types (grass vs. catwalk) are subtle. During bipedalism, gibbons increase both stride length and frequency to increase speed, just as humans and bonobos do, but at a given speed, gibbons take relatively larger strides at lower rates. Bipedal walking in gibbons also appears to be relatively fast—gibbons could keep on walking at speeds where humans have to start running. Apparently, adaptations for arboreal locomotion have not constrained the terrestrial locomotor abilities of gibbons. This may indicate that the step from an arboreal ancestral ape to a terrestrial, upright bipedal hominin might not be difficult and that structural specializations are not a prerequisite for adopting a (non-habitual) bipedal gait.

Human balance, the evolution of bipedalism and dysequilibrium syndrome

A new model of the uniqueness, nature and evolution of human bipedality is presented in the context of the etiology of the balance disorder of dysequilibrium syndrome. Human bipedality is biologically novel in several remarkable respects. Humans are (a) obligate, habitual and diverse in their bipedalism, (b) hold their body carriage spinally erect in a multisegmental "antigravity pole", (c) use their forelimbs exclusively for nonlocomotion, (d) support their body weight exclusively by vertical balance and normally never use prehensile holds. Further, human bipedalism is combined with (e) upper body actions that quickly shift the body's center of mass (e.g. tennis serves, piggy-back carrying of children), (f) use transient unstable erect positions (dance, kicking and fighting), (g) body height that makes falls injurious, (h) stiff gait walking, and (i) endurance running. Underlying these novelties, I conjecture, is a species specific human vertical balance faculty. This faculty synchronizes any action with a skeletomuscular adjustment that corrects its potential destabilizing impact upon the projection of the body's center of mass over its foot support. The balance faculty depends upon internal models of the erect vertical body's geometrical relationship (and its deviations) to its support base. Due to the situation that humans are obligate erect terrestrial animals, two frameworks - the body- and gravity-defined frameworks - are in constant alignment in the vertical z-axis. This alignment allows human balance to adapt egocentric body cognitions to detect body deviations from the gravitational vertical. This link between human balance and the processing of geometrical orientation, I propose, accounts for the close link between balance and spatial cognition found in the cerebral cortex. I argue that cortical areas processing the spatial and other cognitions needed to enable vertical balance was an important reason for brain size expansion of Homo erectus. A novel source of evidence for this conjecture is the rare autosomal recessive condition of dysequilibrium syndrome. In dysequilibrium syndrome, individuals fail to learn to walk bipedally (with this not being due to sensory, vestibular nor motor coordination defects). Dysequilibrium syndrome is associated with severe spatial deficits that I conjecture underlie its balance dysfunction. The associated brain defects and gene mutations of dysequilibrium syndrome provide new opportunities to investigate (i) the neurological processes responsible for the human specific balance faculty, and (ii) through gene dating techniques, its evolution.

Functional analysis of the gibbon foot during terrestrial bipedal walking: Plantar pressure distributions and three‐dimensional ground reaction forces

This paper gives a detailed analysis of bipedal walking in the white-handed gibbon, based on collected pressure and force data. These data were obtained from four gibbons in the Wild Animal Park, Planckendael, Belgium, by using a walkway with integrated force plate and pressure mat. This is the first study that collects and describes dynamic plantar pressure data of bipedally walking gibbons, and combines these with force plate data. The combination of these data with previously described roll-off patterns of gibbons, based on general observations, video images, force plates, and EMG data, gives us a detailed description of foot function during gibbon bipedalism. In addition, we compare the observed characteristics of hylobatid bipedalism with the main characteristics of bonobo and human bipedalism. We found that gibbons are midfoot/heel plantigrade, and lack the typical heel-strike of other hominoids. The hallux is widely abducted and touches down at the onset of the stance phase, which results in an L-shaped course of the center of pressure. The vertical force curve is trapezoid to triangular in shape, with high peak values compared to humans. The braking component is shorter than the accelerating component, and shortens further at higher walking velocities. Speed has a significant influence on the forefoot peak pressures and on most of the defined gait parameters (e.g., vertical force peak), and it alters the foot contact pattern as well. The investigation of existing form-function relationships in nonhuman primates is essential for the interpretation of fossil remains, and might help us understand the evolution of habitual bipedal walking in hominids.

Form and function of the oblique cord (chorda obliqua) in anthropoid primates

The oblique cord of the forearm in humans is a ligament connecting the anterolateral aspect of the ulna proximally to the posteromedial aspect of the radius distally, inserting just below the radial tuberosity. Its functional significance is uncertain, but it has been proposed that the ligament limits supination of the forearm or aids in resisting buckling failure in the curved radius. These functional explanations are unsatisfactory for bipedal humans who do not use their forelimbs in weight support. Furthermore, there are no evolutionary explanations for its presence in humans. The purpose of this study was to investigate the distribution and form of the oblique cord in non-human primates and to explore its possible functional significance and evolutionary origin in humans. Soft tissue dissections of anthropoid primates revealed the presence of an oblique cord in New and Old World monkeys and Asian apes. It was absent in all atelines. Passive manipulation of the two forearm bones in all specimens revealed that the ligament becomes most taut in pronation contrary to the proposed human condition. Isolated radii show that the oblique cord's radial insertion lacks a clear relationship with bone curvature, thus providing little support for the hypothesis of preventing buckling failure. The oblique cord's involvement in reducing bending strain in the curved radius is also unlikely. It is suggested here that the ligament may have a role in maintaining elbow stability in quadrupedal primates. Therefore, the function of the oblique cord in anthropoid primates suggests that its presence in modern humans may be a morphological and functional remnant from a quadrupedal ancestry.

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.

Fossils, feet and the evolution of human bipedal locomotion.

We review the evolution of human bipedal locomotion with a particular emphasis on the evolution of the foot. We begin in the early twentieth century and focus particularly on hypotheses of an ape-like ancestor for humans and human bipedal locomotion put forward by a succession of Gregory, Keith, Morton and Schultz. We give consideration to Morton's (1935) synthesis of foot evolution, in which he argues that the foot of the common ancestor of modern humans and the African apes would be intermediate between the foot of Pan and Hylobates whereas the foot of a hypothetical early hominin would be intermediate between that of a gorilla and a modern human. From this base rooted in comparative anatomy of living primates we trace changing ideas about the evolution of human bipedalism as increasing amounts of postcranial fossil material were discovered. Attention is given to the work of John Napier and John Robinson who were pioneers in the interpretation of Plio-Pleistocene hominin skeletons in the 1960s. This is the period when the wealth of evidence from the southern African australopithecine sites was beginning to be appreciated and Olduvai Gorge was revealing its first evidence for Homo habilis. In more recent years, the discovery of the Laetoli footprint trail, the AL 288-1 (A. afarensis) skeleton, the wealth of postcranial material from Koobi Fora, the Nariokotome Homo ergaster skeleton, Little Foot (Stw 573) from Sterkfontein in South Africa, and more recently tantalizing material assigned to the new and very early taxa Orrorin tugenensis, Ardipithecus ramidus and Sahelanthropus tchadensis has fuelled debate and speculation. The varying interpretations based on this material, together with changing theoretical insights and analytical approaches, is discussed and assessed in the context of new three-dimensional morphometric analyses of australopithecine and Homo foot bones, suggesting that there may have been greater diversity in human bipedalism in the earlier phases of our evolutionary history than previously suspected.

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.

Locomotion in bonobos (Pan paniscus): differences and similarities between bipedal and quadrupedal terrestrial walking, and a comparison with other locomotor modes.

One of the great ongoing debates in palaeo-anthropology is when, and how, hominids acquired habitual bipedal locomotion. The newly adopted bipedal gait and the ancestral quadrupedal gait are most often considered as very distinct, with each habitual locomotor mode showing corresponding anatomical adaptations. Bonobos (Pan paniscus), along with common chimpanzees (P. troglodytes), are the closest living relatives to humans and their locomotion is valuable for comparison with other primates, and to gain an insight in the acquisition of human bipedalism. Bonobos are habitual quadrupeds, but they also engage in bipedal locomotion, both on terrestrial and in arboreal substrates. In terms of kinematics and dynamics, the contrast between bipedal and quadrupedal walking seems to be more subtle than one might expect. Apart from the trunk being approximately 37 degrees more erect during bipedal locomotion, the leg movements are rather similar. Apart from the heel, plantar pressure distributions show subtle differences between bipedal and quadrupedal locomotion. Regardless, variability is high, and various intermediate forms of locomotion (e.g. tripedal walking) exist both in captivity and in the wild. Moreover, there is overlap between the characteristics of walking and other locomotor modes, as we show with new data of walking on an inclined pole and of vertical squat jumps. We suggest that there is great overlap between the many locomotor modes in bonobos, and that the required polyvalence is reflected in their anatomy. This may hamper the development of one highly specialized gait (i.e. bipedalism), which would constrain performance of the other types of locomotion.

Origin of human bipedality

Origin of human bipedality

Human footprints in Pleistocene volcanic ash

These ancient tracks are the oldest known to have been made by fully bipedal humans. We have analysed three fossilized trackways of human footprints in a zeolite-rich pyroclastic flow dated to 385,000–325,000 years ago (kyr), discovered along the western margin of the Roccamonfina volcanic complex in southern Italy. We believe that these tracks are the oldest human footprints found so far and that they were made by hominids who had a fully bipedal, free-standing gait, using their hands only to steady themselves on the difficult descent.

Comparison of Stability against External Perturbations between Human Upright Posture and Ape-like Bent-hip, Bent-knee Posture

In order to better understand the background behind the origin of human bipedality, the functional significance of the human upright posture in terms of stability should be more investigated. Therefore, in this study, we examined postural stability of human upright posture against external perturbations and compared it with that of ape-like bent-hip, bent-knee posture. Five male participants were ask to stand on a stabilometer and forward and backward horizontal surface perturbations were applied. Fluctuation of the center of pressure, electromyographic signals, and body sway were monitored.The results show the human upright posture, despite of its relatively higher location of the center of body mass (COM), is not significantly less stable against the external perturbations than the ape-like flexed posture. Moreover, it is suggested that postural adjustments against external perturbations become more difficult in the flexed posture, since muscles have to be more activated not only to cope with the body sway, but also to sustain the flexed posture against the gravity. Therefore, the upright posture seems to be more advantageous than the mechanically more stable flexed posture as a whole. Due to this inherent advantage, it is deduced that the human ancestors could probably transfer to the upright posture in a relatively early stage, once bipedality becomes necessary.

Physiologic aspects of lymphatic contractile function: current perspectives.

The lymphatic system plays an important role in fluid/macromolecular balance, lipid absorption, and immune functions, and is involved in many different pathologic conditions, like inflammation, spread of cancer cells, and lymphedema. There are several forces that drive lymph centripetally. Extrinsic driving forces, or the passive lymph pump, include lymph formation, arterial pulsations, skeletal muscles contractions, fluctuations of central venous pressure, gastrointestinal peristalsis, and respiration. Intrinsic forces, or the active lymph pump, are the result of coordinated contractions of lymphangions, the morpho-functional units of the lymphatic vessels, which include the valve and portion of the vessel extending to the next valve. The contractions of the lymphangions are initiated by the pacemaker activity of the smooth muscle cells of lymphangion wall. Transmural pressure is an important hydrodynamic factor that modulates pacemaking. Under conditions of low filling, lymphangions might produce negative intraluminal pressures and a suction effect. Because of the complicated hydrodynamic conditions in lymphatic beds, the passive and active lymph pumps sometimes work together to propel lymph centripetally. In other cases (i.e., under conditions of enhanced lymph flow), flow-mediated inhibition of the active lymph pump could serve to decrease lymphatic outflow resistance and save metabolic energy when the driving force of the passive lymph pump is enough to propel lymph. We have recently found that there are profound differences in the pressure and flow sensitivities of lymphatic vessels derived from different tissues, such as the thoracic duct and mesenteric lymphatics. Such results, when considered in light of the controversy surrounding some studies performed in different animals, lead to the idea that the active lymph pumps in humans may have greater regional differences in contractile function than has been seen in animals, because of the upright posture in bipedal humans. This posture creates an additional outflow resistance for lymphatics of the lower part of the body. Thus, despite the ongoing attempts to determine the mechanisms of lymphatic diseases and useful therapies to treat them, there are many disputable or unknown issues regarding the physiology of lymph transport in humans.