Human Bipedalism Research Articles

機械工学から直立二足歩行の起源と進化を探る(<連載講座>機械工学は21世紀を拓けるか?(第4回))

機械工学から直立二足歩行の起源と進化を探る(<連載講座>機械工学は21世紀を拓けるか?(第4回))

Force direction pattern stabilizes sagittal plane mechanics of human walking

The neural control and mechanics of human bipedalism are inadequately understood. The variable at the interface of neural control and body mechanics that is key to upright posture during human walking is the force of the ground on the foot (ground reaction force, F). We present a model that predicts sagittal plane F direction as passing through a divergent point (DP) fixed in a reference frame attached to the person. Four reference frames were tested to identify which provided the simplest and most accurate description of F direction. For all reference frames, the DP model predicted nearly all the observed variation in F direction and whole body angular momentum during single leg stance. The reference frame with vertical orientation and with origin on the pelvis provided the best combination of accuracy and simplicity. The DP was located higher than the CM and the predicted F produced a pattern of torque about the CM that caused body pitch oscillations that disrupted upright posture. Despite those oscillations, that torque was evidence of a stability mechanism that may be a critical component enabling humans to remain upright while walking and performing other tasks.

Sagittal spino-pelvic balance is a crucial analysis for normal and degenerative spine

Human beings are the only vertebrates to maintain an upright, totally vertical, bipedal position submitted to the law of gravity. Unlike other vertebrates, the human spine comprises successive, opposing curves which allows the trunk to maintain an erect position. Lumbar lordosis is found in no other species. Bipedal locomotion in non-human primates is extremely limited. Those species often use their long forelimbs in pronation position to keep balance when trying to perform a bipedal displacement. Erect position is linked with pelvic rotation and enlargement, associated with modifications of the spine sagittal curvatures and muscle adaptation [1]. Human bipedalism is exclusive, stable and ergonomic. In order to maintain this posture for prolonged periods, the erect position should be economical in terms of energy expenditure. To correctly analyze the erect posture, it is necessary to define spinal and pelvic parameters, and to correlate those parameters with global parameters in full standing position. Since Hippocrates provided a precise description of the segments and normal curves of the spine over 2000 years ago, clinicians have attempted to understand the mechanics underlying spinal balance [2]. For an individual to stand, it is necessary that the projection of the centre of the body mass on the ground plane lies behind the toes and is in front of the heels (the base of support). In order to maintain this position, for any individual, there will be an optimal combination of spinal, spino-pelvic and lower limb joint alignments, requiring the least muscle energy expenditure combined with the least discomfort, potentially caused by stretching of ligaments and joint capsules. Understanding the diversity of human spinal shape and orientation is pivotal to the understanding of the achievement of this optimal position, and to its restoration, in cases of sagittal imbalance. Despite ongoing interest in assessing the nature of the ‘normal’ curvature of the spine, there are no universally accepted parameters for its form [5, 6]. The hip joint, instead of being considered as a joint solely capable of allowing femoral movement against a fixed pelvis, should be considered as a joint that allows rotation of the pelvis against fixed femurs [5]. Rotation of the hip joint can, therefore, be considered as a fundamental factor that can interfere with the spinal shape through the pelvic anatomy. The aims of this supplement are to review published data using the current techniques, to compare these different methods and to demonstrate that it is possible today to obtain clear parameters allowing a correct analysis of the sagittal plane of our substantial patient population. Case studies will be used to illustrate the importance of proper consideration of the thoracic kyphosis in relation to the lumbo-pelvic lordosis and the pelvic shape. Those cases will also demonstrate the importance of the degenerative process acting as a new parameter and its influence on modification of the spine balance. The initial spino-pelvic organization will also interfere with the degenerative process of the spine [3]. The biomechanics of the spine differ according to the spino-pelvic shape [4]. Some spinal curvatures and spino-pelvic situations are more frequently subject to specific degenerative evolution [7]. As spine-aging process is also genetically driven, it is easy to understand that the degenerative process can vary dramatically from one patient to another leading to different imbalance situation with a similar spino-pelvic organization at the beginning. Sagittal balance analysis, and the spino-pelvic organization linked to it, is a major tool when attempting to propose a diagnosis and a treatment for lumbalgia and degenerative spine disease. Spinal imbalance should be taken into consideration before initiation of any kind of treatment including conservative or surgical procedures.

Comparative triceps surae morphology in primates: a review.

Primate locomotor evolution, particularly the evolution of bipedalism, is often examined through morphological studies. Many of these studies have examined the uniqueness of the primate forelimb, and others have examined the primate hip and thigh. Few data exist, however, regarding the myology and function of the leg muscles, even though the ankle plantar flexors are highly important during human bipedalism. In this paper, we draw together data on the fiber type and muscle mass variation in the ankle plantar flexors of primates and make comparisons to other mammals. The data suggest that great apes, atelines, and lorisines exhibit similarity in the mass distribution of the triceps surae. We conclude that variation in triceps surae may be related to the shared locomotor mode exhibited by these groups and that triceps surae morphology, which approaches that of humans, may be related to frequent use of semiplantigrade locomotion and vertical climbing.

Évolution du bassin et de la hanche « à travers les âges » : des primates à l’homme moderne

The evolution to a bipedal mode of locomotion was accompanied by a verticalization of the spine and a modification in the shape of the pelvis: horizontal curvature and sagittal rotation. Phylogenesis meets ontogenesis: flat bones in fetuses similar to the monkey, australopithecus features at birth and “human-like” features by 7 or 8 years of age. These anatomical modifications explain the characteristics of human bipedalism: stable, economical, with hip and knee extension in the standing position with little lateral motion. Some pathologies induce a regression to a more archaic mode of bipedal locomotion.

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

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

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.

Knuckle-walking in Sivapithecus? The combined effects of homology and homoplasy with possible implications for pongine dispersals

Sivapithecus is a Miocene great ape from South Asia that is orangutan-like cranially but is distinctive postcranially. Work by others shows that the humerus resembles large terrestrial cercopithecoids proximally and suspensory hominoids distally, but most functional interpretations nevertheless situate Sivapithecus in an arboreal setting. We present a new quantitative analysis of the Sivapithecus capitate and hamate. Though the functional morphology of both bones suggests some degree of arboreality, the overall morphology is most similar to knuckle-walking African apes. Other features of the Sivapithecus humerus and hind limb are also functionally consistent with knuckle-walking, and we suggest that this locomotor behavior is a valid alternative functional interpretation of the postcranial morphology. We speculate that knuckle-walking in Sivapithecus would have evolved independently from African apes, perhaps for similar ecological reasons. The discovery of a possible pongine knuckle-walker challenges the hypotheses that (1) knuckle-walking evolved only once in hominoids and (2) knuckle-walking is too highly specialized to be the positional behavior from which human bipedalism evolved. The possibility of knuckle-walking in Sivapithecus may help to explain not only the curious combination of characters that typify the postcranium but also the unique postcranial morphology of extant Pongo. Furthermore, it may clarify the distribution of fossil pongines across many ecological zones in Eurasia in the Miocene and Pleistocene, as well as, independently, the spread of African apes across a diversity of environments in equatorial Africa.

Interpreting the Role of Climbing in Primate Locomotor Evolution: Are the Biomechanics of Climbing Influenced by Habitual Substrate Use and Anatomy?

Vertical climbing is widely accepted to have played an important role in the origins of both primate locomotion and of human bipedalism. Yet, only a few researchers have compared climbing mechanics in quadrupedal primates that vary in their degree of arboreality. It is assumed that primates using vertical climbing with a relatively high frequency will have morphological and behavioral specializations that facilitate efficient climbing mechanics. We test this assumption by examining whether time spent habitually engaged in climbing influences locomotor parameters such as footfall sequence, peak forces, and joint excursions during vertical climbing. Previous studies have shown that during climbing, the pronograde and semiterrestrial Macaca fuscata differs in these parameters compared to the more arboreal and highly specialized, antipronograde Ateles geoffroyi. Here, we examine whether a fully arboreal, quadrupedal primate that does not regularly arm-swing will exhibit gait and force distribution patterns intermediate between those of Macaca fuscata and Ateles geoffroyi. We collected footfall sequence, limb peak vertical forces, and 3D hindlimb excursion data for Macaca fascicularis during climbing on a stationary pole instrumented with a force transducer. Results show that footfall sequences are similar between macaque species, whereas peak force distributions and hindlimb excursions for Macaca fascicularis are intermediate between values reported for M. fuscata and Ateles geoffroyi. These results support the notion that time spent climbing is reflected in climbing mechanics, even though morphology may not provide for efficient mechanics, and highlight the important role of arboreal locomotor activity in determining the pathways of primate locomotor evolution.

Pre-existent vertebral rotation in the human spine is influenced by body position

Both the humans as well as the quadrupedal spine have been shown to exhibit a pattern of pre-existent rotation that is similar in direction to what is found in the most common types of idiopathic scoliosis. It has been postulated that human bipedalism introduces forces to the spine that increase a tendency of the vertebrae to rotate. The objective of this study was to examine the effect of body position on vertebral rotation in vivo. Thirty asymptomatic volunteers underwent magnetic resonance imaging scanning of the spine (T2-L5) in three different body positions; upright, quadrupedal-like (on hands-and-knees) and supine. Vertebral rotation in the local transverse plane was measured according to a pre-established method and compared at different spinal levels between the three body positions. It was shown that in all three positions the mid- and lower thoracic vertebrae were predominantly rotated to the right. However, vertebral rotation was significantly less in the quadrupedal position than in both the standing upright and supine positions.

Cross-Sectional Morphology of the Femoral Neck of Wild Chimpanzees

To understand the mechanical effects of different modes of locomotion on the femoral neck of chimpanzees, we investigated the cross-sectional morphology of the femoral neck of 4 chimpanzees (Pan troglodytes schweinfurthii) collected from the Mahale Mountains, Tanzania. We performed serial computed tomography (CT) scans of the neck from the femoral head to the base of the neck perpendicular to the long axis of the neck. We measured the cortical thickness of the serial 5 cross sections of the neck region every 45° around the circumference, i.e., 8 points per section, and examined the cross-sectional properties of the mid-section. When we compared the superior and inferior parts of the cortical thickness of the femoral neck, the inferior part exhibited the greatest cortical thickness whereas the superior part had the smallest values in every specimen. Researchers have also observed such regional differences between superior and inferior cortical thicknesses in bipedal humans and other primates, although these differences are not as large in the chimpanzee as in bipedal hominini. The present study differed from the past study on hominini and chimpanzees in that the superior anterior (SA) part exhibited greater cortical thickness in chimpanzees. We believe these observations reflect the structural strengthening of parts of the chimpanzee femoral neck that is needed to accommodate the mechanical loads imposed by arboreal vertical climbing and terrestrial quadrupedal and bipedal locomotion.

Evolution of genomic imprinting in humans: does bipedalism have a role?

Recent studies have indicated that genomic imprinting is less conserved in human placenta and fetuses than in mice. Studies in mice confirm evolutionary predictions that imprinted genes have an important role in fetal growth via their effects on placental function, nutrient demand and transfer. Here, I argue that the development of bipedalism in humans might have contributed to a reduced role for imprinted genes in fetal growth. As a consequence of bipedalism, the shape of the human pelvis has changed, leading to a reduced gestation period and smaller 'premature' babies. This overarching selective pressure could, in turn, lead to a relaxation of the silencing of those imprinted genes that reduce fetal growth, a prediction borne out by current data.

The Great Divides: Ardipithecus ramidus Reveals the Postcrania of Our Last Common Ancestors with African Apes

Genomic comparisons have established the chimpanzee and bonobo as our closest living relatives. However, the intricacies of gene regulation and expression caution against the use of these extant apes in deducing the anatomical structure of the last common ancestor that we shared with them. Evidence for this structure must therefore be sought from the fossil record. Until now, that record has provided few relevant data because available fossils were too recent or too incomplete. Evidence from Ardipithecus ramidus now suggests that the last common ancestor lacked the hand, foot, pelvic, vertebral, and limb structures and proportions specialized for suspension, vertical climbing, and knuckle-walking among extant African apes. If this hypothesis is correct, each extant African ape genus must have independently acquired these specializations from more generalized ancestors who still practiced careful arboreal climbing and bridging. African apes and hominids acquired advanced orthogrady in parallel. Hominoid spinal invagination is an embryogenetic mechanism that reoriented the shoulder girdle more laterally. It was unaccompanied by substantial lumbar spine abbreviation, an adaptation restricted to vertical climbing and/or suspension. The specialized locomotor anatomies and behaviors of chimpanzees and gorillas therefore constitute poor models for the origin and evolution of human bipedality.

Independent evolution of knuckle-walking in African apes shows that humans did not evolve from a knuckle-walking ancestor.

Despite decades of debate, it remains unclear whether human bipedalism evolved from a terrestrial knuckle-walking ancestor or from a more generalized, arboreal ape ancestor. Proponents of the knuckle-walking hypothesis focused on the wrist and hand to find morphological evidence of this behavior in the human fossil record. These studies, however, have not examined variation or development of purported knuckle-walking features in apes or other primates, data that are critical to resolution of this long-standing debate. Here we present novel data on the frequency and development of putative knuckle-walking features of the wrist in apes and monkeys. We use these data to test the hypothesis that all knuckle-walking apes share similar anatomical features and that these features can be used to reliably infer locomotor behavior in our extinct ancestors. Contrary to previous expectations, features long-assumed to indicate knuckle-walking behavior are not found in all African apes, show different developmental patterns across species, and are found in nonknuckle-walking primates as well. However, variation among African ape wrist morphology can be clearly explained if we accept the likely independent evolution of 2 fundamentally different biomechanical modes of knuckle-walking: an extended wrist posture in an arboreal environment (Pan) versus a neutral, columnar hand posture in a terrestrial environment (Gorilla). The presence of purported knuckle-walking features in the hominin wrist can thus be viewed as evidence of arboreality, not terrestriality, and provide evidence that human bipedalism evolved from a more arboreal ancestor occupying the ecological niche common to all living apes.

The Quadrupedal Nature of Human Bipedal Locomotion

During rhythmic movement, arm activity contributes to the neural excitation of leg muscles. These observations are consistent with the emergence of human bipedalism and nonhuman primate arboreal quadrupedal walking. These neural and biomechanical linkages could be exploited in rehabilitation after neurotrauma to allow the arms to give the legs a helping hand during gait rehabilitation.

Simulation study of bipedal locomotion in the Japanese macaque: Towards understanding the mechanisms of adaptive control and the evolution of human bipedalism

Simulation study of bipedal locomotion in the Japanese macaque: Towards understanding the mechanisms of adaptive control and the evolution of human bipedalism

Observations on the Prenatal Development of Human Lymphatic Vessels with Focus on Basic Structural Elements of Lymph Flow

The prenatal development of human lymphatic systems has not attracted enough attention by lymphatic researchers in the past. Yet clearly these critical, early events determine the fate and function of the human lymphatic system. The main focus of these studies was to investigate the embryonic development of human lymphangions including lymphatic valves and muscle cells, to better understand the prenatal formation of basic structural elements of lymph flow. This review in most of its parts is a short summary of the findings. It provides important information necessary for understanding the development and functioning of the human lymphatic system. The structural basis of the active lymph transport system--the lymphatic muscle cells and lymphatic valves--which is absolutely necessary for all functions of lymphatic system, is already formed during the first half of the prenatal development in humans. During the second half of this development maturation of this system is already underway. The enlargement of lymphatic muscle cells together with increases in their quantity leads to formation of the multi-layered lymphatic vessel wall, able to develop contractions strong enough to propel lymph downstream of the lymphatic channels against gravity in bipedal humans. The development of the competent valves in lymphatic vessels occurs at the same time creating the ground for effective net, unidirectional lymph flow. The data summarized here represents some of the first systematic studies of the prenatal development of lymphatic muscle cells and valves in humans.

Cranio-dental evidence of a hominin-like hyper-masticatory apparatus in Oreopithecus bambolii. Was the swamp ape a human ancestor?

The phylogeny of Oreopithecus bambolii has been controversial since Johannes Hurzeler first argued that the Late Miocene (Tortonian) primate was a fossil hominin. While most paleontologist currently exclude Oreopithecus from human ancestry, recent postcranial evidence of hominin-like bipedalism and power precision grips in Oreopithecus has rekindled interest in the fossil Italian hominoid. In this study, a comparative review of hominoid cranio-dental morphology indicates that Oreopithecus possessed a suite of hominin-like characteristics that were apparently functionally related to powerful folivorous mastication. Since the oreopithecine dentition exhibited exceptional adaptations for folivory relative to most other extant and extinct hominoids, the accessory development of a hominin-like hyper-masticatory cranio-mandibular apparatus to further enhance plant comminution and digestibility is not surprising. However, the combination of hominin-like locomotive, manual, and masticatory functional attributes appears to substantiate Hurzeler's designation of O. bambolii as a Late Miocene hominin. Additionally, an extensive compilation of hominoid cranio-dental and postcranial characteristics strongly supports a close phylogenetic relationship between Oreopithecus and the earliest African hominins Sahelanthropus and Australopithecus. The wetland paleoecology of the Tortonian island of Tuscany–Sardinia suggests that Oreopithecus was a specialized semiaquatic folivore who apparently waded bipedally into freshwater swamps to feed on aquatic plants. However, the extensive wear on the oreopithecine canines and incisors along with their manual precision grips may indicate that freshwater invertebrates were also included in their diets. Such specialized aquatic feeding behavior by these ancient Italian swamp apes seems to support Alister Hardy's hypothesis that human bipedalism and power precision grips were inherited from Late Miocene hominin ancestors who originally utilized such functional attributes for aquatic foraging in shallow water environments. Additionally, the remarkably hominin-like cranio-dental morphology of O. bambolii suggests that modern omnivorous humans probably inherited a significant number of their cranio-dental characteristics from these highly specialized aquatic plant eating hominins.

UNERTAN SYNDROME: A CASE SERIES DEMONSTRATING HUMAN DEVOLUTION

A large family with six individuals exhibiting the Unertan syndrome (UTS) was identified residing in southern Turkey. All of the individuals had mental impairments and walked on all four extremities. The practice of intra-familial marriages suggested that the UTS may be an autosomal recessive disorder, similar to previously described cases. The inferior portions of the cerebellum and vermis were absent as evidenced by MRI and CT scans. The height and head circumference of those affected were within normal ranges. Barany's test suggested normal vestibular system function. The subjects could not name objects or their close relatives. The males (n = 4) could understand simple questions and commands, but answered questions with only one or two sounds. The females (n = 2) were superior to the males with respect to language skills and walking, suggesting an association between walking and speaking abilities. One male exhibited three walking patterns at the same time: quadripedal, tiptoe, and scissor walking. Another male used two walking styles: quadripedal and toe-walking. It is emphasized that there are important differences between the UTS and the disequilibrium syndrome. It is suggested that the inability to walk upright in those affected with the UTS may be best explained by a disturbance in lateral-balance mechanisms, without being related to the cerebello-vestibular system. An interruption of locomotor development during the transition from quadripedality to bipedality may result in habitual walking on all four extremities and is normal in some children. Because quadripedal gait is an ancestral trait, individuals with the UTS, exhibiting a manifestation of reverse evolution in humans, may be considered an experiment of nature, useful in understanding the mechanisms underlying the transition from quadripedality to bipedality during human evolution. The proposed mutant gene or gene pool playing a role in human quadrupedality may also be responsible for human bipedality at the same time. Herein there is no intent to insult or injure; rather, this report is an endeavor to better understand human beings. Supplementary materials are available for this article. Go to the publisher's online edition of International Journal of Neuroscience for the following free supplemental resource(s): video clips.

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

Thorpe et al. (Reports, 1 June 2007, p. 1328) concluded that human bipedalism evolved from a type of bipedal posture they observed in extant orangutans with seemingly human-like extended knees. However, humans share knuckle-walking characters with African apes that are absent in orangutans. These are most parsimoniously explained by positing a knuckle-walking precursor to human bipedalism.