Talk:Bipedalism: Difference between revisions

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Discussion

Re: Evolution of bipedalism

Consider:

Comment on "Origin of human bipedalism as an adaptation for locomotion on flexible branches".^[1]

Endurance running and the evolution of Homo^[2]

Resolving head rotation for human bipedalism^[3]

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References

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Begun DR, Richmond BG, Strait DS. Comment on "Origin of human bipedalism as an adaptation for locomotion on flexible branches". Science 2007;318:1066.
- Abstract: 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.
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Bramble DM, Lieberman DE. Endurance running and the evolution of Homo. Nature 2004;432:345-52.
- Abstract: Striding bipedalism is a key derived behaviour of hominids that possibly originated soon after the divergence of the chimpanzee and human lineages. Although bipedal gaits include walking and running, running is generally considered to have played no major role in human evolution because humans, like apes, are poor sprinters compared to most quadrupeds. Here we assess how well humans perform at sustained long-distance running, and review the physiological and anatomical bases of endurance running capabilities in humans and other mammals. Judged by several criteria, humans perform remarkably well at endurance running, thanks to a diverse array of features, many of which leave traces in the skeleton. The fossil evidence of these features suggests that endurance running is a derived capability of the genus Homo, originating about 2 million years ago, and may have been instrumental in the evolution of the human body form.
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Fitzpatrick RC, Butler JE, Day BL. Resolving head rotation for human bipedalism. Curr Biol 2006;16:1509-14.
- Abstract: 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.
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Kuliukas A. Wading for food the driving force of the evolution of bipedalism? Nutr Health 2002;16:267-89.
- Abstract: Evidence is accumulating that suggests that the large human brain is most likely to have evolved in littoral and estuarine habitats rich in naturally occurring essential fatty acids. This paper adds further weight to this view, suggesting that another key human trait, our bipedality might also be best explained as an adaptation to a water-side niche. Evidence is provided here that extant apes, although preferring to keep dry, go into water when driven to do so by hunger. The anecdotal evidence has suggested that they tend to do this bipedally. Here, a new empirical study of captive bonobos found them to exhibit 2% or less bipedality on the ground or in trees but over 90% when wading in water to collect food. The skeletal morphology of AL 288-1 ("Lucy") is shown to indicate a strong ability to abduct and adduct the femur. These traits, together with a remarkably platypelloid pelvis, have not yet been adequately explained by terrestrial or arboreal models for early bipedalism but are consistent with those expected in an ape that adopted a specialist side-to-side 'ice-skating' or sideways wading mode. It is argued that this explanation of A. afarensis locomotor morphology is more parsimonious than others which have plainly failed to produce a consensus. Microwear evidence of Australopithecus dentition is also presented as evidence that the drive for such a wading form of locomotion might well have been waterside foods. This model obtains further support from the paleo-habitats of the earliest known bipeds, which are consistent with the hypothesis that wading contributed to the adaptive pressure towards bipedality.
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Nakatsukasa M. Acquisition of bipedalism: the Miocene hominoid record and modern analogues for bipedal protohominids. J Anat 2004;204:385-402.
- Abstract: The well-known fossil hominoid Proconsul from the Early Miocene of Kenya was a non-specialized arboreal quadruped with strong pollicial/hallucial assisted grasping capability. It lacked most of the suspensory specializations acquired in living hominoids. Nacholapithecus, however, from the Middle Miocene of Kenya, although in part sharing with Proconsul the common primitive anatomical body design, was more specialized for orthograde climbing, 'hoisting' and bridging, with the glenoid fossae of the scapula probably being cranially orientated, the forelimbs proportionally large, and very long toes. Its tail loss suggests relatively slow movement, although tail loss may already have occurred in Proconsul. Nacholapithecus-like positional behaviour might thus have been a basis for development of more suspensory specialized positional behaviour in later hominoids. Unfortunately, after 13 Ma, there is a gap in the hominoid postcranial record in Africa until 6 Ma. Due to this gap, a scenario for later locomotor evolution prior to the divergence of Homo and Pan cannot be determined with certainty. The time gap also causes difficulties when we seek to determine polarities of morphological traits in very early hominids. Interpretation of the form-function relationships of postcranial features in incipient hominids will be difficult because it is predicted that they had incorporated bipedalism only moderately into their total positional repertoires. However, Japanese macaques, which are trained in traditional bipedal performance, may provide useful hints about bipedal adaptation in the protohominids. Kinematic analyses revealed that these macaques walked bipedally with a longer stride and lower stride frequency than used by ordinary macaques, owing to a more extended posture of the hindlimb joints. The body centre of gravity rises during the single-support phase of stance. Energetic studies of locomotion in these bipedal macaques revealed that energetic expenditure was 20-30% higher in bipedalism than in quadrupedalism, regardless of walking velocity.
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Niemitz C. A theory on the evolution of the habitual orthograde human bipedalism--the "Amphibische Generalistentheorie". Anthropol Anz 2002;60:3-66.
- Abstract: The theory is formulated that ubiquitous scarcity of energy is one of the main motors of evolution. It is concluded that our primate ancestors never came down from the trees, but rather they have always been (semi-)terrestrial. This habit is probably an old symplesiomorph trait, older than primates themselves. Terrestrial habits in primates correlate to body weight in small systematic groups (e.g., large genera, families) but are, overall, completely independent from individual body mass. An omnivorous, semiterrestrial quadrupedal locomotor generalist seems to be the most probable morpho- and eco-type for our ancestor at the threshold of a hominoid stage of our evolution. The theory presented here suggests that our hominoid ancestor lived in gallery forests and changed strata in order also to inhabit the savannah habitat as well as the shallow water of the rivers or coasts. Foraging in a wading manner was extremely favourable for an effective and, especially, seasonally independent, animal protein supply. Anatomical adaptations to orthogradism and proportions of the extremities are discussed in relation to the necessary and frequent change of habitat strata. Ultimately, human bipedalism is seen here to be derived as a consequence of the centre of body mass, which is, in primates, near the hind extremities. By contrast to other mammals entering the water, wading primates sink back on their hind limbs. Selective forces for habitat use, limb proportions and wading habits are discussed, as well as the phylogenetic origin of human affinity to water and shores in all peoples through all times, from australopithecine times through the Paleolithic until today.
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Pickford M. Palaeoenvironments and hominoid evolution. Z Morphol Anthropol 2002;83:337-48.
- Abstract: One of the key features that separates humans and their closest relatives (extinct species of the genus Homo and Praeanthropus and the australopithecines Australopithecus and Paranthropus) on the one hand, from the other hominoids, on the other, is their obligate bipedal locomotion when on the ground. This major difference from the generally quadrupedal locomotion practiced by other hominoids (Pan, Gorilla, Pongo and many extinct lineages) is reflected in many parts of the body, including all the major bones in the legs, arms, trunk and cranium. Locomotion has thus been of major interest to those interested in human origins, evolution, classification and phylogeny. A major hurdle to studies of the origins of bipedalism concerns the paucity of African hominoid fossils between 15 Ma, when all the adequately known hominoids were quadrupedal (most were pronograde, but at least one lineage was orthograde), and 4.2 Ma by which time fully bipedal hominids were established in Africa. Examination of Old World geology and palaeontology reveals a great deal about the evolution of palaeoenvironments and faunas during this period, and it is suggested that hominids evolved bipedal locomotion at the same time that there was a fundamental reorganisation of faunas towards the end of the Miocene. This faunal turnover resulted in the establishment of faunal lineages of "modern" aspect in Africa at the expense of "archaic" lineages which either went extinct or suffered a diminution of diversity. Many of the "modern" lineages were adapted to open country habitats in which grass became a major component of the diet as shown by modifications in the cheek teeth. Hominoids, in contrast, retained their traditional diet but were obliged to forage over greater and greater areas in order to do so, and this tactic led to pressures to modify the locomotor system rather than the diet. If bipedal hominids originated during this period, then the family Hominidae (sensu stricto) dates from about 8-7 Ma.
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Preuschoft H. Mechanisms for the acquisition of habitual bipedality: are there biomechanical reasons for the acquisition of upright bipedal posture? J Anat 2004;204:363-84.
- Abstract: Morphology and biomechanics are linked by causal morphogenesis ('Wolff's law') and the interplay of mutations and selection (Darwin's 'survival of the fittest'). Thus shape-based selective pressures can be determined. In both cases we need to know which biomechanical factors lead to skeletal adaptation, and which ones exert selective pressures on body shape. Each bone must be able to sustain the greatest regularly occurring loads. Smaller loads are unlikely to lead to adaptation of morphology. The highest loads occur primarily in posture and locomotion, simply because of the effect of body weight (or its multiple). In the skull, however, it is biting and chewing that result in the greatest loads. Body shape adapted for an arboreal lifestyle also smooths the way towards bipedality. Hindlimb dominance, length of the limbs in relation to the axial skeleton, grasping hands and feet, mass distribution (especially of the limb segments), thoracic shape, rib curvatures, and the position of the centre of gravity are the adaptations to arboreality that also pre-adapt for bipedality. Five divergent locomotor/morphological types have evolved from this base: arm-swinging in gibbons, forelimb-dominated slow climbing in orangutans, quadrupedalism/climbing in the African apes, an unknown mix of climbing and bipedal walking in australopithecines, and the remarkably endurant bipedal walking of humans. All other apes are also facultative bipeds, but it is the biomechanical characteristics of bipedalism in orangutans, the most arboreal great ape, which is closest to that in humans. If not evolutionary accident, what selective factor can explain why two forms adopted bipedality? Most authors tend to connect bipedal locomotion with some aspect of progressively increasing distance between trees because of climatic changes. More precise factors, in accordance with biomechanical requirements, include stone-throwing, thermoregulation or wading in shallow water. Once bipedality has been acquired, development of typical human morphology can readily be explained as adaptations for energy saving over long distances. A paper in this volume shows that load-carrying ability was enhanced from australopithecines to Homo ergaster (early African H. erectus), supporting an earlier proposition that load-carrying was an essential factor in human evolution.
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Richmond BG, Begun DR, Strait DS. Origin of human bipedalism: The knuckle-walking hypothesis revisited. Am J Phys Anthropol 2001;Suppl 33:70-105.
- Abstract: Some of the most long-standing questions in paleoanthropology concern how and why human bipedalism evolved. Over the last century, many hypotheses have been offered on the mode of locomotion from which bipedalism originated. Candidate ancestral adaptations include monkey-like arboreal or terrestrial quadrupedalism, gibbon- or orangutan-like (or other forms of) climbing and suspension, and knuckle-walking. This paper reviews the history of these hypotheses, outlines their predictions, and assesses them in light of current phylogenetic, comparative anatomical, and fossil evidence. The functional significance of characteristics of the shoulder and arm, elbow, wrist, and hand shared by African apes and humans, including their fossil relatives, most strongly supports the knuckle-walking hypothesis, which reconstructs the ancestor as being adapted to knuckle-walking and arboreal climbing. Future fossil discoveries, and a clear understanding of anthropoid locomotor anatomy, are required to ultimately test these hypotheses. If knuckle-walking was an important component of the behavioral repertoire of the prebipedal human ancestor, then we can reject scenarios on the origin of bipedalism that rely on a strictly arboreal ancestor. Moreover, paleoenvironmental data associated with the earliest hominins, and their close relatives, contradict hypotheses that place the agents of selection for bipedality in open savanna habitats. Existing hypotheses must explain why bipedalism would evolve from an ancestor that was already partly terrestrial. Many food acquisition and carrying hypotheses remain tenable in light of current evidence.
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Richmond BG, Jungers WL. Orrorin tugenensis femoral morphology and the evolution of hominin bipedalism. Science 2008;319:1662-5.
- Abstract: Bipedalism is a key human adaptation and a defining feature of the hominin clade. Fossil femora discovered in Kenya and attributed to Orrorin tugenensis, at 6 million years ago, purportedly provide the earliest postcranial evidence of hominin bipedalism, but their functional and phylogenetic affinities are controversial. We show that the O. tugenensis femur differs from those of apes and Homo and most strongly resembles those of Australopithecus and Paranthropus, indicating that O. tugenensis was bipedal but is not more closely related to Homo than to Australopithecus. Femoral morphology indicates that O. tugenensis shared distinctive hip biomechanics with australopiths, suggesting that this complex evolved early in human evolution and persisted for almost 4 million years until modifications of the hip appeared in the late Pliocene in early Homo.
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Schmitt D. Insights into the evolution of human bipedalism from experimental studies of humans and other primates. J Exp Biol 2003;206:1437-48.
- Abstract: An understanding of the evolution of human bipedalism can provide valuable insights into the biomechanical and physiological characteristics of locomotion in modern humans. The walking gaits of humans, other bipeds and most quadrupedal mammals can best be described by using an inverted-pendulum model, in which there is minimal change in flexion of the limb joints during stance phase. As a result, it seems logical that the evolution of bipedalism in humans involved a simple transition from a relatively stiff-legged quadrupedalism in a terrestrial ancestor to relatively stiff-legged bipedalism in early humans. However, experimental studies of locomotion in humans and nonhuman primates have shown that the evolution of bipedalism involved a much more complex series of transitions, originating with a relatively compliant form of quadrupedalism. These studies show that relatively compliant walking gaits allow primates to achieve fast walking speeds using long strides, low stride frequencies, relatively low peak vertical forces, and relatively high impact shock attenuation ratios. A relatively compliant, ape-like bipedal walking style is consistent with the anatomy of early hominids and may have been an effective gait for a small biped with relatively small and less stabilized joints, which had not yet completely forsaken arboreal locomotion. Laboratory-based studies of primates also suggest that human bipedalism arose not from a terrestrial ancestor but rather from a climbing, arboreal forerunner. Experimental data, in conjunction with anatomical data on early human ancestors, show clearly that a relatively stiff modern human gait and associated physiological and anatomical adaptations are not primitive retentions from a primate ancestor, but are instead recently acquired characters of our genus.
↑ Schwartz JH. The origins of human bipedalism. Science 2007;318:1065.
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Sellers WI, Dennis LA, Crompton RH. Predicting the metabolic energy costs of bipedalism using evolutionary robotics. J Exp Biol 2003;206:1127-36.
- Abstract: To understand the evolution of bipedalism among the hominoids in an ecological context we need to be able to estimate the energetic cost of locomotion in fossil forms. Ideally such an estimate would be based entirely on morphology since, except for the rare instances where footprints are preserved, this is the only primary source of evidence available. In this paper we use evolutionary robotics techniques (genetic algorithms, pattern generators and mechanical modeling) to produce a biomimetic simulation of bipedalism based on human body dimensions. The mechanical simulation is a seven-segment, two-dimensional model with motive force provided by tension generators representing the major muscle groups acting around the lower-limb joints. Metabolic energy costs are calculated from the muscle model, and bipedal gait is generated using a finite-state pattern generator whose parameters are produced using a genetic algorithm with locomotor economy (maximum distance for a fixed energy cost) as the fitness criterion. The model is validated by comparing the values it generates with those for modern humans. The result (maximum efficiency of 200 J m(-1)) is within 15% of the experimentally derived value, which is very encouraging and suggests that this is a useful analytic technique for investigating the locomotor behaviour of fossil forms. Initial work suggests that in the future this technique could be used to estimate other locomotor parameters such as top speed. In addition, the animations produced by this technique are qualitatively very convincing, which suggests that this may also be a useful technique for visualizing bipedal locomotion.
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Skoyles JR. Human balance, the evolution of bipedalism and dysequilibrium syndrome. Med Hypotheses 2006;66:1060-8.
- Abstract: 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.
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Sockol MD, Raichlen DA, Pontzer H. Chimpanzee locomotor energetics and the origin of human bipedalism. Proc Natl Acad Sci U S A 2007;104:12265-9.
- Abstract: 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.
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Spoor F, Wood B, Zonneveld F. Implications of early hominid labyrinthine morphology for evolution of human bipedal locomotion. Nature 1994;369:645-8.
- Abstract: The upright posture and obligatory bipedalism of modern humans are unique among living primates. The evolutionary history of this behaviour has traditionally been pursued by functional analysis of the postcranial skeleton and the preserved footprint trails of fossil hominids. Here we report a systematic attempt to reconstruct the locomotor behaviour of early hominids by looking at a major component of the mechanism for the unconscious perception of movement, namely by examining the vestibular system of living primates and early hominids. High-resolution computed tomography was used to generate cross-sectional images of the bony labyrinth. Among the fossil hominids the earliest species to demonstrate the modern human morphology is Homo erectus. In contrast, the semicircular canal dimensions in crania from southern Africa attributed to Australopithecus and Paranthropus resemble those of the extant great apes. Among early Homo specimens, the canal dimensions of Stw 53 are unlike those seen in any of the hominids or great apes, whereas those of SK 847 are modern-human-like.
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Stanford CB. Arboreal bipedalism in wild chimpanzees: implications for the evolution of hominid posture and locomotion. Am J Phys Anthropol 2006;129:225-31.
- Abstract: Field observations of bipedal posture and locomotion in wild chimpanzees (Pan troglodytes) can serve as key evidence for reconstructing the likely origins of bipedalism in the last prehominid human ancestor. This paper reports on a sample of bipedal bouts, recorded ad libitum, in wild chimpanzees in Bwindi Impenetrable National Park in southwestern Uganda. The Ruhija community of chimpanzees in Bwindi displays a high rate of bipedal posture. In 246.7 hr of observation from 2001-2003, 179 instances of bipedal posture lasting 5 sec or longer were recorded, for a rate of 0.73 bouts per observation hour. Bipedalism was observed only on arboreal substrates, and was almost all postural, and not locomotor. Bipedalism was part of a complex series of positional behaviors related to feeding, which included two-legged standing, one-legged standing with arm support, and other intermediate postures. Ninety-six percent of bipedal bouts occurred in a foraging context, always as a chimpanzee reached to pluck fruit from tree limbs. Bipedalism was seen in both male and female adults, less frequently among juveniles, and rarely in infants. Both the frequency and duration of bipedal bouts showed a significant positive correlation with estimated substrate diameter. Neither fruit size nor nearest-neighbor association patterns were significantly correlated with the occurrence of bipedalism. Bipedalism is seen frequently in the Bwindi chimpanzee community, in part because of the unusual observer conditions at Bwindi. Most observations of bipedalism were made when the animals were in treetops and the observer at eye-level across narrow ravines. This suggests that wild chimpanzees may engage in bipedal behavior more often than is generally appreciated. Models of the likely evolutionary origins of bipedalism are considered in the light of Bwindi bipedalism data. Bipedalism among Bwindi chimpanzees suggests the origin of bipedal posture in hominids to be related to foraging advantages in fruit trees. It suggests important arboreal advantages in upright posture. The origin of postural bipedalism may have preceded and been causally disconnected from locomotor bipedalism.
↑ Steudel-Numbers KL. Role of locomotor economy in the origin of bipedal posture and gait. Am J Phys Anthropol 2001;116:171-3.
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Sylvester AD. Locomotor decoupling and the origin of hominin bipedalism. J Theor Biol 2006;242:581-90.
- Abstract: Theoretical adaptive landscapes and mathematical representations of key constraints of evolutionary and primate biology are used to propose a new hypothesis for the origin of hominin bipedalism. These constraints suggest that the selective pressure that produced this novel form of locomotion was the need for effective suspensory and terrestrial movement. This testable hypothesis, termed the Decoupling Hypothesis, posits that bipedalism is an adaptation that enables the shoulder to maintain a high degree of mobility, a feature important to suspensory behaviors, in the face of significant demands for a high degree of stability, a feature important for highly effective terrestrial quadrupedism.
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Thorpe SK, Holder RL, Crompton RH. Origin of human bipedalism as an adaptation for locomotion on flexible branches. Science 2007;316:1328-31.
- Abstract: 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.
↑ Vaughan CL. Theories of bipedal walking: an odyssey. J Biomech 2003;36:513-23. Abstract: In this paper six theories of bipedal walking, and the evidence in support of the theories, are reviewed. They include: evolution, minimising energy consumption, maturation in children, central pattern generators, linking control and effect, and robots on two legs. Specifically, the six theories posit that: (1) bipedalism is the fundamental evolutionary adaptation that sets hominids--and therefore humans--apart from other primates; (2) locomotion is the translation of the centre of gravity along a pathway requiring the least expenditure of energy; (3) when a young child takes its first few halting steps, his or her biomechanical strategy is to minimise the risk of falling; (4) a dedicated network of interneurons in the spinal cord generates the rhythm and cyclic pattern of electromyographic signals that give rise to bipedal gait; (5) bipedal locomotion is generated through global entrainment of the neural system on the one hand, and the musculoskeletal system plus environment on the other; and (6) powered dynamic gait in a bipedal robot can be realised only through a strategy which is based on stability and real-time feedback control. The published record suggests that each of the theories has some measure of support. However, it is important to note that there are other important theories of locomotion which have not been covered in this review. Despite such omissions, this odyssey has explored the wide spectrum of bipedal walking, from its origins through to the integration of the nervous, muscular and skeletal systems.
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Wang WJ, Crompton RH, Li Y, Gunther MM. Energy transformation during erect and 'bent-hip, bent-knee' walking by humans with implications for the evolution of bipedalism. J Hum Evol 2003;44:563-79.
- Abstract: We have previously reported that predictive dynamic modeling suggests that the 'bent-hip, bent-knee' gait, which some attribute to Australopithecus afarensis AL-288-1, would have been much more expensive in mechanical terms for this hominid than an upright gait. Normal walking by modern adult humans owes much of its efficiency to conservation of energy by transformation between its potential and kinetic states. These findings suggest the question if, and to what extent, energy transformation exists in 'bent-hip, bent-knee' gait.This study calculates energy transformation in humans walking upright, at three different speeds, and walking 'bent-hip, bent-knee'. Kinematic data were gathered from video sequences and kinetic (ground reaction force) data from synchronous forceplate measurement. Applying Newtonian mechanics to our experimental data, the fluctuations of kinetic and potential energy in the body centre of mass were obtained and the effects of energy transformation evaluated and compared. In erect walking the fluctuations of two forms of energy are indeed largely out-of-phase, so that energy transformation occurs and total energy is conserved. In 'bent-hip, bent-knee' walking, however, the fluctuations of the kinetic and potential energy are much more in-phase, so that energy transformation occurs to a much lesser extent. Among all modes of walking the highest energy recovery is obtained in subjectively 'comfortable' walking, the next highest in subjectively 'fast' or 'slow' walking, and the least lowest in 'bent-hip, bent-knee' walking. The results imply that if 'bent-hip, bent-knee' gait was indeed habitually practiced by early bipedal hominids, a very substantial (and in our view as yet unidentified) selective advantage would have had to accrue, to offset the selective disadvantages of 'bent-hip, bent-knee' gait in terms of energy transformation.

[1] Begun DR, Richmond BG, Strait DS. Comment on "Origin of human bipedalism as an adaptation for locomotion on flexible branches". Science 2007;318:1066.
Abstract: 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.

[2] Abstract: 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.

[2] Bramble DM, Lieberman DE. Endurance running and the evolution of Homo. Nature 2004;432:345-52.
Abstract: Striding bipedalism is a key derived behaviour of hominids that possibly originated soon after the divergence of the chimpanzee and human lineages. Although bipedal gaits include walking and running, running is generally considered to have played no major role in human evolution because humans, like apes, are poor sprinters compared to most quadrupeds. Here we assess how well humans perform at sustained long-distance running, and review the physiological and anatomical bases of endurance running capabilities in humans and other mammals. Judged by several criteria, humans perform remarkably well at endurance running, thanks to a diverse array of features, many of which leave traces in the skeleton. The fossil evidence of these features suggests that endurance running is a derived capability of the genus Homo, originating about 2 million years ago, and may have been instrumental in the evolution of the human body form.

[4] Abstract: Striding bipedalism is a key derived behaviour of hominids that possibly originated soon after the divergence of the chimpanzee and human lineages. Although bipedal gaits include walking and running, running is generally considered to have played no major role in human evolution because humans, like apes, are poor sprinters compared to most quadrupeds. Here we assess how well humans perform at sustained long-distance running, and review the physiological and anatomical bases of endurance running capabilities in humans and other mammals. Judged by several criteria, humans perform remarkably well at endurance running, thanks to a diverse array of features, many of which leave traces in the skeleton. The fossil evidence of these features suggests that endurance running is a derived capability of the genus Homo, originating about 2 million years ago, and may have been instrumental in the evolution of the human body form.

[3] Fitzpatrick RC, Butler JE, Day BL. Resolving head rotation for human bipedalism. Curr Biol 2006;16:1509-14.
Abstract: 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.

[6] Abstract: 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.

[4] Kuliukas A. Wading for food the driving force of the evolution of bipedalism? Nutr Health 2002;16:267-89.
Abstract: Evidence is accumulating that suggests that the large human brain is most likely to have evolved in littoral and estuarine habitats rich in naturally occurring essential fatty acids. This paper adds further weight to this view, suggesting that another key human trait, our bipedality might also be best explained as an adaptation to a water-side niche. Evidence is provided here that extant apes, although preferring to keep dry, go into water when driven to do so by hunger. The anecdotal evidence has suggested that they tend to do this bipedally. Here, a new empirical study of captive bonobos found them to exhibit 2% or less bipedality on the ground or in trees but over 90% when wading in water to collect food. The skeletal morphology of AL 288-1 ("Lucy") is shown to indicate a strong ability to abduct and adduct the femur. These traits, together with a remarkably platypelloid pelvis, have not yet been adequately explained by terrestrial or arboreal models for early bipedalism but are consistent with those expected in an ape that adopted a specialist side-to-side 'ice-skating' or sideways wading mode. It is argued that this explanation of A. afarensis locomotor morphology is more parsimonious than others which have plainly failed to produce a consensus. Microwear evidence of Australopithecus dentition is also presented as evidence that the drive for such a wading form of locomotion might well have been waterside foods. This model obtains further support from the paleo-habitats of the earliest known bipeds, which are consistent with the hypothesis that wading contributed to the adaptive pressure towards bipedality.

[8] Abstract: Evidence is accumulating that suggests that the large human brain is most likely to have evolved in littoral and estuarine habitats rich in naturally occurring essential fatty acids. This paper adds further weight to this view, suggesting that another key human trait, our bipedality might also be best explained as an adaptation to a water-side niche. Evidence is provided here that extant apes, although preferring to keep dry, go into water when driven to do so by hunger. The anecdotal evidence has suggested that they tend to do this bipedally. Here, a new empirical study of captive bonobos found them to exhibit 2% or less bipedality on the ground or in trees but over 90% when wading in water to collect food. The skeletal morphology of AL 288-1 ("Lucy") is shown to indicate a strong ability to abduct and adduct the femur. These traits, together with a remarkably platypelloid pelvis, have not yet been adequately explained by terrestrial or arboreal models for early bipedalism but are consistent with those expected in an ape that adopted a specialist side-to-side 'ice-skating' or sideways wading mode. It is argued that this explanation of A. afarensis locomotor morphology is more parsimonious than others which have plainly failed to produce a consensus. Microwear evidence of Australopithecus dentition is also presented as evidence that the drive for such a wading form of locomotion might well have been waterside foods. This model obtains further support from the paleo-habitats of the earliest known bipeds, which are consistent with the hypothesis that wading contributed to the adaptive pressure towards bipedality.

[5] Nakatsukasa M. Acquisition of bipedalism: the Miocene hominoid record and modern analogues for bipedal protohominids. J Anat 2004;204:385-402.
Abstract: The well-known fossil hominoid Proconsul from the Early Miocene of Kenya was a non-specialized arboreal quadruped with strong pollicial/hallucial assisted grasping capability. It lacked most of the suspensory specializations acquired in living hominoids. Nacholapithecus, however, from the Middle Miocene of Kenya, although in part sharing with Proconsul the common primitive anatomical body design, was more specialized for orthograde climbing, 'hoisting' and bridging, with the glenoid fossae of the scapula probably being cranially orientated, the forelimbs proportionally large, and very long toes. Its tail loss suggests relatively slow movement, although tail loss may already have occurred in Proconsul. Nacholapithecus-like positional behaviour might thus have been a basis for development of more suspensory specialized positional behaviour in later hominoids. Unfortunately, after 13 Ma, there is a gap in the hominoid postcranial record in Africa until 6 Ma. Due to this gap, a scenario for later locomotor evolution prior to the divergence of Homo and Pan cannot be determined with certainty. The time gap also causes difficulties when we seek to determine polarities of morphological traits in very early hominids. Interpretation of the form-function relationships of postcranial features in incipient hominids will be difficult because it is predicted that they had incorporated bipedalism only moderately into their total positional repertoires. However, Japanese macaques, which are trained in traditional bipedal performance, may provide useful hints about bipedal adaptation in the protohominids. Kinematic analyses revealed that these macaques walked bipedally with a longer stride and lower stride frequency than used by ordinary macaques, owing to a more extended posture of the hindlimb joints. The body centre of gravity rises during the single-support phase of stance. Energetic studies of locomotion in these bipedal macaques revealed that energetic expenditure was 20-30% higher in bipedalism than in quadrupedalism, regardless of walking velocity.

[10] Abstract: The well-known fossil hominoid Proconsul from the Early Miocene of Kenya was a non-specialized arboreal quadruped with strong pollicial/hallucial assisted grasping capability. It lacked most of the suspensory specializations acquired in living hominoids. Nacholapithecus, however, from the Middle Miocene of Kenya, although in part sharing with Proconsul the common primitive anatomical body design, was more specialized for orthograde climbing, 'hoisting' and bridging, with the glenoid fossae of the scapula probably being cranially orientated, the forelimbs proportionally large, and very long toes. Its tail loss suggests relatively slow movement, although tail loss may already have occurred in Proconsul. Nacholapithecus-like positional behaviour might thus have been a basis for development of more suspensory specialized positional behaviour in later hominoids. Unfortunately, after 13 Ma, there is a gap in the hominoid postcranial record in Africa until 6 Ma. Due to this gap, a scenario for later locomotor evolution prior to the divergence of Homo and Pan cannot be determined with certainty. The time gap also causes difficulties when we seek to determine polarities of morphological traits in very early hominids. Interpretation of the form-function relationships of postcranial features in incipient hominids will be difficult because it is predicted that they had incorporated bipedalism only moderately into their total positional repertoires. However, Japanese macaques, which are trained in traditional bipedal performance, may provide useful hints about bipedal adaptation in the protohominids. Kinematic analyses revealed that these macaques walked bipedally with a longer stride and lower stride frequency than used by ordinary macaques, owing to a more extended posture of the hindlimb joints. The body centre of gravity rises during the single-support phase of stance. Energetic studies of locomotion in these bipedal macaques revealed that energetic expenditure was 20-30% higher in bipedalism than in quadrupedalism, regardless of walking velocity.

[6] Niemitz C. A theory on the evolution of the habitual orthograde human bipedalism--the "Amphibische Generalistentheorie". Anthropol Anz 2002;60:3-66.
Abstract: The theory is formulated that ubiquitous scarcity of energy is one of the main motors of evolution. It is concluded that our primate ancestors never came down from the trees, but rather they have always been (semi-)terrestrial. This habit is probably an old symplesiomorph trait, older than primates themselves. Terrestrial habits in primates correlate to body weight in small systematic groups (e.g., large genera, families) but are, overall, completely independent from individual body mass. An omnivorous, semiterrestrial quadrupedal locomotor generalist seems to be the most probable morpho- and eco-type for our ancestor at the threshold of a hominoid stage of our evolution. The theory presented here suggests that our hominoid ancestor lived in gallery forests and changed strata in order also to inhabit the savannah habitat as well as the shallow water of the rivers or coasts. Foraging in a wading manner was extremely favourable for an effective and, especially, seasonally independent, animal protein supply. Anatomical adaptations to orthogradism and proportions of the extremities are discussed in relation to the necessary and frequent change of habitat strata. Ultimately, human bipedalism is seen here to be derived as a consequence of the centre of body mass, which is, in primates, near the hind extremities. By contrast to other mammals entering the water, wading primates sink back on their hind limbs. Selective forces for habitat use, limb proportions and wading habits are discussed, as well as the phylogenetic origin of human affinity to water and shores in all peoples through all times, from australopithecine times through the Paleolithic until today.

[12] Abstract: The theory is formulated that ubiquitous scarcity of energy is one of the main motors of evolution. It is concluded that our primate ancestors never came down from the trees, but rather they have always been (semi-)terrestrial. This habit is probably an old symplesiomorph trait, older than primates themselves. Terrestrial habits in primates correlate to body weight in small systematic groups (e.g., large genera, families) but are, overall, completely independent from individual body mass. An omnivorous, semiterrestrial quadrupedal locomotor generalist seems to be the most probable morpho- and eco-type for our ancestor at the threshold of a hominoid stage of our evolution. The theory presented here suggests that our hominoid ancestor lived in gallery forests and changed strata in order also to inhabit the savannah habitat as well as the shallow water of the rivers or coasts. Foraging in a wading manner was extremely favourable for an effective and, especially, seasonally independent, animal protein supply. Anatomical adaptations to orthogradism and proportions of the extremities are discussed in relation to the necessary and frequent change of habitat strata. Ultimately, human bipedalism is seen here to be derived as a consequence of the centre of body mass, which is, in primates, near the hind extremities. By contrast to other mammals entering the water, wading primates sink back on their hind limbs. Selective forces for habitat use, limb proportions and wading habits are discussed, as well as the phylogenetic origin of human affinity to water and shores in all peoples through all times, from australopithecine times through the Paleolithic until today.

[7] Pickford M. Palaeoenvironments and hominoid evolution. Z Morphol Anthropol 2002;83:337-48.
Abstract: One of the key features that separates humans and their closest relatives (extinct species of the genus Homo and Praeanthropus and the australopithecines Australopithecus and Paranthropus) on the one hand, from the other hominoids, on the other, is their obligate bipedal locomotion when on the ground. This major difference from the generally quadrupedal locomotion practiced by other hominoids (Pan, Gorilla, Pongo and many extinct lineages) is reflected in many parts of the body, including all the major bones in the legs, arms, trunk and cranium. Locomotion has thus been of major interest to those interested in human origins, evolution, classification and phylogeny. A major hurdle to studies of the origins of bipedalism concerns the paucity of African hominoid fossils between 15 Ma, when all the adequately known hominoids were quadrupedal (most were pronograde, but at least one lineage was orthograde), and 4.2 Ma by which time fully bipedal hominids were established in Africa. Examination of Old World geology and palaeontology reveals a great deal about the evolution of palaeoenvironments and faunas during this period, and it is suggested that hominids evolved bipedal locomotion at the same time that there was a fundamental reorganisation of faunas towards the end of the Miocene. This faunal turnover resulted in the establishment of faunal lineages of "modern" aspect in Africa at the expense of "archaic" lineages which either went extinct or suffered a diminution of diversity. Many of the "modern" lineages were adapted to open country habitats in which grass became a major component of the diet as shown by modifications in the cheek teeth. Hominoids, in contrast, retained their traditional diet but were obliged to forage over greater and greater areas in order to do so, and this tactic led to pressures to modify the locomotor system rather than the diet. If bipedal hominids originated during this period, then the family Hominidae (sensu stricto) dates from about 8-7 Ma.

[14] Abstract: One of the key features that separates humans and their closest relatives (extinct species of the genus Homo and Praeanthropus and the australopithecines Australopithecus and Paranthropus) on the one hand, from the other hominoids, on the other, is their obligate bipedal locomotion when on the ground. This major difference from the generally quadrupedal locomotion practiced by other hominoids (Pan, Gorilla, Pongo and many extinct lineages) is reflected in many parts of the body, including all the major bones in the legs, arms, trunk and cranium. Locomotion has thus been of major interest to those interested in human origins, evolution, classification and phylogeny. A major hurdle to studies of the origins of bipedalism concerns the paucity of African hominoid fossils between 15 Ma, when all the adequately known hominoids were quadrupedal (most were pronograde, but at least one lineage was orthograde), and 4.2 Ma by which time fully bipedal hominids were established in Africa. Examination of Old World geology and palaeontology reveals a great deal about the evolution of palaeoenvironments and faunas during this period, and it is suggested that hominids evolved bipedal locomotion at the same time that there was a fundamental reorganisation of faunas towards the end of the Miocene. This faunal turnover resulted in the establishment of faunal lineages of "modern" aspect in Africa at the expense of "archaic" lineages which either went extinct or suffered a diminution of diversity. Many of the "modern" lineages were adapted to open country habitats in which grass became a major component of the diet as shown by modifications in the cheek teeth. Hominoids, in contrast, retained their traditional diet but were obliged to forage over greater and greater areas in order to do so, and this tactic led to pressures to modify the locomotor system rather than the diet. If bipedal hominids originated during this period, then the family Hominidae (sensu stricto) dates from about 8-7 Ma.

[8] Preuschoft H. Mechanisms for the acquisition of habitual bipedality: are there biomechanical reasons for the acquisition of upright bipedal posture? J Anat 2004;204:363-84.
Abstract: Morphology and biomechanics are linked by causal morphogenesis ('Wolff's law') and the interplay of mutations and selection (Darwin's 'survival of the fittest'). Thus shape-based selective pressures can be determined. In both cases we need to know which biomechanical factors lead to skeletal adaptation, and which ones exert selective pressures on body shape. Each bone must be able to sustain the greatest regularly occurring loads. Smaller loads are unlikely to lead to adaptation of morphology. The highest loads occur primarily in posture and locomotion, simply because of the effect of body weight (or its multiple). In the skull, however, it is biting and chewing that result in the greatest loads. Body shape adapted for an arboreal lifestyle also smooths the way towards bipedality. Hindlimb dominance, length of the limbs in relation to the axial skeleton, grasping hands and feet, mass distribution (especially of the limb segments), thoracic shape, rib curvatures, and the position of the centre of gravity are the adaptations to arboreality that also pre-adapt for bipedality. Five divergent locomotor/morphological types have evolved from this base: arm-swinging in gibbons, forelimb-dominated slow climbing in orangutans, quadrupedalism/climbing in the African apes, an unknown mix of climbing and bipedal walking in australopithecines, and the remarkably endurant bipedal walking of humans. All other apes are also facultative bipeds, but it is the biomechanical characteristics of bipedalism in orangutans, the most arboreal great ape, which is closest to that in humans. If not evolutionary accident, what selective factor can explain why two forms adopted bipedality? Most authors tend to connect bipedal locomotion with some aspect of progressively increasing distance between trees because of climatic changes. More precise factors, in accordance with biomechanical requirements, include stone-throwing, thermoregulation or wading in shallow water. Once bipedality has been acquired, development of typical human morphology can readily be explained as adaptations for energy saving over long distances. A paper in this volume shows that load-carrying ability was enhanced from australopithecines to Homo ergaster (early African H. erectus), supporting an earlier proposition that load-carrying was an essential factor in human evolution.

[16] Abstract: Morphology and biomechanics are linked by causal morphogenesis ('Wolff's law') and the interplay of mutations and selection (Darwin's 'survival of the fittest'). Thus shape-based selective pressures can be determined. In both cases we need to know which biomechanical factors lead to skeletal adaptation, and which ones exert selective pressures on body shape. Each bone must be able to sustain the greatest regularly occurring loads. Smaller loads are unlikely to lead to adaptation of morphology. The highest loads occur primarily in posture and locomotion, simply because of the effect of body weight (or its multiple). In the skull, however, it is biting and chewing that result in the greatest loads. Body shape adapted for an arboreal lifestyle also smooths the way towards bipedality. Hindlimb dominance, length of the limbs in relation to the axial skeleton, grasping hands and feet, mass distribution (especially of the limb segments), thoracic shape, rib curvatures, and the position of the centre of gravity are the adaptations to arboreality that also pre-adapt for bipedality. Five divergent locomotor/morphological types have evolved from this base: arm-swinging in gibbons, forelimb-dominated slow climbing in orangutans, quadrupedalism/climbing in the African apes, an unknown mix of climbing and bipedal walking in australopithecines, and the remarkably endurant bipedal walking of humans. All other apes are also facultative bipeds, but it is the biomechanical characteristics of bipedalism in orangutans, the most arboreal great ape, which is closest to that in humans. If not evolutionary accident, what selective factor can explain why two forms adopted bipedality? Most authors tend to connect bipedal locomotion with some aspect of progressively increasing distance between trees because of climatic changes. More precise factors, in accordance with biomechanical requirements, include stone-throwing, thermoregulation or wading in shallow water. Once bipedality has been acquired, development of typical human morphology can readily be explained as adaptations for energy saving over long distances. A paper in this volume shows that load-carrying ability was enhanced from australopithecines to Homo ergaster (early African H. erectus), supporting an earlier proposition that load-carrying was an essential factor in human evolution.

[9] Richmond BG, Begun DR, Strait DS. Origin of human bipedalism: The knuckle-walking hypothesis revisited. Am J Phys Anthropol 2001;Suppl 33:70-105.
Abstract: Some of the most long-standing questions in paleoanthropology concern how and why human bipedalism evolved. Over the last century, many hypotheses have been offered on the mode of locomotion from which bipedalism originated. Candidate ancestral adaptations include monkey-like arboreal or terrestrial quadrupedalism, gibbon- or orangutan-like (or other forms of) climbing and suspension, and knuckle-walking. This paper reviews the history of these hypotheses, outlines their predictions, and assesses them in light of current phylogenetic, comparative anatomical, and fossil evidence. The functional significance of characteristics of the shoulder and arm, elbow, wrist, and hand shared by African apes and humans, including their fossil relatives, most strongly supports the knuckle-walking hypothesis, which reconstructs the ancestor as being adapted to knuckle-walking and arboreal climbing. Future fossil discoveries, and a clear understanding of anthropoid locomotor anatomy, are required to ultimately test these hypotheses. If knuckle-walking was an important component of the behavioral repertoire of the prebipedal human ancestor, then we can reject scenarios on the origin of bipedalism that rely on a strictly arboreal ancestor. Moreover, paleoenvironmental data associated with the earliest hominins, and their close relatives, contradict hypotheses that place the agents of selection for bipedality in open savanna habitats. Existing hypotheses must explain why bipedalism would evolve from an ancestor that was already partly terrestrial. Many food acquisition and carrying hypotheses remain tenable in light of current evidence.

[18] Abstract: Some of the most long-standing questions in paleoanthropology concern how and why human bipedalism evolved. Over the last century, many hypotheses have been offered on the mode of locomotion from which bipedalism originated. Candidate ancestral adaptations include monkey-like arboreal or terrestrial quadrupedalism, gibbon- or orangutan-like (or other forms of) climbing and suspension, and knuckle-walking. This paper reviews the history of these hypotheses, outlines their predictions, and assesses them in light of current phylogenetic, comparative anatomical, and fossil evidence. The functional significance of characteristics of the shoulder and arm, elbow, wrist, and hand shared by African apes and humans, including their fossil relatives, most strongly supports the knuckle-walking hypothesis, which reconstructs the ancestor as being adapted to knuckle-walking and arboreal climbing. Future fossil discoveries, and a clear understanding of anthropoid locomotor anatomy, are required to ultimately test these hypotheses. If knuckle-walking was an important component of the behavioral repertoire of the prebipedal human ancestor, then we can reject scenarios on the origin of bipedalism that rely on a strictly arboreal ancestor. Moreover, paleoenvironmental data associated with the earliest hominins, and their close relatives, contradict hypotheses that place the agents of selection for bipedality in open savanna habitats. Existing hypotheses must explain why bipedalism would evolve from an ancestor that was already partly terrestrial. Many food acquisition and carrying hypotheses remain tenable in light of current evidence.

[10] Richmond BG, Jungers WL. Orrorin tugenensis femoral morphology and the evolution of hominin bipedalism. Science 2008;319:1662-5.
Abstract: Bipedalism is a key human adaptation and a defining feature of the hominin clade. Fossil femora discovered in Kenya and attributed to Orrorin tugenensis, at 6 million years ago, purportedly provide the earliest postcranial evidence of hominin bipedalism, but their functional and phylogenetic affinities are controversial. We show that the O. tugenensis femur differs from those of apes and Homo and most strongly resembles those of Australopithecus and Paranthropus, indicating that O. tugenensis was bipedal but is not more closely related to Homo than to Australopithecus. Femoral morphology indicates that O. tugenensis shared distinctive hip biomechanics with australopiths, suggesting that this complex evolved early in human evolution and persisted for almost 4 million years until modifications of the hip appeared in the late Pliocene in early Homo.

[20] Abstract: Bipedalism is a key human adaptation and a defining feature of the hominin clade. Fossil femora discovered in Kenya and attributed to Orrorin tugenensis, at 6 million years ago, purportedly provide the earliest postcranial evidence of hominin bipedalism, but their functional and phylogenetic affinities are controversial. We show that the O. tugenensis femur differs from those of apes and Homo and most strongly resembles those of Australopithecus and Paranthropus, indicating that O. tugenensis was bipedal but is not more closely related to Homo than to Australopithecus. Femoral morphology indicates that O. tugenensis shared distinctive hip biomechanics with australopiths, suggesting that this complex evolved early in human evolution and persisted for almost 4 million years until modifications of the hip appeared in the late Pliocene in early Homo.

[11] Schmitt D. Insights into the evolution of human bipedalism from experimental studies of humans and other primates. J Exp Biol 2003;206:1437-48.
Abstract: An understanding of the evolution of human bipedalism can provide valuable insights into the biomechanical and physiological characteristics of locomotion in modern humans. The walking gaits of humans, other bipeds and most quadrupedal mammals can best be described by using an inverted-pendulum model, in which there is minimal change in flexion of the limb joints during stance phase. As a result, it seems logical that the evolution of bipedalism in humans involved a simple transition from a relatively stiff-legged quadrupedalism in a terrestrial ancestor to relatively stiff-legged bipedalism in early humans. However, experimental studies of locomotion in humans and nonhuman primates have shown that the evolution of bipedalism involved a much more complex series of transitions, originating with a relatively compliant form of quadrupedalism. These studies show that relatively compliant walking gaits allow primates to achieve fast walking speeds using long strides, low stride frequencies, relatively low peak vertical forces, and relatively high impact shock attenuation ratios. A relatively compliant, ape-like bipedal walking style is consistent with the anatomy of early hominids and may have been an effective gait for a small biped with relatively small and less stabilized joints, which had not yet completely forsaken arboreal locomotion. Laboratory-based studies of primates also suggest that human bipedalism arose not from a terrestrial ancestor but rather from a climbing, arboreal forerunner. Experimental data, in conjunction with anatomical data on early human ancestors, show clearly that a relatively stiff modern human gait and associated physiological and anatomical adaptations are not primitive retentions from a primate ancestor, but are instead recently acquired characters of our genus.

[22] Abstract: An understanding of the evolution of human bipedalism can provide valuable insights into the biomechanical and physiological characteristics of locomotion in modern humans. The walking gaits of humans, other bipeds and most quadrupedal mammals can best be described by using an inverted-pendulum model, in which there is minimal change in flexion of the limb joints during stance phase. As a result, it seems logical that the evolution of bipedalism in humans involved a simple transition from a relatively stiff-legged quadrupedalism in a terrestrial ancestor to relatively stiff-legged bipedalism in early humans. However, experimental studies of locomotion in humans and nonhuman primates have shown that the evolution of bipedalism involved a much more complex series of transitions, originating with a relatively compliant form of quadrupedalism. These studies show that relatively compliant walking gaits allow primates to achieve fast walking speeds using long strides, low stride frequencies, relatively low peak vertical forces, and relatively high impact shock attenuation ratios. A relatively compliant, ape-like bipedal walking style is consistent with the anatomy of early hominids and may have been an effective gait for a small biped with relatively small and less stabilized joints, which had not yet completely forsaken arboreal locomotion. Laboratory-based studies of primates also suggest that human bipedalism arose not from a terrestrial ancestor but rather from a climbing, arboreal forerunner. Experimental data, in conjunction with anatomical data on early human ancestors, show clearly that a relatively stiff modern human gait and associated physiological and anatomical adaptations are not primitive retentions from a primate ancestor, but are instead recently acquired characters of our genus.

[12] Schwartz JH. The origins of human bipedalism. Science 2007;318:1065.

[13] Sellers WI, Dennis LA, Crompton RH. Predicting the metabolic energy costs of bipedalism using evolutionary robotics. J Exp Biol 2003;206:1127-36.
Abstract: To understand the evolution of bipedalism among the hominoids in an ecological context we need to be able to estimate the energetic cost of locomotion in fossil forms. Ideally such an estimate would be based entirely on morphology since, except for the rare instances where footprints are preserved, this is the only primary source of evidence available. In this paper we use evolutionary robotics techniques (genetic algorithms, pattern generators and mechanical modeling) to produce a biomimetic simulation of bipedalism based on human body dimensions. The mechanical simulation is a seven-segment, two-dimensional model with motive force provided by tension generators representing the major muscle groups acting around the lower-limb joints. Metabolic energy costs are calculated from the muscle model, and bipedal gait is generated using a finite-state pattern generator whose parameters are produced using a genetic algorithm with locomotor economy (maximum distance for a fixed energy cost) as the fitness criterion. The model is validated by comparing the values it generates with those for modern humans. The result (maximum efficiency of 200 J m(-1)) is within 15% of the experimentally derived value, which is very encouraging and suggests that this is a useful analytic technique for investigating the locomotor behaviour of fossil forms. Initial work suggests that in the future this technique could be used to estimate other locomotor parameters such as top speed. In addition, the animations produced by this technique are qualitatively very convincing, which suggests that this may also be a useful technique for visualizing bipedal locomotion.

[25] Abstract: To understand the evolution of bipedalism among the hominoids in an ecological context we need to be able to estimate the energetic cost of locomotion in fossil forms. Ideally such an estimate would be based entirely on morphology since, except for the rare instances where footprints are preserved, this is the only primary source of evidence available. In this paper we use evolutionary robotics techniques (genetic algorithms, pattern generators and mechanical modeling) to produce a biomimetic simulation of bipedalism based on human body dimensions. The mechanical simulation is a seven-segment, two-dimensional model with motive force provided by tension generators representing the major muscle groups acting around the lower-limb joints. Metabolic energy costs are calculated from the muscle model, and bipedal gait is generated using a finite-state pattern generator whose parameters are produced using a genetic algorithm with locomotor economy (maximum distance for a fixed energy cost) as the fitness criterion. The model is validated by comparing the values it generates with those for modern humans. The result (maximum efficiency of 200 J m(-1)) is within 15% of the experimentally derived value, which is very encouraging and suggests that this is a useful analytic technique for investigating the locomotor behaviour of fossil forms. Initial work suggests that in the future this technique could be used to estimate other locomotor parameters such as top speed. In addition, the animations produced by this technique are qualitatively very convincing, which suggests that this may also be a useful technique for visualizing bipedal locomotion.

[14] Skoyles JR. Human balance, the evolution of bipedalism and dysequilibrium syndrome. Med Hypotheses 2006;66:1060-8.
Abstract: 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.

[27] Abstract: 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.

[15] Sockol MD, Raichlen DA, Pontzer H. Chimpanzee locomotor energetics and the origin of human bipedalism. Proc Natl Acad Sci U S A 2007;104:12265-9.
Abstract: 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.

[29] Abstract: 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.

[16] Spoor F, Wood B, Zonneveld F. Implications of early hominid labyrinthine morphology for evolution of human bipedal locomotion. Nature 1994;369:645-8.
Abstract: The upright posture and obligatory bipedalism of modern humans are unique among living primates. The evolutionary history of this behaviour has traditionally been pursued by functional analysis of the postcranial skeleton and the preserved footprint trails of fossil hominids. Here we report a systematic attempt to reconstruct the locomotor behaviour of early hominids by looking at a major component of the mechanism for the unconscious perception of movement, namely by examining the vestibular system of living primates and early hominids. High-resolution computed tomography was used to generate cross-sectional images of the bony labyrinth. Among the fossil hominids the earliest species to demonstrate the modern human morphology is Homo erectus. In contrast, the semicircular canal dimensions in crania from southern Africa attributed to Australopithecus and Paranthropus resemble those of the extant great apes. Among early Homo specimens, the canal dimensions of Stw 53 are unlike those seen in any of the hominids or great apes, whereas those of SK 847 are modern-human-like.

[31] Abstract: The upright posture and obligatory bipedalism of modern humans are unique among living primates. The evolutionary history of this behaviour has traditionally been pursued by functional analysis of the postcranial skeleton and the preserved footprint trails of fossil hominids. Here we report a systematic attempt to reconstruct the locomotor behaviour of early hominids by looking at a major component of the mechanism for the unconscious perception of movement, namely by examining the vestibular system of living primates and early hominids. High-resolution computed tomography was used to generate cross-sectional images of the bony labyrinth. Among the fossil hominids the earliest species to demonstrate the modern human morphology is Homo erectus. In contrast, the semicircular canal dimensions in crania from southern Africa attributed to Australopithecus and Paranthropus resemble those of the extant great apes. Among early Homo specimens, the canal dimensions of Stw 53 are unlike those seen in any of the hominids or great apes, whereas those of SK 847 are modern-human-like.

[17] Stanford CB. Arboreal bipedalism in wild chimpanzees: implications for the evolution of hominid posture and locomotion. Am J Phys Anthropol 2006;129:225-31.
Abstract: Field observations of bipedal posture and locomotion in wild chimpanzees (Pan troglodytes) can serve as key evidence for reconstructing the likely origins of bipedalism in the last prehominid human ancestor. This paper reports on a sample of bipedal bouts, recorded ad libitum, in wild chimpanzees in Bwindi Impenetrable National Park in southwestern Uganda. The Ruhija community of chimpanzees in Bwindi displays a high rate of bipedal posture. In 246.7 hr of observation from 2001-2003, 179 instances of bipedal posture lasting 5 sec or longer were recorded, for a rate of 0.73 bouts per observation hour. Bipedalism was observed only on arboreal substrates, and was almost all postural, and not locomotor. Bipedalism was part of a complex series of positional behaviors related to feeding, which included two-legged standing, one-legged standing with arm support, and other intermediate postures. Ninety-six percent of bipedal bouts occurred in a foraging context, always as a chimpanzee reached to pluck fruit from tree limbs. Bipedalism was seen in both male and female adults, less frequently among juveniles, and rarely in infants. Both the frequency and duration of bipedal bouts showed a significant positive correlation with estimated substrate diameter. Neither fruit size nor nearest-neighbor association patterns were significantly correlated with the occurrence of bipedalism. Bipedalism is seen frequently in the Bwindi chimpanzee community, in part because of the unusual observer conditions at Bwindi. Most observations of bipedalism were made when the animals were in treetops and the observer at eye-level across narrow ravines. This suggests that wild chimpanzees may engage in bipedal behavior more often than is generally appreciated. Models of the likely evolutionary origins of bipedalism are considered in the light of Bwindi bipedalism data. Bipedalism among Bwindi chimpanzees suggests the origin of bipedal posture in hominids to be related to foraging advantages in fruit trees. It suggests important arboreal advantages in upright posture. The origin of postural bipedalism may have preceded and been causally disconnected from locomotor bipedalism.

[33] Abstract: Field observations of bipedal posture and locomotion in wild chimpanzees (Pan troglodytes) can serve as key evidence for reconstructing the likely origins of bipedalism in the last prehominid human ancestor. This paper reports on a sample of bipedal bouts, recorded ad libitum, in wild chimpanzees in Bwindi Impenetrable National Park in southwestern Uganda. The Ruhija community of chimpanzees in Bwindi displays a high rate of bipedal posture. In 246.7 hr of observation from 2001-2003, 179 instances of bipedal posture lasting 5 sec or longer were recorded, for a rate of 0.73 bouts per observation hour. Bipedalism was observed only on arboreal substrates, and was almost all postural, and not locomotor. Bipedalism was part of a complex series of positional behaviors related to feeding, which included two-legged standing, one-legged standing with arm support, and other intermediate postures. Ninety-six percent of bipedal bouts occurred in a foraging context, always as a chimpanzee reached to pluck fruit from tree limbs. Bipedalism was seen in both male and female adults, less frequently among juveniles, and rarely in infants. Both the frequency and duration of bipedal bouts showed a significant positive correlation with estimated substrate diameter. Neither fruit size nor nearest-neighbor association patterns were significantly correlated with the occurrence of bipedalism. Bipedalism is seen frequently in the Bwindi chimpanzee community, in part because of the unusual observer conditions at Bwindi. Most observations of bipedalism were made when the animals were in treetops and the observer at eye-level across narrow ravines. This suggests that wild chimpanzees may engage in bipedal behavior more often than is generally appreciated. Models of the likely evolutionary origins of bipedalism are considered in the light of Bwindi bipedalism data. Bipedalism among Bwindi chimpanzees suggests the origin of bipedal posture in hominids to be related to foraging advantages in fruit trees. It suggests important arboreal advantages in upright posture. The origin of postural bipedalism may have preceded and been causally disconnected from locomotor bipedalism.

[18] Steudel-Numbers KL. Role of locomotor economy in the origin of bipedal posture and gait. Am J Phys Anthropol 2001;116:171-3.

[19] Sylvester AD. Locomotor decoupling and the origin of hominin bipedalism. J Theor Biol 2006;242:581-90.
Abstract: Theoretical adaptive landscapes and mathematical representations of key constraints of evolutionary and primate biology are used to propose a new hypothesis for the origin of hominin bipedalism. These constraints suggest that the selective pressure that produced this novel form of locomotion was the need for effective suspensory and terrestrial movement. This testable hypothesis, termed the Decoupling Hypothesis, posits that bipedalism is an adaptation that enables the shoulder to maintain a high degree of mobility, a feature important to suspensory behaviors, in the face of significant demands for a high degree of stability, a feature important for highly effective terrestrial quadrupedism.

[36] Abstract: Theoretical adaptive landscapes and mathematical representations of key constraints of evolutionary and primate biology are used to propose a new hypothesis for the origin of hominin bipedalism. These constraints suggest that the selective pressure that produced this novel form of locomotion was the need for effective suspensory and terrestrial movement. This testable hypothesis, termed the Decoupling Hypothesis, posits that bipedalism is an adaptation that enables the shoulder to maintain a high degree of mobility, a feature important to suspensory behaviors, in the face of significant demands for a high degree of stability, a feature important for highly effective terrestrial quadrupedism.

[20] Thorpe SK, Holder RL, Crompton RH. Origin of human bipedalism as an adaptation for locomotion on flexible branches. Science 2007;316:1328-31.
Abstract: 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.

[38] Abstract: 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.

[21] Vaughan CL. Theories of bipedal walking: an odyssey. J Biomech 2003;36:513-23. Abstract: In this paper six theories of bipedal walking, and the evidence in support of the theories, are reviewed. They include: evolution, minimising energy consumption, maturation in children, central pattern generators, linking control and effect, and robots on two legs. Specifically, the six theories posit that: (1) bipedalism is the fundamental evolutionary adaptation that sets hominids--and therefore humans--apart from other primates; (2) locomotion is the translation of the centre of gravity along a pathway requiring the least expenditure of energy; (3) when a young child takes its first few halting steps, his or her biomechanical strategy is to minimise the risk of falling; (4) a dedicated network of interneurons in the spinal cord generates the rhythm and cyclic pattern of electromyographic signals that give rise to bipedal gait; (5) bipedal locomotion is generated through global entrainment of the neural system on the one hand, and the musculoskeletal system plus environment on the other; and (6) powered dynamic gait in a bipedal robot can be realised only through a strategy which is based on stability and real-time feedback control. The published record suggests that each of the theories has some measure of support. However, it is important to note that there are other important theories of locomotion which have not been covered in this review. Despite such omissions, this odyssey has explored the wide spectrum of bipedal walking, from its origins through to the integration of the nervous, muscular and skeletal systems.

[22] Wang WJ, Crompton RH, Li Y, Gunther MM. Energy transformation during erect and 'bent-hip, bent-knee' walking by humans with implications for the evolution of bipedalism. J Hum Evol 2003;44:563-79.
Abstract: We have previously reported that predictive dynamic modeling suggests that the 'bent-hip, bent-knee' gait, which some attribute to Australopithecus afarensis AL-288-1, would have been much more expensive in mechanical terms for this hominid than an upright gait. Normal walking by modern adult humans owes much of its efficiency to conservation of energy by transformation between its potential and kinetic states. These findings suggest the question if, and to what extent, energy transformation exists in 'bent-hip, bent-knee' gait.This study calculates energy transformation in humans walking upright, at three different speeds, and walking 'bent-hip, bent-knee'. Kinematic data were gathered from video sequences and kinetic (ground reaction force) data from synchronous forceplate measurement. Applying Newtonian mechanics to our experimental data, the fluctuations of kinetic and potential energy in the body centre of mass were obtained and the effects of energy transformation evaluated and compared. In erect walking the fluctuations of two forms of energy are indeed largely out-of-phase, so that energy transformation occurs and total energy is conserved. In 'bent-hip, bent-knee' walking, however, the fluctuations of the kinetic and potential energy are much more in-phase, so that energy transformation occurs to a much lesser extent. Among all modes of walking the highest energy recovery is obtained in subjectively 'comfortable' walking, the next highest in subjectively 'fast' or 'slow' walking, and the least lowest in 'bent-hip, bent-knee' walking. The results imply that if 'bent-hip, bent-knee' gait was indeed habitually practiced by early bipedal hominids, a very substantial (and in our view as yet unidentified) selective advantage would have had to accrue, to offset the selective disadvantages of 'bent-hip, bent-knee' gait in terms of energy transformation.

[41] Abstract: We have previously reported that predictive dynamic modeling suggests that the 'bent-hip, bent-knee' gait, which some attribute to Australopithecus afarensis AL-288-1, would have been much more expensive in mechanical terms for this hominid than an upright gait. Normal walking by modern adult humans owes much of its efficiency to conservation of energy by transformation between its potential and kinetic states. These findings suggest the question if, and to what extent, energy transformation exists in 'bent-hip, bent-knee' gait.This study calculates energy transformation in humans walking upright, at three different speeds, and walking 'bent-hip, bent-knee'. Kinematic data were gathered from video sequences and kinetic (ground reaction force) data from synchronous forceplate measurement. Applying Newtonian mechanics to our experimental data, the fluctuations of kinetic and potential energy in the body centre of mass were obtained and the effects of energy transformation evaluated and compared. In erect walking the fluctuations of two forms of energy are indeed largely out-of-phase, so that energy transformation occurs and total energy is conserved. In 'bent-hip, bent-knee' walking, however, the fluctuations of the kinetic and potential energy are much more in-phase, so that energy transformation occurs to a much lesser extent. Among all modes of walking the highest energy recovery is obtained in subjectively 'comfortable' walking, the next highest in subjectively 'fast' or 'slow' walking, and the least lowest in 'bent-hip, bent-knee' walking. The results imply that if 'bent-hip, bent-knee' gait was indeed habitually practiced by early bipedal hominids, a very substantial (and in our view as yet unidentified) selective advantage would have had to accrue, to offset the selective disadvantages of 'bent-hip, bent-knee' gait in terms of energy transformation.

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+== Re: Evolution of bipedalism ==
+Consider:
+*Comment on "Origin of human bipedalism as an adaptation for locomotion on flexible branches".<ref>	Begun DR, Richmond BG, Strait DS. Comment on "Origin of human bipedalism as an adaptation for locomotion on flexible branches". Science 2007;318:1066.
+:*Abstract: 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.</ref>
+*Endurance running and the evolution of Homo<ref>Bramble DM, Lieberman DE. Endurance running and the evolution of Homo. Nature 2004;432:345-52.
+:*Abstract: Striding bipedalism is a key derived behaviour of hominids that possibly originated soon after the divergence of the chimpanzee and human lineages. Although bipedal gaits include walking and running, running is generally considered to have played no major role in human evolution because humans, like apes, are poor sprinters compared to most quadrupeds. Here we assess how well humans perform at sustained long-distance running, and review the physiological and anatomical bases of endurance running capabilities in humans and other mammals. Judged by several criteria, humans perform remarkably well at endurance running, thanks to a diverse array of features, many of which leave traces in the skeleton. The fossil evidence of these features suggests that endurance running is a derived capability of the genus Homo, originating about 2 million years ago, and may have been instrumental in the evolution of the human body form.</ref>
+*Resolving head rotation for human bipedalism<ref>	Fitzpatrick RC, Butler JE, Day BL. Resolving head rotation for human bipedalism. Curr Biol 2006;16:1509-14.
+:*Abstract: 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.</ref>
+<ref>	Kuliukas A. Wading for food the driving force of the evolution of bipedalism? Nutr Health 2002;16:267-89.
+:*Abstract: Evidence is accumulating that suggests that the large human brain is most likely to have evolved in littoral and estuarine habitats rich in naturally occurring essential fatty acids. This paper adds further weight to this view, suggesting that another key human trait, our bipedality might also be best explained as an adaptation to a water-side niche. Evidence is provided here that extant apes, although preferring to keep dry, go into water when driven to do so by hunger. The anecdotal evidence has suggested that they tend to do this bipedally. Here, a new empirical study of captive bonobos found them to exhibit 2% or less bipedality on the ground or in trees but over 90% when wading in water to collect food. The skeletal morphology of AL 288-1 ("Lucy") is shown to indicate a strong ability to abduct and adduct the femur. These traits, together with a remarkably platypelloid pelvis, have not yet been adequately explained by terrestrial or arboreal models for early bipedalism but are consistent with those expected in an ape that adopted a specialist side-to-side 'ice-skating' or sideways wading mode. It is argued that this explanation of A. afarensis locomotor morphology is more parsimonious than others which have plainly failed to produce a consensus. Microwear evidence of Australopithecus dentition is also presented as evidence that the drive for such a wading form of locomotion might well have been waterside foods. This model obtains further support from the paleo-habitats of the earliest known bipeds, which are consistent with the hypothesis that wading contributed to the adaptive pressure towards bipedality.</ref>
+<ref>	Nakatsukasa M. Acquisition of bipedalism: the Miocene hominoid record and modern analogues for bipedal protohominids. J Anat 2004;204:385-402.
+:*Abstract: The well-known fossil hominoid Proconsul from the Early Miocene of Kenya was a non-specialized arboreal quadruped with strong pollicial/hallucial assisted grasping capability. It lacked most of the suspensory specializations acquired in living hominoids. Nacholapithecus, however, from the Middle Miocene of Kenya, although in part sharing with Proconsul the common primitive anatomical body design, was more specialized for orthograde climbing, 'hoisting' and bridging, with the glenoid fossae of the scapula probably being cranially orientated, the forelimbs proportionally large, and very long toes. Its tail loss suggests relatively slow movement, although tail loss may already have occurred in Proconsul. Nacholapithecus-like positional behaviour might thus have been a basis for development of more suspensory specialized positional behaviour in later hominoids. Unfortunately, after 13 Ma, there is a gap in the hominoid postcranial record in Africa until 6 Ma. Due to this gap, a scenario for later locomotor evolution prior to the divergence of Homo and Pan cannot be determined with certainty. The time gap also causes difficulties when we seek to determine polarities of morphological traits in very early hominids. Interpretation of the form-function relationships of postcranial features in incipient hominids will be difficult because it is predicted that they had incorporated bipedalism only moderately into their total positional repertoires. However, Japanese macaques, which are trained in traditional bipedal performance, may provide useful hints about bipedal adaptation in the protohominids. Kinematic analyses revealed that these macaques walked bipedally with a longer stride and lower stride frequency than used by ordinary macaques, owing to a more extended posture of the hindlimb joints. The body centre of gravity rises during the single-support phase of stance. Energetic studies of locomotion in these bipedal macaques revealed that energetic expenditure was 20-30% higher in bipedalism than in quadrupedalism, regardless of walking velocity.</ref>
+<ref>	Niemitz C. A theory on the evolution of the habitual orthograde human bipedalism--the "Amphibische Generalistentheorie". Anthropol Anz 2002;60:3-66.
+:*Abstract: The theory is formulated that ubiquitous scarcity of energy is one of the main motors of evolution. It is concluded that our primate ancestors never came down from the trees, but rather they have always been (semi-)terrestrial. This habit is probably an old symplesiomorph trait, older than primates themselves. Terrestrial habits in primates correlate to body weight in small systematic groups (e.g., large genera, families) but are, overall, completely independent from individual body mass. An omnivorous, semiterrestrial quadrupedal locomotor generalist seems to be the most probable morpho- and eco-type for our ancestor at the threshold of a hominoid stage of our evolution. The theory presented here suggests that our hominoid ancestor lived in gallery forests and changed strata in order also to inhabit the savannah habitat as well as the shallow water of the rivers or coasts. Foraging in a wading manner was extremely favourable for an effective and, especially, seasonally independent, animal protein supply. Anatomical adaptations to orthogradism and proportions of the extremities are discussed in relation to the necessary and frequent change of habitat strata. Ultimately, human bipedalism is seen here to be derived as a consequence of the centre of body mass, which is, in primates, near the hind extremities. By contrast to other mammals entering the water, wading primates sink back on their hind limbs. Selective forces for habitat use, limb proportions and wading habits are discussed, as well as the phylogenetic origin of human affinity to water and shores in all peoples through all times, from australopithecine times through the Paleolithic until today.</ref>
+<ref>	Pickford M. Palaeoenvironments and hominoid evolution. Z Morphol Anthropol 2002;83:337-48.
+:*Abstract: One of the key features that separates humans and their closest relatives (extinct species of the genus Homo and Praeanthropus and the australopithecines Australopithecus and Paranthropus) on the one hand, from the other hominoids, on the other, is their obligate bipedal locomotion when on the ground. This major difference from the generally quadrupedal locomotion practiced by other hominoids (Pan, Gorilla, Pongo and many extinct lineages) is reflected in many parts of the body, including all the major bones in the legs, arms, trunk and cranium. Locomotion has thus been of major interest to those interested in human origins, evolution, classification and phylogeny. A major hurdle to studies of the origins of bipedalism concerns the paucity of African hominoid fossils between 15 Ma, when all the adequately known hominoids were quadrupedal (most were pronograde, but at least one lineage was orthograde), and 4.2 Ma by which time fully bipedal hominids were established in Africa. Examination of Old World geology and palaeontology reveals a great deal about the evolution of palaeoenvironments and faunas during this period, and it is suggested that hominids evolved bipedal locomotion at the same time that there was a fundamental reorganisation of faunas towards the end of the Miocene. This faunal turnover resulted in the establishment of faunal lineages of "modern" aspect in Africa at the expense of "archaic" lineages which either went extinct or suffered a diminution of diversity. Many of the "modern" lineages were adapted to open country habitats in which grass became a major component of the diet as shown by modifications in the cheek teeth. Hominoids, in contrast, retained their traditional diet but were obliged to forage over greater and greater areas in order to do so, and this tactic led to pressures to modify the locomotor system rather than the diet. If bipedal hominids originated during this period, then the family Hominidae (sensu stricto) dates from about 8-7 Ma.</ref>
+<ref>	Preuschoft H. Mechanisms for the acquisition of habitual bipedality: are there biomechanical reasons for the acquisition of upright bipedal posture? J Anat 2004;204:363-84.
+:*Abstract: Morphology and biomechanics are linked by causal morphogenesis ('Wolff's law') and the interplay of mutations and selection (Darwin's 'survival of the fittest'). Thus shape-based selective pressures can be determined. In both cases we need to know which biomechanical factors lead to skeletal adaptation, and which ones exert selective pressures on body shape. Each bone must be able to sustain the greatest regularly occurring loads. Smaller loads are unlikely to lead to adaptation of morphology. The highest loads occur primarily in posture and locomotion, simply because of the effect of body weight (or its multiple). In the skull, however, it is biting and chewing that result in the greatest loads. Body shape adapted for an arboreal lifestyle also smooths the way towards bipedality. Hindlimb dominance, length of the limbs in relation to the axial skeleton, grasping hands and feet, mass distribution (especially of the limb segments), thoracic shape, rib curvatures, and the position of the centre of gravity are the adaptations to arboreality that also pre-adapt for bipedality. Five divergent locomotor/morphological types have evolved from this base: arm-swinging in gibbons, forelimb-dominated slow climbing in orangutans, quadrupedalism/climbing in the African apes, an unknown mix of climbing and bipedal walking in australopithecines, and the remarkably endurant bipedal walking of humans. All other apes are also facultative bipeds, but it is the biomechanical characteristics of bipedalism in orangutans, the most arboreal great ape, which is closest to that in humans. If not evolutionary accident, what selective factor can explain why two forms adopted bipedality? Most authors tend to connect bipedal locomotion with some aspect of progressively increasing distance between trees because of climatic changes. More precise factors, in accordance with biomechanical requirements, include stone-throwing, thermoregulation or wading in shallow water. Once bipedality has been acquired, development of typical human morphology can readily be explained as adaptations for energy saving over long distances. A paper in this volume shows that load-carrying ability was enhanced from australopithecines to Homo ergaster (early African H. erectus), supporting an earlier proposition that load-carrying was an essential factor in human evolution.</ref>
+<ref>	Richmond BG, Begun DR, Strait DS. Origin of human bipedalism: The knuckle-walking hypothesis revisited. Am J Phys Anthropol 2001;Suppl 33:70-105.
+:*Abstract: Some of the most long-standing questions in paleoanthropology concern how and why human bipedalism evolved. Over the last century, many hypotheses have been offered on the mode of locomotion from which bipedalism originated. Candidate ancestral adaptations include monkey-like arboreal or terrestrial quadrupedalism, gibbon- or orangutan-like (or other forms of) climbing and suspension, and knuckle-walking. This paper reviews the history of these hypotheses, outlines their predictions, and assesses them in light of current phylogenetic, comparative anatomical, and fossil evidence. The functional significance of characteristics of the shoulder and arm, elbow, wrist, and hand shared by African apes and humans, including their fossil relatives, most strongly supports the knuckle-walking hypothesis, which reconstructs the ancestor as being adapted to knuckle-walking and arboreal climbing. Future fossil discoveries, and a clear understanding of anthropoid locomotor anatomy, are required to ultimately test these hypotheses. If knuckle-walking was an important component of the behavioral repertoire of the prebipedal human ancestor, then we can reject scenarios on the origin of bipedalism that rely on a strictly arboreal ancestor. Moreover, paleoenvironmental data associated with the earliest hominins, and their close relatives, contradict hypotheses that place the agents of selection for bipedality in open savanna habitats. Existing hypotheses must explain why bipedalism would evolve from an ancestor that was already partly terrestrial. Many food acquisition and carrying hypotheses remain tenable in light of current evidence.</ref>
+<ref>	Richmond BG, Jungers WL. Orrorin tugenensis femoral morphology and the evolution of hominin bipedalism. Science 2008;319:1662-5.
+:*Abstract: Bipedalism is a key human adaptation and a defining feature of the hominin clade. Fossil femora discovered in Kenya and attributed to Orrorin tugenensis, at 6 million years ago, purportedly provide the earliest postcranial evidence of hominin bipedalism, but their functional and phylogenetic affinities are controversial. We show that the O. tugenensis femur differs from those of apes and Homo and most strongly resembles those of Australopithecus and Paranthropus, indicating that O. tugenensis was bipedal but is not more closely related to Homo than to Australopithecus. Femoral morphology indicates that O. tugenensis shared distinctive hip biomechanics with australopiths, suggesting that this complex evolved early in human evolution and persisted for almost 4 million years until modifications of the hip appeared in the late Pliocene in early Homo.</ref>
+<ref>	Schmitt D. Insights into the evolution of human bipedalism from experimental studies of humans and other primates. J Exp Biol 2003;206:1437-48.
+:*Abstract: An understanding of the evolution of human bipedalism can provide valuable insights into the biomechanical and physiological characteristics of locomotion in modern humans. The walking gaits of humans, other bipeds and most quadrupedal mammals can best be described by using an inverted-pendulum model, in which there is minimal change in flexion of the limb joints during stance phase. As a result, it seems logical that the evolution of bipedalism in humans involved a simple transition from a relatively stiff-legged quadrupedalism in a terrestrial ancestor to relatively stiff-legged bipedalism in early humans. However, experimental studies of locomotion in humans and nonhuman primates have shown that the evolution of bipedalism involved a much more complex series of transitions, originating with a relatively compliant form of quadrupedalism. These studies show that relatively compliant walking gaits allow primates to achieve fast walking speeds using long strides, low stride frequencies, relatively low peak vertical forces, and relatively high impact shock attenuation ratios. A relatively compliant, ape-like bipedal walking style is consistent with the anatomy of early hominids and may have been an effective gait for a small biped with relatively small and less stabilized joints, which had not yet completely forsaken arboreal locomotion. Laboratory-based studies of primates also suggest that human bipedalism arose not from a terrestrial ancestor but rather from a climbing, arboreal forerunner. Experimental data, in conjunction with anatomical data on early human ancestors, show clearly that a relatively stiff modern human gait and associated physiological and anatomical adaptations are not primitive retentions from a primate ancestor, but are instead recently acquired characters of our genus.</ref>
+<ref>	Schwartz JH. The origins of human bipedalism. Science 2007;318:1065.</ref>
+<ref>	Sellers WI, Dennis LA, Crompton RH. Predicting the metabolic energy costs of bipedalism using evolutionary robotics. J Exp Biol 2003;206:1127-36.
+:*Abstract: To understand the evolution of bipedalism among the hominoids in an ecological context we need to be able to estimate the energetic cost of locomotion in fossil forms. Ideally such an estimate would be based entirely on morphology since, except for the rare instances where footprints are preserved, this is the only primary source of evidence available. In this paper we use evolutionary robotics techniques (genetic algorithms, pattern generators and mechanical modeling) to produce a biomimetic simulation of bipedalism based on human body dimensions. The mechanical simulation is a seven-segment, two-dimensional model with motive force provided by tension generators representing the major muscle groups acting around the lower-limb joints. Metabolic energy costs are calculated from the muscle model, and bipedal gait is generated using a finite-state pattern generator whose parameters are produced using a genetic algorithm with locomotor economy (maximum distance for a fixed energy cost) as the fitness criterion. The model is validated by comparing the values it generates with those for modern humans. The result (maximum efficiency of 200 J m(-1)) is within 15% of the experimentally derived value, which is very encouraging and suggests that this is a useful analytic technique for investigating the locomotor behaviour of fossil forms. Initial work suggests that in the future this technique could be used to estimate other locomotor parameters such as top speed. In addition, the animations produced by this technique are qualitatively very convincing, which suggests that this may also be a useful technique for visualizing bipedal locomotion.</ref>
+<ref>	Skoyles JR. Human balance, the evolution of bipedalism and dysequilibrium syndrome.
+Med Hypotheses 2006;66:1060-8.
+:*Abstract: 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.</ref>
+<ref>	Sockol MD, Raichlen DA, Pontzer H. Chimpanzee locomotor energetics and the origin of human bipedalism. Proc Natl Acad Sci U S A 2007;104:12265-9.
+:*Abstract: 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.</ref>
+<ref>	Spoor F, Wood B, Zonneveld F. Implications of early hominid labyrinthine morphology for evolution of human bipedal locomotion. Nature 1994;369:645-8.
+:*Abstract: The upright posture and obligatory bipedalism of modern humans are unique among living primates. The evolutionary history of this behaviour has traditionally been pursued by functional analysis of the postcranial skeleton and the preserved footprint trails of fossil hominids. Here we report a systematic attempt to reconstruct the locomotor behaviour of early hominids by looking at a major component of the mechanism for the unconscious perception of movement, namely by examining the vestibular system of living primates and early hominids. High-resolution computed tomography was used to generate cross-sectional images of the bony labyrinth. Among the fossil hominids the earliest species to demonstrate the modern human morphology is Homo erectus. In contrast, the semicircular canal dimensions in crania from southern Africa attributed to Australopithecus and Paranthropus resemble those of the extant great apes. Among early Homo specimens, the canal dimensions of Stw 53 are unlike those seen in any of the hominids or great apes, whereas those of SK 847 are modern-human-like.</ref>
+<ref>	Stanford CB. Arboreal bipedalism in wild chimpanzees: implications for the evolution of hominid posture and locomotion. Am J Phys Anthropol 2006;129:225-31.
+:*Abstract: Field observations of bipedal posture and locomotion in wild chimpanzees (Pan troglodytes) can serve as key evidence for reconstructing the likely origins of bipedalism in the last prehominid human ancestor. This paper reports on a sample of bipedal bouts, recorded ad libitum, in wild chimpanzees in Bwindi Impenetrable National Park in southwestern Uganda. The Ruhija community of chimpanzees in Bwindi displays a high rate of bipedal posture. In 246.7 hr of observation from 2001-2003, 179 instances of bipedal posture lasting 5 sec or longer were recorded, for a rate of 0.73 bouts per observation hour. Bipedalism was observed only on arboreal substrates, and was almost all postural, and not locomotor. Bipedalism was part of a complex series of positional behaviors related to feeding, which included two-legged standing, one-legged standing with arm support, and other intermediate postures. Ninety-six percent of bipedal bouts occurred in a foraging context, always as a chimpanzee reached to pluck fruit from tree limbs. Bipedalism was seen in both male and female adults, less frequently among juveniles, and rarely in infants. Both the frequency and duration of bipedal bouts showed a significant positive correlation with estimated substrate diameter. Neither fruit size nor nearest-neighbor association patterns were significantly correlated with the occurrence of bipedalism. Bipedalism is seen frequently in the Bwindi chimpanzee community, in part because of the unusual observer conditions at Bwindi. Most observations of bipedalism were made when the animals were in treetops and the observer at eye-level across narrow ravines. This suggests that wild chimpanzees may engage in bipedal behavior more often than is generally appreciated. Models of the likely evolutionary origins of bipedalism are considered in the light of Bwindi bipedalism data. Bipedalism among Bwindi chimpanzees suggests the origin of bipedal posture in hominids to be related to foraging advantages in fruit trees. It suggests important arboreal advantages in upright posture. The origin of postural bipedalism may have preceded and been causally disconnected from locomotor bipedalism.</ref>
+<ref>	Steudel-Numbers KL. Role of locomotor economy in the origin of bipedal posture and gait. Am J Phys Anthropol 2001;116:171-3.</ref>
+<ref>	Sylvester AD. Locomotor decoupling and the origin of hominin bipedalism. J Theor Biol 2006;242:581-90.
+:*Abstract: Theoretical adaptive landscapes and mathematical representations of key constraints of evolutionary and primate biology are used to propose a new hypothesis for the origin of hominin bipedalism. These constraints suggest that the selective pressure that produced this novel form of locomotion was the need for effective suspensory and terrestrial movement. This testable hypothesis, termed the Decoupling Hypothesis, posits that bipedalism is an adaptation that enables the shoulder to maintain a high degree of mobility, a feature important to suspensory behaviors, in the face of significant demands for a high degree of stability, a feature important for highly effective terrestrial quadrupedism.</ref>
+<ref>	Thorpe SK, Holder RL, Crompton RH. Origin of human bipedalism as an adaptation for locomotion on flexible branches. Science 2007;316:1328-31.
+:*Abstract: 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.</ref>
+<ref>Vaughan CL. Theories of bipedal walking: an odyssey. J Biomech 2003;36:513-23.
+Abstract: In this paper six theories of bipedal walking, and the evidence in support of the theories, are reviewed. They include: evolution, minimising energy consumption, maturation in children, central pattern generators, linking control and effect, and robots on two legs. Specifically, the six theories posit that: (1) bipedalism is the fundamental evolutionary adaptation that sets hominids--and therefore humans--apart from other primates; (2) locomotion is the translation of the centre of gravity along a pathway requiring the least expenditure of energy; (3) when a young child takes its first few halting steps, his or her biomechanical strategy is to minimise the risk of falling; (4) a dedicated network of interneurons in the spinal cord generates the rhythm and cyclic pattern of electromyographic signals that give rise to bipedal gait; (5) bipedal locomotion is generated through global entrainment of the neural system on the one hand, and the musculoskeletal system plus environment on the other; and (6) powered dynamic gait in a bipedal robot can be realised only through a strategy which is based on stability and real-time feedback control. The published record suggests that each of the theories has some measure of support. However, it is important to note that there are other important theories of locomotion which have not been covered in this review. Despite such omissions, this odyssey has explored the wide spectrum of bipedal walking, from its origins through to the integration of the nervous, muscular and skeletal systems.</ref>
+<ref>Wang WJ, Crompton RH, Li Y, Gunther MM. Energy transformation during erect and 'bent-hip, bent-knee' walking by humans with implications for the evolution of bipedalism. J Hum Evol 2003;44:563-79.
+:*Abstract: We have previously reported that predictive dynamic modeling suggests that the 'bent-hip, bent-knee' gait, which some attribute to Australopithecus afarensis AL-288-1, would have been much more expensive in mechanical terms for this hominid than an upright gait. Normal walking by modern adult humans owes much of its efficiency to conservation of energy by transformation between its potential and kinetic states. These findings suggest the question if, and to what extent, energy transformation exists in 'bent-hip, bent-knee' gait.This study calculates energy transformation in humans walking upright, at three different speeds, and walking 'bent-hip, bent-knee'. Kinematic data were gathered from video sequences and kinetic (ground reaction force) data from synchronous forceplate measurement. Applying Newtonian mechanics to our experimental data, the fluctuations of kinetic and potential energy in the body centre of mass were obtained and the effects of energy transformation evaluated and compared. In erect walking the fluctuations of two forms of energy are indeed largely out-of-phase, so that energy transformation occurs and total energy is conserved. In 'bent-hip, bent-knee' walking, however, the fluctuations of the kinetic and potential energy are much more in-phase, so that energy transformation occurs to a much lesser extent. Among all modes of walking the highest energy recovery is obtained in subjectively 'comfortable' walking, the next highest in subjectively 'fast' or 'slow' walking, and the least lowest in 'bent-hip, bent-knee' walking. The results imply that if 'bent-hip, bent-knee' gait was indeed habitually practiced by early bipedal hominids, a very substantial (and in our view as yet unidentified) selective advantage would have had to accrue, to offset the selective disadvantages of 'bent-hip, bent-knee' gait in terms of energy transformation.</ref>
+==References==
+<references />

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