Human bipedal instability in tree canopy environments is reduced by "light touch" fingertip support.

Whether tree canopy habitats played a sustained role in the ecology of ancestral bipedal hominins is unresolved. Some argue that arboreal bipedalism was prohibitively risky for hominins whose increasingly modern anatomy prevented them from gripping branches with their feet. Balancing on two legs is indeed challenging for humans under optimal conditions let alone in forest canopy, which is physically and visually highly dynamic. Here we quantify the impact of forest canopy characteristics on postural stability in humans. Viewing a movie of swaying branches while standing on a branch-like bouncy springboard destabilised the participants as much as wearing a blindfold. However "light touch", a sensorimotor strategy based on light fingertip support, significantly enhanced their balance and lowered their thigh muscle activity by up to 30%. This demonstrates how a light touch strategy could have been central to our ancestor's ability to avoid falls and reduce the mechanical and metabolic cost of arboreal feeding and movement. Our results may also indicate that some adaptations in the hand that facilitated continued access to forest canopy may have complemented, rather than opposed, adaptations that facilitated precise manipulation and tool use.

Fibre type composition in the lumbar perivertebral muscles of primates: implications for the evolution of orthogrady in hominoids.

The axial musculoskeletal system is important for the static and dynamic control of the body during both locomotor and non-locomotor behaviour. As a consequence, major evolutionary changes in the positional habits of a species are reflected by morpho-functional adaptations of the axial system. Because of the remarkable phenotypic plasticity of muscle tissue, a close relationship exists between muscle morphology and function. One way to explore major evolutionary transitions in muscle function is therefore by comparative analysis of fibre type composition. In this study, the three-dimensional distribution of slow and fast muscle fibres was analysed in the lumbar perivertebral muscles of two lemuriform (mouse lemur, brown lemur) and four hominoid primate species (white-handed gibbon, orangutan, bonobo, chimpanzee) in order to develop a plausible scenario for the evolution of the contractile properties of the axial muscles in hominoids and to discern possible changes in muscle physiology that were associated with the evolution of orthogrady. Similar to all previously studied quadrupedal mammals, the lemuriform primates in this study exhibited a morpho-functional dichotomy between deep slow contracting local stabilizer muscles and superficial fast contracting global mobilizers and stabilizers and thus retained the fibre distribution pattern typical for quadrupedal non-primates. In contrast, the hominoid primates showed no regionalization of the fibre types, similar to previous observations in Homo. We suggest that this homogeneous fibre composition is associated with the high functional versatility of the axial musculature that was brought about by the evolution of orthograde behaviours and reflects the broad range of mechanical demands acting on the trunk in orthograde hominoids. Because orthogrady is a derived character of euhominoids, the uniform fibre type distribution is hypothesized to coincide with the evolution of orthograde behaviours.

Postural strategies employed by orangutans (Pongo abelii) during feeding in the terminal branch niche.

Obtaining food in an arboreal habitat is complex due to the irregular and flexible nature of the supports available. As the largest predominantly arboreal primate, orangutans are expected to have developed particular postural strategies to enable them to feed successfully. In particular, they need to be able to cope within the terminal branch niche (TBN) as this is where the smallest, most compliant supports are, and also where the majority of the fruit and leaves are situated. We recorded feeding posture, along with a number of ecological and behavioral variables from different age-sex classes to enable analysis of the interactions between these and the compliance of the supports (as estimated from stiffness score). Suspensory postures with a pronograde orientation were used on the most compliant supports for all age-sex classes and appeared to play a particular role in facilitating safe use of the TBN by distributing body weight and using limbs for balance across multiple supports. This contradicts the idea that orthograde suspension evolved in response to the demands of feeding in the TBN. Adult males appear to use the same postures and feeding zones as the other age-sex classes, but appear to use stiffer supports where possible due to their larger body mass. Feeding method differed between the age-sex classes in relation to support stiffness, with larger adult males taking fewer risks due to their larger size, compared to infants and juveniles.

A new method for recording complex positional behaviours and habitat interactions in primates.

In an arboreal habitat, primates have to cope with a complex meshwork of flexible supports in order to obtain food, find mates and avoid predators. To understand how animals interact with such complex environments we can study their positional behaviour. However, due to the intricate variation in locomotion and posture it can be difficult to capture details such as limb use (i.e. weight and balance), limb flexion and substrate use. This paper presents a suitable method replicable for any primate species, based on the movement notation technique, Sutton Movement Writing (SMW), aiming to record the spatial arrangement of limbs during positional behaviours on multiple, compliant supports. This method was piloted during a year-long field study of wild orangutans (Pongo abelii) and validated and tested for inter- and intraobserver reliability using videos from the field. Overall, SMW shows considerable promise for increasing the resolution with which positional behaviours can be recorded under field conditions and provides a way to extract numerical data for use in statistical analyses. This will facilitate our understanding of how behaviours vary in response to the environment, and the capabilities of primates to perform key tasks in their distinct niches.

Operative treatment of Linburg-Comstock syndrome.

Linburg-Comstock syndrome is characterised by an anomalous tendon slip from the flexor pollicis longus to the flexor digitorum profundus, usually of the index finger. An incidence as high as 60% to 70% has been reported. Post-traumatic inflammation of inter-tendinous connections between the flexor pollicis longus and flexor digitorum profundus, usually of the index finger, may cause unexplained chronic pain in the distal forearm. A total of 11 patients (eight females, three males), mean age 29.1 years (14 to 47) with a clinical diagnosis of Linburg-Comstock syndrome underwent surgical release of the inter-tendinous connection. The mean follow-up was for 27 months (2 to 48). Ten patients reported excellent relief of pain in the forearm, with independent flexion of flexor pollicis longus and flexor digitorum profundus to the index finger. Surgical release was an effective treatment for the Linburg-Comstock syndrome in this series.

Locomotion and posture from the common hominoid ancestor to fully modern hominins, with special reference to the last common panin/hominin ancestor.

Based on our knowledge of locomotor biomechanics and ecology we predict the locomotion and posture of the last common ancestors of (a) great and lesser apes and their close fossil relatives (hominoids); (b) chimpanzees, bonobos and modern humans (hominines); and (c) modern humans and their fossil relatives (hominins). We evaluate our propositions against the fossil record in the context of a broader review of evolution of the locomotor system from the earliest hominoids of modern aspect (crown hominoids) to early modern Homo sapiens. While some early East African stem hominoids were pronograde, it appears that the adaptations which best characterize the crown hominoids are orthogrady and an ability to abduct the arm above the shoulder - rather than, as is often thought, manual suspension sensu stricto. At 7-9 Ma (not much earlier than the likely 4-8 Ma divergence date for panins and hominins, see Bradley, 2008) there were crown hominoids in southern Europe which were adapted to moving in an orthograde posture, supported primarily on the hindlimb, in an arboreal, and possibly for Oreopithecus, a terrestrial context. By 7 Ma, Sahelanthropus provides evidence of a Central African hominin, panin or possibly gorilline adapted to orthogrady, and both orthogrady and habitually highly extended postures of the hip are evident in the arboreal East African protohominin Orrorin at 6 Ma. If the traditional idea that hominins passed through a terrestrial 'knuckle-walking' phase is correct, not only does it have to be explained how a quadrupedal gait typified by flexed postures of the hindlimb could have preadapted the body for the hominin acquisition of straight-legged erect bipedality, but we would have to accept a transition from stem-hominoid pronogrady to crown hominoid orthogrady, back again to pronogrady in the African apes and then back to orthogrady in hominins. Hand-assisted arboreal bipedality, which is part of a continuum of orthograde behaviours, is used by modern orangutans to forage among the small branches at the periphery of trees where the core hominoid dietary resource, ripe fruit, is most often to be found. Derivation of habitual terrestrial bipedality from arboreal hand-assisted bipedality requires fewer transitions, and is also kinematically and kinetically more parsimonious.

Origin of human bipedalism as an adaptation for locomotion on flexible branches.

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

Orangutans use compliant branches to lower the energetic cost of locomotion.

Within the forest canopy, the shortest gaps between tree crowns lie between slender terminal branches. While the compliance of these supports has previously been shown to increase the energetic cost of gap crossing in arboreal animals (e.g. Alexander 1991 Z. Morphol. Anthropol. 78, 315-320; Demes et al. 1995 Am. J. Phys. Anthropol. 96, 419-429), field observations suggest that some primates may be able to use support compliance to increase the energetic efficiency of locomotion. Here, we calculate the energetic cost of alternative methods of gap crossing in orangutans (Pongo abelii). Tree sway (in which orangutans oscillate a compliant tree trunk with increasing magnitude to bridge a gap) was found to be less than half as costly as jumping, and an order of magnitude less costly than descending the tree, walking to the vine and climbing it. Observations of wild orangutans suggest that they actually use support compliance in many aspects of their locomotor behaviour. This study seems to be the first to show that elastic compliance in arboreal supports can be used to reduce the energetic cost of gap crossing.

Morphological analysis of the hindlimb in apes and humans. II. Moment arms.

Flexion/extension moment arms were obtained for the major muscles crossing the hip, knee and ankle joints in the orang-utan, gibbon, gorilla (Eastern and Western lowland) and bonobo. Moment arms varied with joint motion and were generally longer in proximal limb muscles than distal limb muscles. The shape of the moment arm curves (i.e. the plots of moment arm against joint angle) differed in different hindlimb muscles and in the same muscle in different subjects (both in the same and in different ape species). Most moment arms increased with increasing joint flexion, a finding which may be understood in the context of the employment of flexed postures by most non-human apes (except orang-utans) during both terrestrial and arboreal locomotion. When compared with humans, non-human great apes tended to have muscles better designed for moving the joints through large ranges. This was particularly true of the pedal digital flexors in orang-utans. In gibbons, the only lesser ape studied here, many of the moment arms measured were relatively short compared with those of great apes. This study was performed on a small sample of apes and thus differences noted here warrant further investigation in larger populations.

Morphological analysis of the hindlimb in apes and humans. I. Muscle architecture.

We present quantitative data on the hindlimb musculature of Pan paniscus, Gorilla gorilla gorilla, Gorilla gorilla graueri, Pongo pygmaeus abelii and Hylobates lar and discuss the findings in relation to the locomotor habits of each. Muscle mass and fascicle length data were obtained for all major hindlimb muscles. Physiological cross-sectional area (PCSA) was estimated. Data were normalized assuming geometric similarity to allow for comparison of animals of different size/species. Muscle mass scaled closely to (body mass)(1.0) and fascicle length scaled closely to (body mass)(0.3) in most species. However, human hindlimb muscles were heavy and had short fascicles per unit body mass when compared with non-human apes. Gibbon hindlimb anatomy shared some features with human hindlimbs that were not observed in the non-human great apes: limb circumferences tapered from proximal-to-distal, fascicle lengths were short per unit body mass and tendons were relatively long. Non-human great ape hindlimb muscles were, by contrast, characterized by long fascicles arranged in parallel, with little/no tendon of insertion. Such an arrangement of muscle architecture would be useful for locomotion in a three dimensionally complex arboreal environment.

Stresses exerted in the hindlimb muscles of common chimpanzees (Pan troglodytes) during bipedal locomotion.

Recent studies have indicated that chimpanzee bipedality is mechanically inefficient and dynamically unlike that of humans, thus undermining the chimpanzee analogy for mechanical aspects of the early evolution of hominid bipedalism. This paper continues this theme by measuring the forces and stresses engendered by the muscles during bipedal locomotion, for an untrained chimpanzee and for data from chimpanzees which have been encouraged to walk bipedally, presented in the literature. Peak stresses in the triceps surae were lower for the untrained chimpanzee than for the trained subjects because during the late stance phase, when peak ankle moments occur, the centre of pressure of the ground reaction force on the foot of the untrained chimpanzee stayed close to the ankle joint. In contrast, for the trained subjects it moved closer to the toes, as in human bipedalism. Quadriceps and hip extensor stresses are approximately 30% larger for the untrained chimpanzee than for the trained subjects, because the trained chimpanzees walked with a more erect posture. These results may reflect the way in which muscles can develop in response to training, since research on humans has shown that muscle physiological cross-sectional area increases as a result of exercise, resulting in smaller stresses for a given muscle force. During a slow walk, untrained chimpanzees were found to exert far greater muscle stresses than humans do when running at moderate speed, particularly in the muscles that extend the hip, because of the bent-hip, bent-knee posture.

Dimensions and moment arms of the hind- and forelimb muscles of common chimpanzees (Pan troglodytes).

This paper supplies quantitative data on the hind- and forelimb musculature of common chimpanzees (Pan troglodytes) and calculates maximum joint moments of force as a contribution to a better understanding of the differences between chimpanzee and human locomotion. We dissected three chimpanzees, and recorded muscle mass, fascicle length, and physiological cross-sectional area (PCSA). We also obtained flexion/extension moment arms of the major muscles about the limb joints. We find that in the hindlimb, chimpanzees possess longer fascicles in most muscles but smaller PCSAs than are predicted for humans of equal body mass, suggesting that the adaptive emphasis in chimpanzees is on joint mobility at the expense of tension production. In common chimpanzee bipedalism, both hips and knees are significantly more flexed than in humans, necessitating muscles capable of exerting larger moments at the joints for the same ground force. However, we find that when subject to the same stresses, chimpanzee hindlimb muscles provide far smaller moments at the joints than humans, particularly the quadriceps and plantar flexors. In contrast, all forelimb muscle masses, fascicle lengths, and PCSAs are smaller in humans than in chimpanzees, reflecting the use of the forelimbs in chimpanzee, but not human, locomotion. When subject to the same stresses, chimpanzee forelimb muscles provide larger moments at the joints than humans, presumably because of the demands on the forelimbs during locomotion. These differences in muscle architecture and function help to explain why chimpanzees are restricted in their ability to walk, and particularly to run bipedally.

Stresses in human leg muscles in running and jumping determined by force plate analysis and from published magnetic resonance images.

Calculation of the stresses exerted by human muscles requires knowledge of their physiological cross-sectional area (PCSA). Magnetic resonance imaging (MRI) has made it possible to measure PCSAs of leg muscles of healthy human subjects, which are much larger than the PCSAs of cadaveric leg muscles that have been used in previous studies. We have used published MRI data, together with our own force-plate records and films of running and jumping humans, to calculate stresses in the major groups of leg muscles. Peak stresses in the triceps surae ranged from 100 kN m-2 during take off for standing high jumps to 150 kN m-2 during running at 4 m s-1. In the quadriceps, peak stresses ranged from 190 kN m-2 during standing long jumps to 280 kN m-2 during standing high jumps. Similar stresses were calculated from published measurements of joint moments. These stresses are lower than those previously calculated from cadaveric data, but are in the range expected from physiological experiments on isolated muscles.