Foot use during vertical climbing in chimpanzees (Pan troglodytes).

Upright bipedalism is a hallmark of hominin locomotion, however debates continue regarding the extent of arboreal locomotion and the nature of bipedalism practiced by early hominins. Pedal form and function play a prominent role in these debates, as the foot is the element that directly interacts with the locomotor substrate. Recent finds have substantially increased the availability of associated foot remains of early hominins and emphasized the enigmatic nature of the early evolution of human bipedalism. New discoveries of associated forefoot remains have afforded the opportunity to assess relative proportions across the forefoot of fossil hominins and illuminated the need for data on relative loading across the forefoot in extant hominoids. In order to provide functional data with which to examine the relationship between bony features and load distribution across the forefoot during climbing, we present the first analysis of plantar pressure distribution across the forefoot of chimpanzees climbing a vertical support. Chimpanzees load the medial metatarsals and first toe disproportionately during vertical climbing. Peak pressures on these elements occur at the end of stance phase during climbing and are higher than on any other elements of the foot. Toe pressures are considerably higher during vertical climbing than during knuckle-walking or movement on horizontal poles, supporting the notion that the plantarly-broad and dorsally-narrow metatarsal heads in chimpanzees and some early hominins are associated with close-packing of the metatarsophalangeal joint during climbing.

Dynamics of locomotor transitions from arboreal to terrestrial substrates in Verreaux's sifaka (Propithecus verreauxi).

Most primates are able to move with equal facility on the ground and in trees, but most use the same quadrupedal gaits in both environments. A few specialized primates, however, use a suspensory or leaping mode of locomotion when in the trees but a bipedal gait while on the ground. This is a rare behavioral pattern among mammals, and the extent to which the bipedal gaits of these primates converge and are constrained by the anatomical and neurological adaptations associated with arboreal locomotion is poorly understood. Sifakas (Propithecus), primates living only in Madagascar, are highly committed vertical clingers and leapers that also spend a substantial amount of time on the ground. When moving terrestrially sifakas use a unique bipedal galloping gait seen in no other mammals. Little research has examined the mechanics of these gaits, and most of that research has been restricted to controlled captive conditions. The energetic costs associated with leaping and bipedal galloping are unknown. This study begins to fill that gap using triaxial accelerometry to characterize and compare the dynamics of sifakas' leaping and bipedal galloping behavior. As this is a relatively novel approach, the first goal of this article is to explore the feasibility of collecting such data on free-roaming animals and attempt to automate the identification of leaping and bipedal behavior within the output. The second goal is to compare the overall accelerations of the body and to use that as an approximation of aspects of energetic costs during leaping and bipedalism. To achieve this, a lightweight accelerometer was mounted on freely moving sifakas. The resulting acceleration profiles were processed, and sequences of leaps (bouts) were automatically extracted from the waveforms with 85% accuracy. Both vector dynamic body acceleration and overall dynamic body acceleration (ODBA) were used to characterize locomotor patterns and energy expenditure during leaping and bipedalism. The unique kinematics of the gait of sifakas, and the mechanics of bouts involving a string of successive leaps or gallops, appear to minimize redirections of the center of mass as well as the number of acceleration peaks and ODBAs. These results suggest that bipedal galloping is not only a reflection of the unique anatomical configuration of a leaping primate, but it may also provide a musculoskeletal and an energetic advantage to sifakas. In that sense, bipedal galloping represents an advantageous way for sifakas to move when transitioning from arboreal leaping to terrestrial locomotion.

The foot of Homo floresiensis.

Homo floresiensis is an endemic hominin species that occupied Liang Bua, a limestone cave on Flores in eastern Indonesia, during the Late Pleistocene epoch. The skeleton of the type specimen (LB1) of H. floresiensis includes a relatively complete left foot and parts of the right foot. These feet provide insights into the evolution of bipedalism and, together with the rest of the skeleton, have implications for hominin dispersal events into Asia. Here we show that LB1's foot is exceptionally long relative to the femur and tibia, proportions never before documented in hominins but seen in some African apes. Although the metatarsal robusticity sequence is human-like and the hallux is fully adducted, other intrinsic proportions and pedal features are more ape-like. The postcranial anatomy of H. floresiensis is that of a biped, but the unique lower-limb proportions and surprising combination of derived and primitive pedal morphologies suggest kinematic and biomechanical differences from modern human gait. Therefore, LB1 offers the most complete glimpse of a bipedal hominin foot that lacks the full suite of derived features characteristic of modern humans and whose mosaic design may be primitive for the genus Homo. These new findings raise the possibility that the ancestor of H. floresiensis was not Homo erectus but instead some other, more primitive, hominin whose dispersal into southeast Asia is still undocumented.

Manual digital pressures during knuckle-walking in chimpanzees (Pan troglodytes).

Considerable attention has been given to hand morphology and function associated with knuckle-walking in the African apes because of the implications they have for the evolution of bipedalism in early hominins. Knuckle-walking is associated with a unique suite of musculoskeletal features of the wrist and hand, and numerous studies have hypothesized that these anatomical features are associated with the dynamics of load distribution across the digits during knuckle-walking. We collected dynamic digital pressures on two chimpanzees during terrestrial and simulated arboreal locomotion. Comparisons were made across substrates, limb positions, hand positions, and age categories. Peak digital pressures were similar on the pole and on the ground but were distributed differently across the digits on each substrate. In young animals, pressure was equally high on digits 2-4 on the ground but higher on digits 3 and 4 on the pole. Older animals experience higher pressures on digits 2 and 3 on the ground. Hand posture (palm-in vs. palm-back) influenced the distribution and timing of peak pressures. Age-related increases in body mass also result in higher overall pressures and increased variation across the digital row. In chimpanzees, digit 5 typically bears relatively little load regardless of hand position or substrate. These are the first quantitative data on digital pressures during knuckle-walking in hominoids, and they afford the opportunity to develop hypotheses about variation among hominoids and biomechanical models of wrist and forearm loading.

The hands and feet of Archaeolemur: metrical affinities and their functional significance.

Recent expeditions to Madagascar have recovered abundant skeletal remains of Archaeolemur, one of the so-called "monkey lemurs" known from Holocene deposits scattered across the island. These new skeletons are sufficiently complete to permit reassembly of entire hands and feet--postcranial elements crucial to drawing inferences about substrate preferences and positional behavior. Univariate and multivariate analysis of intrinsic hand and foot proportions, phalangeal indices, relative pollex and hallux lengths, phalangeal curvature, and distal phalangeal shape reveal a highly derived and unique morphology for an extinct strepsirrhine that diverges dramatically from that of living lemurs and converges in some respects on that of Old World monkeys (e.g., mandrills, but not baboons or geladas). The hands and feet of Archaeolemur are relatively short (extremely so relative to body size); the carpus and tarsus are both "long" relative to total hand and foot lengths, respectively; phalangeal indices of both the hands and feet are low; both pollex and hallux are reduced; the apical tufts of the distal phalanges are very broad; and the proximal phalanges are slightly curved (but more so than in baboons). Overall grasping capabilities may have been compromised to some extent, and dexterous handling of small objects seems improbable. Deliberate and noncursorial quadrupedalism was most likely practiced on both the ground and in the trees. A flexible locomotor repertoire in conjunction with a eurytopic trophic adaptation allowed Archaeolemur to inhabit much of Madagascar and may explain why it was one of the latest surviving subfossil lemurs.

Gender differences in adult foot shape: implications for shoe design.

PURPOSE: To analyze gender differences in foot shape in a large sample of young individuals. METHODS: Univariate t-tests and multivariate discriminant analyses were used to assess 1) significant differences between men and women for each foot and leg dimension, standardized to foot length, 2) the reliability of classification into gender classes using the absolute and standardized variable sets, and 3) the relative importance of each variable to the discrimination between men and women. RESULTS: Men have longer and broader feet than women for a given stature. After normalization of the measurements by foot length, men and women were found to differ significantly in two calf, five ankle, and four foot shape variables. Classification by gender using absolute values was correct at least 93% of the time. Using the variables standardized to foot length, gender was correctly classified 85% of the time. CONCLUSIONS: This study demonstrates that female feet and legs are not simply scaled-down versions of male feet but rather differ in a number of shape characteristics, particularly at the arch, the lateral side of the foot, the first toe, and the ball of the foot. These differences should be taken into account in the design and manufacture of women's sport shoes.

Reappraisal of the postcranium of Hadropithecus (Primates, Indroidea).

Hadropithecus stenognathus (Lorenz von Liburnau [1899] Anz. Akad. Wiss. Wien 36:255-257), a giant extinct lemur from Madagascar, has been reconstructed as primarily terrestrial and probably cursorial on the basis of its postcranial anatomy, especially long bone gracility and interlimb proportions. We show here that aspects of this reconstruction are almost certainly incorrect. Hindlimb bones of Archaeolemur have been misattributed to Hadropithecus, and new hindlimb allocations (including newly recognized elements such as the calcaneus) indicate that Hadropithecus had a robust body build and lacked osteological specializations for cursoriality. We review the evidence for the existence of "Bradylemur" and offer a view of archaeolemurid positional behavior that includes terrestrial and arboreal components. Body size and limb proportions of Hadropithecus are reassessed in light of our new allocations.

New pedal remains of Megaladapis and their functional significance.

New remains of Megaladapis from the caves within the Ankarana Range of northern Madagascar and the cave site of Ankilitelo near Toliara in southwestern Madagascar add considerably to the present sample of pedal remains for this genus. Here we describe and analyze the new pedal material and discuss the function of the Megaladapis foot in terms of positional behavior and substrate use. The northern specimens belong to the M. madagascariensis /M. grandidieri group in terms of size and morphology whereas the new southwestern fossils are assigned to M. madagascariensis. The new specimens demonstrate that the small and intermediate sized M. madagascariensis and M. grandidieri were very similar in anatomy and inferred locomotor function, findings that also support the prior suggestion that they belong to a single widespread subgenus (Megaladapis). The new fossils provide the first examples of many pedal elements and present the first opportunity to analyze the whole pedal complex from associated remains. The foot of Megaladapis is distinctive among primates in numerous features. Intrinsic proportions of the hindlimb indicate that the foot is relatively longer than that of any other primate. The first complete calcanei reveal a large and highly modified hindfoot. The calcaneus is reduced distally, indicating an emphasis on climbing over leaping or quadrupedal walking and running. Proximally, a large, medially directed calcaneal tuberosity suggests both a strong inversion component to plantarflexion and a well-developed abductor mechanism and recalls the calcaneal morphology of the larger lorisines in some respects. Talar shape is consistent with considerable tibial rotation during plantarflexion and dorsiflexion. The subtalar joint is designed to emphasize supination/pronation and medial/lateral rotation over proximodistal translation. The distal tarsals are extremely reduced in length, and they form a high transverse arch and a serial tarsus; this configuration promotes inversion/eversion at the transverse tarsal joint. The phalanges are long and moderately curved, and the hallux is very long, robust, and abducted. Pedal morphology suggests that Megaladapis (subgenus Megaladapis) was well adapted to exploit an arboreal environment.(ABSTRACT TRUNCATED)

Changes in astroglial scar formation in rat optic nerve as a function of development.

Although astroglial scar formation is a common response to almost any type of injury to the adult central nervous system, lesions in fetal and neonatal rats have been reported to induce little or no scar formation. To examine this developmental difference further, rats ranging in age from 1 to 65 days postnatal were unilaterally enucleated, a surgical procedure that causes the axons in the optic nerve to degenerate. The optic nerves were processed for light and electron microscopy at times ranging from 7 to 365 days postenucleation. Pronounced and permanent glial scars were formed in every age group examined, including the neonates. However, the time course for removal of the degenerating axonal debris and formation of a compact, debris-free glial scar varied as a function of developmental age. In neonatal rats, a compact glial scar formed in 1-2 weeks whereas 3-5 months were required for compact glial scar formation in juveniles and adults. Changes in cross-sectional area were also associated with optic nerve degeneration and glial scar formation. Whereas lesioned neonatal optic nerves underwent little change in area, there was a substantial decrease in area in the juvenile and adult. Morphometric analysis showed that irrespective of the age of the animal at the time of enucleation, the final area of the compact glial scar was 10-20% of the unlesioned adult control. These results suggest that conflict in the literature over the ability of neonatal astrocytes to form a glial scar may be due to the nature of the lesion or the method of detection since astrocytes in the neonatal rat optic nerve clearly have the capacity to become reactive and form a glial scar.