ABSTRACT Adaptive explanations for modern human foot anatomy have long fascinated evolutionary biologists because of the dramatic differences between our feet and those of our closest living relatives, the great apes. Morphological features, including hallucal opposability, toe length and the longitudinal arch, have traditionally been used to dichotomize human and great ape feet as being adapted for bipedal walking and arboreal locomotion, respectively. However, recent biomechanical models of human foot function and experimental investigations of great ape locomotion have undermined this simple dichotomy. Here, we review this research, focusing on the biomechanics of foot strike, push-off and elastic energy storage in the foot, and show that humans and great apes share some underappreciated, surprising similarities in foot function, such as use of plantigrady and ability to stiffen the midfoot. We also show that several unique features of the human foot, including a spring-like longitudinal arch and short toes, are likely adaptations to long distance running. We use this framework to interpret the fossil record and argue that the human foot passed through three evolutionary stages: first, a great ape-like foot adapted for arboreal locomotion but with some adaptations for bipedal walking; second, a foot adapted for effective bipedal walking but retaining some arboreal grasping adaptations; and third, a human-like foot adapted for enhanced economy during long-distance walking and running that had lost its prehensility. Based on this scenario, we suggest that selection for bipedal running played a major role in the loss of arboreal adaptations.

SUMMARY In the present study we have estimated the temporal elongation of the plantar aponeurosis (PA) during normal walking using a subject-specific multi-segment rigid-body model of the foot. As previous studies have suggested that muscular forces at the ankle can pre-load the PA prior to heel-strike,the main purpose of the current study was to test, through modelling, whether there is any tension present in the PA during early stance phase. Reflective markers were attached to bony landmarks to track the kinematics of the calcaneus, metatarsus and toes during barefoot walking. Ultrasonography measurements were performed on three subjects to determine both the location of the origin of the PA on the plantar aspect of the calcaneus, and the radii of the metatarsal heads. Starting with the foot in a neutral, unloaded position, inverse kinematics allowed calculation of the tension in the five slips of the PA during the whole duration of the stance phase. The results show that the PA experienced tension significantly above rest during early stance phase in all subjects ( P <0.01), thus providing support for the PA-preloading hypothesis. The amount of preloading and the maximum elongation of the slips of the PA decreased from medial to lateral. The mean maximum tension exerted by the PA was 1.5 BW (body weight) over the three subjects.

SUMMARY The phalangeal portion of the forefoot is extremely short relative to body mass in humans. This derived pedal proportion is thought to have evolved in the context of committed bipedalism, but the benefits of shorter toes for walking and/or running have not been tested previously. Here, we propose a biomechanical model of toe function in bipedal locomotion that suggests that shorter pedal phalanges improve locomotor performance by decreasing digital flexor force production and mechanical work, which might ultimately reduce the metabolic cost of flexor force production during bipedal locomotion. We tested this model using kinematic, force and plantar pressure data collected from a human sample representing normal variation in toe length ( N =25). The effect of toe length on peak digital flexor forces, impulses and work outputs was evaluated during barefoot walking and running using partial correlations and multiple regression analysis, controlling for the effects of body mass,whole-foot and phalangeal contact times and toe-out angle. Our results suggest that there is no significant increase in digital flexor output associated with longer toes in walking. In running, however, multiple regression analyses based on the sample suggest that increasing average relative toe length by as little as 20% doubles peak digital flexor impulses and mechanical work,probably also increasing the metabolic cost of generating these forces. The increased mechanical cost associated with long toes in running suggests that modern human forefoot proportions might have been selected for in the context of the evolution of endurance running.