Modern Human Foot Research Articles

Variable gearing at the ankle during walking in adults and young children: implications for foot development and evolution

Introduction: The human foot has evolved over the past seven million years from a relatively mobile, grasping appendage to a highly derived structure with a heel pad and longitudinal arch that can absorb shock at heel strike and weight-bearing yet also function as a powerful lever at toe-off. It has been proposed that the modern human foot evolved to allow our species to use “variable gearing” during walking and running. In this model, the gears of the human foot are defined relative to the ankle center of rotation as R, the distance from the ground reaction resultant vector, and r, the distance from the calf muscle vector. The gear ratio defines the torque generated to propel the body or stretch the triceps surae muscles. We test the hypothesis that variable gearing is associated with modern human pedal anatomy and a heel-to-toe rollover that allows a shift from “low gear” to “high gear” during stance.Methods: Using force plate and video analysis, we examined variable gearing in adults and children engaging in adult heel strike (AHS = 35), flat foot contact (FFC = 39), or initial heel contact (IHC = 26).Results and Discussion: Our hypothesis was partly supported. Although variable gearing was observed in IHC steps and was greater than in FFC steps, it was not as developed as in AHS steps. This may be related to anatomical and motor control differences between juvenile and adult feet, suggesting that adult anatomy, including a high arch, and neural control are critical for full use of variable gearing and that this feature would have evolved in later hominins around two million years ago with the appearance of a fully modern foot.

Evolutionary selection and morphological integration in the foot of modern humans

AbstractObjectivesTo advance our understanding of the evolution of the hominin foot by quantifying integration and responses to selection in the foot of modern humans.Materials and MethodsThe sample includes 247 female and male adult individuals from Euro‐American, Afro‐American, European, and Amerindian populations. We collected 190 linear measurements from the 26 skeletal elements that constitute the modern human foot. With these data, we calculated the magnitudes of integration and the ability of the foot to respond to selection demands.ResultsThe results revealed that distal phalanges are less integrated, more evolvable, and more flexible than proximal elements (i.e., proximal phalanges and metatarsals). Also, bones from the medial ray (e.g., hallux) show stronger integration and weaker evolvability than their counterparts from the lateral column (e.g., fifth ray), following this trend from medial to lateral positions. Among the tarsals, the talus and calcaneus are the most integrated, least evolvable, and flexible elements from that module.DiscussionThese results suggest that selection for bipedalism would have reorganized the variance/covariance matrix of the foot. The hallux might have been under strong functional selection pressures for bipedal requirements, resulting in a strong integration and low evolvability. Also, differences in the developmental process of each bone seem to have played an essential role in the degree of evolvability, showing those elements that develop earlier have less ability to respond to selection demands.

Take a load off: skeletal implications of sedentism in the feet of modern body donors.

Modern biocultural environments continue to place selective pressures on our skeletons. In the past century, a major cultural pressure has been the rise in sedentism. However, studies considering the effects of sedentism on the foot have largely considered pathological changes to the gross foot without particular regard for the pedal skeleton. To address this gap in the literature, temporal trends in the development of osteoarthritis and entheseal changes on the tarsals and metatarsals were analyzed in the context of biodemographic data for recent modern humans. The sample utilized for this project is comprised of 71 individuals from the William M. Bass Donated Skeletal Collection, with birth years ranging from 1909 to 1993. Temporal trends in osteoarthritis and entheseal changes were determined via ANCOVA, using year of birth as the explanatory variable and biodemographic variables (age, sex, stature, body mass index and tibial robusticity) as covariates. Results indicate that entheseal changes and osteoarthritis have decreased over time, and these trends are statistically significant. Temporal trends in pedal entheseal changes and osteoarthritis vary by sex. The increase in sedentary behavior over time has usually been framed as a net negative for human health and well-being. However, considered in isolation, the decrease in entheseal changes and osteoarthritis presented here might be considered a positive development as they suggest overall less stress on the modern human foot. This study also has the potential to inform the health sciences and general public about biocultural contributors to modern foot health.

Mechanics of heel-strike plantigrady in African apes

The initiation of a walking step with a heel strike is a defining characteristic of humans and great apes but is rarely found in other mammals. Despite the considerable importance of heel strike to an understanding of human locomotor evolution, no one has explicitly tested the fundamental mechanical question of why great apes use a heel strike. In this report, we test two hypotheses (1) that heel strike is a function of hip protraction and/or knee extension and (2) that short-legged apes with a midfoot that dorsiflexes at heel lift and long digits for whom digitigrady is not an option use heel-strike plantigrady. This strategy increases hip translation while potentially moderating the cost of redirecting the center of mass (‘collisional costs’) during stance via rollover along the full foot from the heel to toes. We quantified hind limb kinematics and relative hip translation in ten species of primates, including lemurs, terrestrial and arboreal monkeys, chimpanzees, and gorillas. Chimpanzees and gorillas walked with relatively extended knees but only with moderately protracted hips or hind limbs, partially rejecting the first hypothesis. Nonetheless, chimpanzees attained relative hip translations comparable with those of digitigrade primates. Heel-strike plantigrady may be a natural result of a need for increased hip translations when forelimbs are relatively long and digitigrady is morphologically restricted. In addition, foot rollover from the heel to toe in large, short-legged apes may reduce energetic costs of redirecting the center of mass at the step-to-step transition as it appears to do in humans. Heel strike appears to have been an important mechanism for increasing hip translation, and possibly reducing energetic costs, in early hominins and was fundamental to the evolution of the modern human foot and human bipedalism.

Radiographic measurements of the talus and calcaneus in the adult pes planus foot type.

A distinctive feature of the modern human foot is the presence of a medial longitudinal arch when weight-bearing. Although the talus and calcaneus play a major role in the structure and function of the human foot, the association between the morphology of these bones and longitudinal arch height has not been fully investigated. A better understanding of this relationship may assist in the interpretation of pedal remains of fossil hominins, where features of the foot and ankle morphology have been described as providing evidence for the presence of a longitudinal arch. For this study, weight-bearing radiographs of 103 patients from an urban US Level 1 trauma center, taken as part of a clinical examination for medical evaluation, were selected. These radiographs were classified as to foot type by arch height as defined using the calcaneal inclination angle. From this group, 68 radiographs were suitable for linear and angular measurements of the talus and 74 of the calcaneus. The relationships between these measurements and arch height were explored using least squared linear regression analysis. The results demonstrate that angular measurements of the calcaneus (particularly those that reflect the relationship of the talar articular facets to each other and the tilt of the calcaneocuboid joint to the longitudinal axis of the calcaneus) are predictive of arch height (r2 = .29-.44 p ≤ .001). All angular measurements of the talus and all examined linear measurements of both the talus and calcaneus were not predictive of arch height. These results suggest that certain angular measurements of the calcaneus are associated with arch height in the modern human foot. While this information is useful in the interpretation of hominin pedal remains, the relationship of the morphology of these bones, as well as other bones of the foot, to arch height is complex, requiring further investigation.

Early Upper Paleolithic human foot bones from Manot Cave, Israel

The transition from the Middle Paleolithic to the Upper Paleolithic in the Levant represents a major event in human prehistory with regards to the dispersal of modern human populations. Unfortunately, the scarcity of human remains from this period has hampered our ability to study the anatomy of Upper Paleolithic populations. This study describes and examines pedal bones recovered from the Early Upper Paleolithic period at Manot Cave, Israel, from 2014 to 2017. The Manot Cave foot bones include a partial, left foot skeleton comprising a talus, a calcaneus, a cuboid, a first metatarsal, a second metatarsal, a fifth metatarsal, and a hallucal sesamoid. All these remains were found in the same archaeological unit of the cave and belong to a young adult. Shape and size comparisons with Neanderthals, Anatomically Modern Human and modern human foot bones indicate a modern human morphology. In some characteristics, however, the Manot Cave foot bones display a Neanderthal-like pattern. Notably, the Manot Cave foot is remarkable in its overall gracility. A healed traumatic injury in the second metatarsal (Lisfranc's fracture) is most likely due to a remote impact to the dorsum of the foot. This injury, its subsequent debility, and the individual's apparent recovery suggest that the members of the Manot Cave community had a supportive environment, one with mutual responsibilities among the members.

The African ape-like foot of Ardipithecus ramidus and its implications for the origin of bipedalism.

The ancestral condition from which humans evolved is critical for understanding the adaptive origin of bipedal locomotion. The 4.4 million-year-old hominin partial skeleton attributed to Ardipithecus ramidus preserves a foot that purportedly shares morphometric affinities with monkeys, but this interpretation remains controversial. Here I show that the foot of Ar. ramidus is most similar to living chimpanzee and gorilla species among a large sample of anthropoid primates. The foot morphology of Ar. ramidus suggests that the evolutionary precursor of hominin bipedalism was African ape-like terrestrial quadrupedalism and climbing. The elongation of the midfoot and phalangeal reduction in Ar. ramidus relative to the African apes is consistent with hypotheses of increased propulsive capabilities associated with an early form of bipedalism. This study provides evidence that the modern human foot was derived from an ancestral form adapted to terrestrial plantigrade quadrupedalism.

Rethinking the evolution of the human foot: insights from experimental research.

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.

The Relationship Between Linear Osteological and Radiographic Measurements of the Human Calcaneus and Talus

Plain radiographs of the foot are a common form of examination in individuals when medical evaluation is indicated. Their availability makes them useful for studying human variation where large sample sizes are crucial. Calcaneal and talar morphology are critical in the understanding of human foot form and function, but few studies have examined the accuracy of linear measurements of these bones taken from radiographs. If linear measurements are to be used in quantitative analysis, their accuracy must first be demonstrated. For this study, 54 human feet from amputated limbs were imaged in standard radiographic views and skeletonized. Selected linear measurements were made on each skeletonized talus and calcaneus. Corresponding landmarks were identified on the radiographic images and the distances between these points determined. Using the measurements taken from the physical bone, the accuracy of the radiographic measurements was evaluated employing paired Student's t-tests and correlation analysis. Most measurements were not significantly different (P ≥ 0.05). Overall, the sagittal plane measurements were correlated (r2 > 0.61-0.88, all P < 0.001) with the radiographic measurements; transverse plane measurements less so (r2 = 0.20-0.63, P < 0.001). These results demonstrate that measurements from radiographic images provide useful information concerning both the calcaneus and talus, indicating that radiographs of living individuals can be evaluated quantitatively and compared to osteology collections of modern humans. This information can be applied to the understanding of functional anatomy of the modern human foot, interpreting the hominin fossil pedal record, and establishing identification in forensic analysis. Anat Rec, 2017. © 2017 Wiley Periodicals, Inc. Anat Rec, 301:21-33, 2018. © 2017 Wiley Periodicals, Inc.

An investigation of the dynamic relationship between navicular drop and first metatarsophalangeal joint dorsal excursion

The modern human foot is a complex biomechanical structure that must act both as a shock absorber and as a propulsive strut during the stance phase of gait. Understanding the ways in which foot segments interact can illuminate the mechanics of foot function in healthy and pathological humans. It has been proposed that increased values of medial longitudinal arch deformation can limit metatarsophalangeal joint excursion via tension in the plantar aponeurosis. However, this model has not been tested directly in a dynamic setting. In this study, we tested the hypothesis that during the stance phase, subtalar pronation (stretching of the plantar aponeurosis and subsequent lowering of the medial longitudinal arch) will negatively affect the amount of first metatarsophalangeal joint excursion occurring at push-off. Vertical descent of the navicular (a proxy for subtalar pronation) and first metatarsophalangeal joint dorsal excursion were measured during steady locomotion over a flat substrate on a novel sample consisting of asymptomatic adult males and females, many of whom are habitually unshod. Least-squares regression analyses indicated that, contrary to the hypothesis, navicular drop did not explain a significant amount of variation in first metatarsophalangeal joint dorsal excursion. These results suggest that, in an asymptomatic subject, the plantar aponeurosis and the associated foot bones can function effectively within the normal range of subtalar pronation that takes place during walking gait. From a clinical standpoint, this study highlights the need for investigating the in vivo kinematic relationship between subtalar pronation and metatarsophalangeal joint dorsiflexion in symptomatic populations, and also the need to explore other factors that may affect the kinematics of asymptomatic feet.

The Jinniushan hominin pedal skeleton from the late Middle Pleistocene of China

Bipedalism has long been recognized as the seminal adaptation of the hominin radiation and thus used to distinguish hominins from great ape fossils. Notwithstanding preconceptions and varied interpretations, the distinctive features of the modern human foot and accompanying striding gait, appear to be recent innovations that are largely absent in the earliest facultative bipeds. These distinctive features are mainly components of fixed longitudinal and transverse pedal arches, and of a uniquely derived hallucal metatarsophalangeal joint. They enhance ankle joint plantar flexor function and accommodate localized peak plantar pressures at the medial ball during terminal stance. To date, the paleontological record has yielded very little of the hominin foot, especially of the Middle Pleistocene hominins. New specimens from this time interval should help provide insights into the timing and pattern of what appears to be a mosaic pattern of evolution of the modern human foot features. Here we describe the fossil hominin foot skeleton recovered from the Jinniushan site, Liaoning Province, People's Republic of China. It affords a singular glimpse of the pedal morphology of a late Middle Pleistocene hominin (c.f. Homo heidlebergensis). Dated to 200 ka or older, this foot offers the earliest evidence for increased stability of the medial longitudinal arch, while retaining a number of primitive features apparently characteristic of robust premodern hominins, including lower arches and a less stable hallucal metatarsophalangeal joint (medial ball) than in modern humans. These features reflect different foot capabilities and suggest the bipedal stride of the Jinniushan hominin differed subtlety from that of modern humans.

Ichnotaxonomy of the Laetoli trackways: The earliest hominin footprints

At 3.6 Ma, the Laetoli Pliocene hominin trackways are the earliest direct evidence of hominin bipedalism. Three decades since their discovery, not only is the question of their attribution still discussed, but marked differences in interpretation concerning the footprints’ qualitative features and the inferred nature of the early hominin foot morphology remain. Here, we establish a novel ichnotaxon, Praehominipes laetoliensis, for these tracks and clarify the distinctions of these footprints from those of later hominins, especially modern humans. We also contrast hominin, human, and ape footprints to establish morphological features of these footprints correlated with a midtarsal break versus a stiff longitudinal arch. Original photos, including stereo photographs, and casts of footprints from the 1978 Laetoli excavation, confirm midtarsal flexibility, and repeatedly indicate an associated midfoot pressure ridge. In contrast, the modern human footprint reflects the derived arched-foot architecture, combined with a stiff-legged striding gait. Fossilized footprints of unshod modern human pedestrians in Hawaii and Nicaragua unambiguously illustrate these contrasts. Some points of comparisons with ape footprints are complicated by a variable hallucal position and the distinct manner of ape facultative bipedalism. In contrast to the comparatively rigid platform of the modern human foot, midtarsal flexibility is present in the chimpanzee foot. In ape locomotion, flexion at the transverse tarsal joint, referred to as the “midtarsal break,” uncouples the respective functions of the prehensile forefoot and the propulsive hindfoot during grasp-climbing. At some point after the transition to habitual bipedalism, these grasp-climb adaptations, presumed to be present in the last common ancestor of apes and humans, were initially compromised by the loss of divergence of the hallux. An analogous trajectory is evident along an array of increasingly terrestrial extant ape species. However, a flexible midfoot was retained, presumably to spare lateral toes from bending stresses by concentrating push-off from the forefoot, beneath the metatarsals. Only later did the evolution of the longitudinal arch permit increased mechanical advantage of the plantarflexors for speed and improved economy of endurance distance walking and running.

Early Hominin Foot Morphology Based on 1.5-Million-Year-Old Footprints from Ileret, Kenya

Hominin footprints offer evidence about gait and foot shape, but their scarcity, combined with an inadequate hominin fossil record, hampers research on the evolution of the human gait. Here, we report hominin footprints in two sedimentary layers dated at 1.51 to 1.53 million years ago (Ma) at Ileret, Kenya, providing the oldest evidence of an essentially modern human-like foot anatomy, with a relatively adducted hallux, medial longitudinal arch, and medial weight transfer before push-off. The size of the Ileret footprints is consistent with stature and body mass estimates for Homo ergaster/erectus, and these prints are also morphologically distinct from the 3.75-million-year-old footprints at Laetoli, Tanzania. The Ileret prints show that by 1.5 Ma, hominins had evolved an essentially modern human foot function and style of bipedal locomotion.

The mechanics of the gibbon foot and its potential for elastic energy storage during bipedalism

The mechanics of the modern human foot and its specialization for habitual bipedalism are well understood. The windlass mechanism gives it the required stability for propulsion generation, and flattening of the arch and stretching of the plantar aponeurosis leads to energy saving. What is less well understood is how an essentially flat and mobile foot, as found in protohominins and extant apes, functions during bipedalism. This study evaluates the hypothesis that an energy-saving mechanism, by stretch and recoil of plantar connective tissues, is present in the mobile gibbon foot and provides a two-dimensional analysis of the internal joint mechanics of the foot during spontaneous bipedalism of gibbons using a four-link segment foot model. Available force and pressure data are combined with detailed foot kinematics, recorded with a high-speed camera at 250 Hz, to calculate the external joint moments at the metatarsophalangeal (MP), tarsometatarsal (TM) and talocrural (TC) joints. In addition, instantaneous joint powers are estimated to obtain insight into the propulsion-generating capacities of the internal foot joints. It is found that, next to a wide range of motion at the TC joint, substantial motion is observed at the TM and MP joint, underlining the importance of using a multi-segment foot model in primate gait analyses. More importantly, however, this study shows that although a compliant foot is less mechanically effective for push-off than a ;rigid' arched foot, it can contribute to the generation of propulsion in bipedal locomotion via stretch and recoil of the plantarflexor tendons and plantar ligaments.

HOW GIBBONS WALK ON FLOPPY FEET

![Figure][1] The human foot is a miracle of evolution. We can keep striding for miles on our well-sprung feet. There is nothing else like them, not even amongst our closest living relatives. According to Evie Vereecke, from the University of Liverpool, the modern human foot first appeared

Evolutionary aspects of foot disorders

The modern human foot can be viewed as the basic prehensile grasping foot of humans' arboreal ancestors, modified by the need for a propulsive lever to serve bipedal walking. The structure of the human foot serves its function remarkably well. However, in spite of its evolutionary success, the foot remains susceptible to clinical disorders that can be related to its development from nonhuman ancestors. An understanding of the evolution of the foot offers a different perspective on some common human foot disorders, and can provide a framework for planning the treatment of those disorders.

The evolutionary basis of some clinical disorders of the human foot: a comparative survey of the living primates.

The living primates are a highly diverse group of essentially arboreal animals whose feet are variously adapted for grasping, climbing, and leaping in trees. One of the most remarkable aspects of the anatomical variation in the feet of the extant primates is that this diversity can be arranged in a graduated sequence ranging from the primitive transtarsal-opposition type of grasping foot found in the lemurs, through the specialized transmetarsal-adduction type that characterizes the higher primates, to the unique nongrasping foot of humans. The comparative study of this graded series makes it possible, without recourse to the fossil record, to appreciate the adaptive and functional stages through which the human foot passed in its evolution. It is hypothesized that the initial stage of human erect posture was characterized by a foot which was adapted to both hallucial grasping and short distance bipedal walking. In many respects, the structure of the foot of this primitive human was probably similar to that of the living highland gorilla. When compared with the feet of our closest living relatives, the African apes, the human foot is characterized by two major evolutionary specializations: the longitudinal and transverse arches, and the parallel first and second metatarsals. These two morphological adaptations, together with several associated specializations, are the structural basis of the bipedal human foot. Reconstructing the evolutionary and adaptive history of these specializations creates a better understanding of some of the more common clinical podiatric disorders. The modern human foot is structurally so well adapted to prolonged bipedal walking and standing that even slight deviations from its evolutionarily established pattern will produce debilitating clinical manifestations. In most cases, successful treatment of such disorders involves restoration of the foot's basic adaptive configuration.