Foot strength and stiffness are related to footwear use in a comparison of minimally- vs. conventionally-shod populations

The longitudinal arch (LA) helps stiffen the foot during walking, but many people in developed countries suffer from flat foot, a condition characterized by reduced LA stiffness that can impair gait. Studies have found this condition is rare in people who are habitually barefoot or wear minimal shoes compared to people who wear conventional modern shoes, but the basis for this difference remains unknown. Here we test the hypothesis that the use of shoes with features that restrict foot motion (e.g. arch supports, toe boxes) is associated with weaker foot muscles and reduced foot stiffness. We collected data from minimally-shod men from northwestern Mexico and men from urban/suburban areas in the United States who wear ‘conventional’ shoes. We measured dynamic LA stiffness during walking using kinematic and kinetic data, and the cross-sectional areas of three intrinsic foot muscles using ultrasound. Compared to conventionally-shod individuals, minimally-shod individuals had higher and stiffer LAs, and larger abductor hallucis and abductor digiti minimi muscles. Additionally, abductor hallucis size was positively associated with LA stiffness during walking. Our results suggest that use of conventional modern shoes is associated with weaker intrinsic foot muscles that may predispose individuals to reduced foot stiffness and potentially flat foot

Heel impact forces during barefoot versus minimally shod walking among Tarahumara subsistence farmers and urban Americans

Despite substantial recent interest in walking barefoot and in minimal footwear, little is known about potential differences in walking biomechanics when unshod versus minimally shod. To test the hypothesis that heel impact forces are similar during barefoot and minimally shod walking, we analysed ground reaction forces recorded in both conditions with a pedography platform among indigenous subsistence farmers, the Tarahumara of Mexico, who habitually wear minimal sandals, as well as among urban Americans wearing commercially available minimal sandals. Among both the Tarahumara (n = 35) and Americans (n = 30), impact peaks generated in sandals had significantly (p < 0.05) higher force magnitudes, slower loading rates and larger vertical impulses than during barefoot walking. These kinetic differences were partly due to individuals' significantly greater effective mass when walking in sandals. Our results indicate that, in general, people tread more lightly when walking barefoot than in minimal footwear. Further research is needed to test if the variations in impact peaks generated by walking barefoot or in minimal shoes have consequences for musculoskeletal health

Effect of the upward curvature of toe springs on walking biomechanics in humans

Although most features of modern footwear have been intensively studied, there has been almost no research on the effects of toe springs. This nearly ubiquitous upward curvature of the sole at the front of the shoe elevates the toe box dorsally above the ground and thereby holds the toes in a constantly dorsiflexed position. While it is generally recognized that toe springs facilitate the forefoot’s ability to roll forward at the end of stance, toe springs may also have some effect on natural foot function. This study investigated the effects of toe springs on foot biomechanics in a controlled experiment in which participants walked in specially-designed sandals with varying curvature in the toe region to simulate toe springs ranging from 10 to 40 degrees of curvature. Using inverse dynamics techniques, we found that toe springs alter the joint moments and work at the toes such that greater degrees of toe spring curvature resulted in lower work requirements during walking. Our results help explain why toe springs have been a pervasive feature in shoes for centuries but also suggest that toe springs may contribute to weakening of the foot muscles and possibly to increased susceptibility to common pathological conditions such as plantar fasciitis

Foot callus thickness does not trade off protection for tactile sensitivity during walking

Until relatively recently, humans, similar to other animals, were habitually barefoot. Therefore, the soles of our feet were the only direct contact between the body and the ground when walking. There is indirect evidence that footwear such as sandals and moccasins were first invented within the past 40 thousand years , the oldest recovered footwear dates to eight thousand years ago and inexpensive shoes with cushioned heels were not developed until the Industrial Revolution . Because calluses—thickened and hardened areas of the epidermal layer of the skin—are the evolutionary solution to protecting the foot, we wondered whether they differ from shoes in maintaining tactile sensitivity during walking, especially at initial foot contact, to improve safety on surfaces that can be slippery, abrasive or otherwise injurious or uncomfortable. Here we show that, as expected, people from Kenya and the United States who frequently walk barefoot have thicker and harder calluses than those who typically use footwear. However, in contrast to shoes, callus thickness does not trade-off protection, measured as hardness and stiffness, for the ability to perceive tactile stimuli at frequencies experienced during walking. Additionally, unlike cushioned footwear, callus thickness does not affect how hard the feet strike the ground during walking, as indicated by impact forces. Along with providing protection and comfort at the cost of tactile sensitivity, cushioned footwear also lowers rates of loading at impact but increases force impulses, with unknown effects on the skeleton that merit future study. 1 2

Evolutionary anatomy of the plantar aponeurosis in primates, including humans

The plantar aponeurosis in the human foot has been extensively studied and thoroughly described, in part, because of the incidence of plantar fasciitis in humans. It is commonly assumed that the human plantar aponeurosis is a unique adaptation to bipedalism that evolved in concert with the longitudinal arch. However, the comparative anatomy of the plantar aponeurosis is poorly known in most mammals, even among non-human primates, hindering efforts to understand its function. Here, we review previous anatomical descriptions of 40 primate species and use phylogenetic comparative methods to reconstruct the evolution of the plantar aponeurosis and its relationship to the plantaris muscle in primates. Ancestral state reconstructions suggest that the overall organization of the human plantar aponeurosis is shared with chimpanzees and that a similar anatomical configuration evolved independently in different primate clades as an adaptation to terrestrial locomotion. The presence of a plantar aponeurosis with clearly developed lateral and central bands in the African apes suggests that this structure is not prohibitive to suspensory locomotion and that these species possess versatile feet adapted for both terrestrial and arboreal locomotion. This plantar aponeurosis configuration would have been advantageous in enhancing foot stiffness for bipedal locomotion in the earliest hominins, prior to the evolution of a longitudinal arch. Hominins may have subsequently evolved thicker and stiffer plantar aponeuroses alongside the arch to enable a windlass mechanism and elastic energy storage for bipedal walking and running, although this idea requires further testing

Thoracic adaptations for ventilation during locomotion in humans and other mammals

Bipedal humans, like canids and some other cursorial mammals, are thought to have been selected for endurance running, which requires the ability to sustain aerobic metabolism over long distances by inspiring large volumes of air for prolonged periods of time. Here, we tested the general hypothesis that humans and other mammals selected for vigorous endurance activities evolved derived thoracic features to increase ventilatory capacity. To do so, we investigated whether humans and dogs rely on thoracic motion to increase tidal volume during running to a greater extent than goats, a species that was not selected for endurance locomotion. We found that while all three species use diaphragmatic breathing to increase tidal volume with increasing oxygen demand, humans also use both dorsoventral and mediolateral expansion of the thorax. Dogs use increased dorsoventral expansion of the thorax, representing an intermediate between humans and goats. 3D analyses of joint morphology of 10 species across four mammalian orders also showed that endurance-adapted cursorial species independently evolved more concavo-convex costovertebral joint morphologies that allow for increased rib mobility for thoracic expansion. Evidence for similarly derived concavo-convex costovertebral joints in Homo erectus corresponds with other evidence for the evolution of endurance running in the genus Homo

Form and function of the human and chimpanzee forefoot: implications for early hominin bipedalism

During bipedal walking, modern humans dorsiflex their forefoot at the metatarsophalangeal joints (MTPJs) prior to push off, which tightens the plantar soft tissues to convert the foot into a stiff propulsive lever. Particular features of metatarsal head morphology such as “dorsal doming” are thought to facilitate this stiffening mechanism. In contrast, chimpanzees are believed to possess MTPJ morphology that precludes high dorsiflexion excursions during terrestrial locomotion. The morphological affinity of the metatarsal heads has been used to reconstruct locomotor behavior in fossil hominins, but few studies have provided detailed empirical data to validate the assumed link between morphology and function at the MTPJs. Using three-dimensional kinematic and morphometric analyses, we show that humans push off with greater peak dorsiflexion angles at all MTPJs than do chimpanzees during bipedal and quadrupedal walking, with the greatest disparity occurring at MTPJ 1. Among MTPJs 2–5, both species exhibit decreasing peak angles from medial to lateral. This kinematic pattern is mirrored in the morphometric analyses of metatarsal head shape. Analyses of Australopithecus afarensis metatarsals reveal morphology intermediate between humans and chimpanzees, suggesting that this species used different bipedal push-off kinematics than modern humans, perhaps resulting in a less efficient form of bipedalism

Mobility of the human foot's medial arch helps enable upright bipedal locomotion.

Developing the ability to habitually walk and run upright on two feet is one of the most significant transformations to have occurred in human evolution. Many musculoskeletal adaptations enabled bipedal locomotion, including dramatic structural changes to the foot and, in particular, the evolution of an elevated medial arch. The foot's arched structure has previously been assumed to play a central role in directly propelling the center of mass forward and upward through leverage about the toes and a spring-like energy recoil. However, it is unclear whether or how the plantarflexion mobility and height of the medial arch support its propulsive lever function. We use high-speed biplanar x-ray measurements of foot bone motion on seven participants while walking and running and compare their motion to a subject-specific model without arch recoil. We show that regardless of intraspecific differences in medial arch height, arch recoil enables a longer contact time and favorable propulsive conditions at the ankle for walking upright on an extended leg. The generally overlooked navicular-medial cuneiform joint is primarily responsible for arch recoil in human arches. The mechanism through which arch recoil enables an upright ankle posture may have helped drive the evolution of the longitudinal arch after our last common ancestor with chimpanzees, who lack arch plantarflexion mobility during push-off. Future morphological investigations of the navicular-medial cuneiform joint will likely provide new interpretations of the fossil record. Our work further suggests that enabling medial arch recoil in footwear and surgical interventions may be critical for maintaining the ankle's natural propulsive ability

Does plantar skin abrasion affect cutaneous mechanosensation?

Abstract In humans, plantar cutaneous mechanoreceptors provide critical input signals for postural control during walking and running. Because these receptors are located within the dermis, the mechanical properties of the overlying epidermis likely affect the transmission of external stimuli. Epidermal layers are highly adaptable and can form hard and thick protective calluses, but their effects on plantar sensitivity are currently disputed. Some research has shown no effect of epidermal properties on sensitivity to vibrations, whereas other research suggests that vibration and touch sensitivity diminishes with a thicker and harder epidermis. To address this conflict, we conducted an intervention study where 26 participants underwent a callus abrasion while an age‐matched control group (n = 16) received no treatment. Skin hardness and thickness as well as vibration perception thresholds and touch sensitivity thresholds were collected before and after the intervention. The Callus abrasion significantly decreased skin properties. The intervention group exhibited no change in vibration sensitivity but had significantly better touch sensitivity. We argue that touch sensitivity was impeded by calluses because hard skin disperses the monofilament's standardized pressure used to stimulate the mechanoreceptors over a larger area, decreasing indentation depth and therefore stimulus intensity. However, vibration sensitivity was unaffected because the vibrating probe was adjusted to reach specific indentation depths, and thus stimulus intensity was not affected by skin properties. Since objects underfoot necessarily indent plantar skin during weight‐bearing, calluses should not affect mechanosensation during standing, walking, or running

Supporting data from Heel impact forces during barefoot versus minimally shod walking among Tarahumara subsistence farmers and urban Americans

Spreadsheet of data collected and analysed