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.

Cultural variation in running techniques among non-industrial societies.

Research among non-industrial societies suggests that body kinematics adopted during running vary between groups according to the cultural importance of running. Among groups in which running is common and an important part of cultural identity, runners tend to adopt what exercise scientists and coaches consider to be good technique for avoiding injury and maximising performance. In contrast, among groups in which running is not particularly culturally important, people tend to adopt suboptimal technique. This paper begins by describing key elements of good running technique, including landing with a forefoot or midfoot strike pattern and leg oriented roughly vertically. Next, we review evidence from non-industrial societies that cultural attitudes about running associate with variation in running techniques. Then, we present new data from Tsimane forager-horticulturalists in Bolivia. Our findings suggest that running is neither a common activity among the Tsimane nor is it considered an important part of cultural identity. We also demonstrate that when Tsimane do run, they tend to use suboptimal technique, specifically landing with a rearfoot strike pattern and leg protracted ahead of the knee (called overstriding). Finally, we discuss processes by which culture might influence variation in running techniques among non-industrial societies, including self-optimisation and social learning.

Does plantar skin abrasion affect cutaneous mechanosensation?

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.

Forest terrains influence walking kinematics among indigenous Tsimane of the Bolivian Amazon.

Laboratory-based studies indicate that a major evolutionary advantage of bipedalism is enabling humans to walk with relatively low energy expenditure. However, such studies typically record subjects walking on even surfaces or treadmills that do not represent the irregular terrains our species encounters in natural environments. To date, few studies have quantified walking kinematics on natural terrains. Here we used high-speed video to record marker-based kinematics of 21 individuals from a Tsimane forager-horticulturalist community in the Bolivian Amazon walking on three different terrains: a dirt field, a forest trail and an unbroken forest transect. Compared with the field, in the unbroken forest participants contacted the ground with more protracted legs and flatter foot postures, had more inclined trunks, more flexed hips and knees, and raised their feet higher during leg swing. In contrast, kinematics were generally similar between trail and field walking. These results provide preliminary support for the idea that irregular natural surfaces like those in forests cause humans to alter their walking kinematics, such that travel in these environments could be more energetically expensive than would be assumed from laboratory-based data. These findings have important implications for the evolutionary energetics of human foraging in environments with challenging terrains.

Effects of footwear cushioning on leg and longitudinal arch stiffness during running.

During running, humans increase leg stiffness on more compliant surfaces through an in-series spring relationship to maintain constant support mechanics. Following this notion, the compliant midsole material of standard footwear may cause individuals to increase leg stiffness while running, especially in footwear with very thick midsoles. Recently, researchers have also proposed that footwear stiffness can affect the stiffness of the foot's longitudinal arch (LA) via a similar mechanism. To test these ideas, we used 3D motion capture to record 20 participants running on a forceplate-instrumented treadmill while barefoot, and while wearing three types of sandals composed of materials ranging an order of magnitude in Young's modulus: ethylene vinyl acetate (EVA), and two varieties of polyurethane rubber (R30 and R60). We calculated leg stiffness using standard methods, and measured LA stiffness based on medial midfoot kinematics. While there was an overall significant effect of footwear on leg stiffness (P = 0.047), post-hoc tests revealed no significant differences among individual pairs of conditions, and there was no effect of footwear on LA stiffness. However, participants exhibited significantly greater LA compression when barefoot than when running in EVA (P = 0.004) or R30 (P = 0.036) sandals. These results indicate that standard footwear midsole materials are too stiff to appreciably affect leg stiffness during running, meaning that increasing midsole thickness is unlikely to cause individuals to alter their leg stiffness. However, use of footwear does cause individuals to restrict LA compression when compared to running barefoot, and further research is needed to understand why.

Trunk muscle endurance, strength and flexibility in rural subsistence farmers and urban industrialized adults in western Kenya.

OBJECTIVES: High trunk muscle endurance, strength, and moderate flexibility reportedly help maintain musculoskeletal health, but there is evidence for tradeoffs among these variables as well as sex differences in trunk muscle endurance and strength. To test if these observations extend similarly to both men and women in nonindustrial and industrial environments, we investigated intra-individual associations and group and sex differences in trunk muscle endurance, strength, and flexibility among 74 (35 F, 39 M; age range: 18-61 years) adults from the same Kalenjin-speaking population in western Kenya. We specifically compared men and women from an urban community with professions that do not involve manual labor with rural subsistence farmers, including women who frequently carry heavy loads. METHODS: Trunk muscle endurance, strength, and flexibility were measured with exercise tests and electromyography (EMG). RESULTS: We found a positive correlation between trunk extensor strength and endurance (R = .271, p ≤ .05) and no associations between strength or endurance and flexibility. Rural women had higher trunk extensor and flexor endurance, EMG-determined longissimus lumborum endurance, and trunk extensor strength than urban women (all p ≤ .05). Rural women had higher trunk extensor and flexor endurance than rural men (both p ≤ .05). Urban women had lower trunk flexor and extensor endurance than urban men (both p ≤ .01). CONCLUSIONS: High levels of physical activity among nonindustrial subsistence farmers, particularly head carrying among women, appear to be associated with high trunk muscle endurance and strength, which may have important benefits for helping maintain musculoskeletal health.

The energetics of uniquely human subsistence strategies.

The suite of derived human traits, including enlarged brains, elevated fertility rates, and long developmental periods and life spans, imposes extraordinarily high energetic costs relative to other great apes. How do human subsistence strategies accommodate our expanded energy budgets? We found that relative to other great apes, human hunter-gatherers and subsistence farmers spend more energy but less time on subsistence, acquire substantially more energy per hour, and achieve similar energy efficiencies. These findings revise our understanding of human energetic evolution by indicating that humans afford expanded energy budgets primarily by increasing rates of energy acquisition, not through energy-saving adaptations such as economical bipedalism or sophisticated tool use that decrease subsistence costs and improve the energetic efficiency of subsistence. We argue that the time saved by human subsistence strategies provides more leisure time for social interaction and social learning in central-place locations and would have been critical for cumulative cultural evolution.

Urbanization and Knee Cartilage Growth Among Children and Adolescents in Western Kenya.

OBJECTIVE: Previous studies have demonstrated that low physical activity levels during youth are associated with the development of thin knee cartilage, which may increase susceptibility to osteoarthritis later in life. Here, we propose and test the hypothesis that reductions in physical activity impair knee cartilage growth among people in developing countries experiencing urbanization and increased market integration. METHODS: Ultrasonography was used to measure knee cartilage thickness in 168 children and adolescents (aged 8-17 years) from two groups in western Kenya: a rural, physically active group from a small-scale farming community and an urban, less physically active group from the nearby city of Eldoret. We used general linear models to assess the relative effects of age on cartilage thickness in these two groups, controlling for sex and leg length. RESULTS: Both groups exhibited significant reductions in knee cartilage thickness with increasing age (P < 0.0001; 95% confidence interval [CI] 0.15-0.06 mm), yet the rate of reduction was significantly less in the rural than in the urban group (P = 0.012; 95% CI 0.01-0.10 mm). CONCLUSION: The results support our hypothesis by showing that individuals from the more physically active rural group exhibited less knee cartilage loss during youth than the more sedentary urban group. Our findings suggest that reduced physical activity associated with urbanization in developing nations may affect adult knee cartilage thickness and thus could be a factor that increases susceptibility to osteoarthritis.

The human foot functions like a spring of adjustable stiffness during running.

Like other animals, humans use their legs like springs to save energy during running. One potential contributor to leg stiffness in humans is the longitudinal arch (LA) of the foot. Studies of cadaveric feet have demonstrated that the LA can function like a spring, but it is unknown whether humans can adjust LA stiffness in coordination with more proximal joints to help control leg stiffness during running. Here, we used 3D motion capture to record 27 adult participants running on a forceplate-instrumented treadmill, and calculated LA stiffness using beam bending and midfoot kinematics models of the foot. Because changing stride frequency causes humans to adjust overall leg stiffness, we had participants run at their preferred frequency and frequencies 35% above and 20% below preferred frequency to test for similar adjustments in the LA. Regardless of which foot model we used, we found that participants increased LA quasi-stiffness significantly between low and high frequency runs, mirroring changes at the ankle, knee and leg overall. However, among foot models, we found that the model incorporating triceps surae force into bending force on the foot produced unrealistically high LA work estimates, leading us to discourage this modeling approach. Additionally, we found that there was not a consistent correlation between LA height and quasi-stiffness values among the participants, indicating that static LA height measurements are not good predictors of dynamic function. Overall, our findings support the hypothesis that humans dynamically adjust LA stiffness during running in concert with other structures of the leg.

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.

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.

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 years1, the oldest recovered footwear dates to eight thousand years ago2 and inexpensive shoes with cushioned heels were not developed until the Industrial Revolution3. 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.

Step width and frontal plane trunk motion in bipedal chimpanzee and human walking.

Human bipedalism is characterized by mediolateral oscillations of the center of mass (CoM) between the feet. The preferred step widths and CoM oscillations used by humans likely represent a trade-off of several factors (e.g., stance and swing phase costs). However, it is difficult to assess whether human frontal plane control strategies are unique given few detailed data on frontal plane motion during facultative bipedalism in apes. Here, we collected three-dimensional kinematic and kinetic data in humans and chimpanzees to investigate the relationship between step width, mediolateral CoM motion, frontal plane trunk kinematics, and CoM power during bipedalism. Chimpanzee bipedalism entails mediolateral CoM oscillations and step widths that are (scaled to lower/hind limb length) three times larger than those of humans. Chimpanzees use a combination of linear and angular motion of the trunk and list the entire trunk, and especially thorax, over the stance side foot, generating large mediolateral shifts in the CoM, whereas humans utilize little angular motion within the trunk. Larger mediolateral CoM motions do not have a significant effect on CoM power. Similarities between bipedal chimpanzees and other bipedal non-human primates (macaques and gibbons) indicate that narrow CoM motions are unique to humans and are likely due to our adducted hips and valgus knees. Valgus knees appear early in the human fossil record (∼3.6 Ma), contemporaneous with the Laetoli footprints. However, fossils attributed to Ardipithecus ramidus (∼4.4 Ma) suggest that the earliest hominins may have lacked a hominin-like degree of knee valgus. If correct, this suggests that this species may have used wide steps, larger mediolateral CoM motions, and perhaps larger trunk motions during bipedal walking. Finally, we present a novel means to estimate mediolateral CoM motion from trackway step width, and estimate that the Laetoli G track maker used CoM motions within the human range.

Locomotor constraints favour the evolution of the human pygmy phenotype in tropical rainforests.

The convergent evolution of the human pygmy phenotype in tropical rainforests is widely assumed to reflect adaptation in response to the distinct ecological challenges of this habitat (e.g. high levels of heat and humidity, high pathogen load, low food availability, and dense forest structure), yet few precise adaptive benefits of this phenotype have been proposed. Here, we describe and test a biomechanical model of how the rainforest environment can alter gait kinematics such that short stature is advantageous in dense habitats. We hypothesized that environmental constraints on step length in rainforests alter walking mechanics such that taller individuals are expected to walk more slowly due to their inability to achieve preferred step lengths in the rainforest. We tested predictions from this model with experimental field data from two short-statured populations that regularly forage in the rainforest: the Batek of Peninsular Malaysia and the Tsimane of the Bolivian Amazon. In accordance with model expectations, we found stature-dependent constraints on step length in the rainforest and concomitant reductions in walking speed that are expected to compromise foraging efficiency. These results provide the first evidence that the human pygmy phenotype is beneficial in terms of locomotor performance and highlight the value of applying laboratory-derived biomechanical models to field settings for testing evolutionary hypotheses.

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.

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.

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.

Chimpanzee ankle and foot joint kinematics: Arboreal versus terrestrial locomotion.

OBJECTIVES: Many aspects of chimpanzee ankle and midfoot joint morphology are believed to reflect adaptations for arboreal locomotion. However, terrestrial travel also constitutes a significant component of chimpanzee locomotion, complicating functional interpretations of chimpanzee and fossil hominin foot morphology. Here we tested hypotheses of foot motion and, in keeping with general assumptions, we predicted that chimpanzees would use greater ankle and midfoot joint ranges of motion during travel on arboreal supports than on the ground. METHODS: We used a high-speed motion capture system to measure three-dimensional kinematics of the ankle and midfoot joints in two male chimpanzees during three locomotor modes: terrestrial quadrupedalism on a flat runway, arboreal quadrupedalism on a horizontally oriented tree trunk, and climbing on a vertically oriented tree trunk. RESULTS: Chimpanzees used relatively high ankle joint dorsiflexion angles during all three locomotor modes, although dorsiflexion was greatest in arboreal modes. They used higher subtalar joint coronal plane ranges of motion during terrestrial and arboreal quadrupedalism than during climbing, due in part to their use of high eversion angles in the former. Finally, they used high midfoot inversion angles during arboreal locomotor modes, but used similar midfoot sagittal plane kinematics across all locomotor modes. DISCUSSION: The results indicate that chimpanzees use large ranges of motion at their various ankle and midfoot joints during both terrestrial and arboreal locomotion. Therefore, we argue that chimpanzee foot anatomy enables a versatile locomotor repertoire, and urge caution when using foot joint morphology to reconstruct arboreal behavior in fossil hominins.

Chimpanzee super strength and human skeletal muscle evolution.

Since at least the 1920s, it has been reported that common chimpanzees (Pan troglodytes) differ from humans in being capable of exceptional feats of "super strength," both in the wild and in captive environments. A mix of anecdotal and more controlled studies provides some support for this view; however, a critical review of available data suggests that chimpanzee mass-specific muscular performance is a more modest 1.5 times greater than humans on average. Hypotheses for the muscular basis of this performance differential have included greater isometric force-generating capabilities, faster maximum shortening velocities, and/or a difference in myosin heavy chain (MHC) isoform content in chimpanzee relative to human skeletal muscle. Here, we show that chimpanzee muscle is similar to human muscle in its single-fiber contractile properties, but exhibits a much higher fraction of MHC II isoforms. Unlike humans, chimpanzee muscle is composed of ∼67% fast-twitch fibers (MHC IIa+IId). Computer simulations of species-specific whole-muscle models indicate that maximum dynamic force and power output is 1.35 times higher in a chimpanzee muscle than a human muscle of similar size. Thus, the superior mass-specific muscular performance of chimpanzees does not stem from differences in isometric force-generating capabilities or maximum shortening velocities-as has long been suggested-but rather is due in part to differences in MHC isoform content and fiber length. We propose that the hominin lineage experienced a decline in maximum dynamic force and power output during the past 7-8 million years in response to selection for repetitive, low-cost contractile behavior.