Mechanics of the human foot during walking on different slopes.

When humans walk on slopes, the ankle, knee, and hip joints modulate their mechanical work to accommodate the mechanical demands. Yet, it is unclear if the foot modulates its work output during uphill and downhill walking. Therefore, we quantified the mechanical work performed by the foot and its subsections of twelve adults walked on five randomized slopes (-10°, -5°, 0°, +5°, +10°). We estimated the work of distal-to-hindfoot and distal-to-forefoot structures using unified deformable segment analysis and the work of the midtarsal, ankle, knee, and hip joints using a six-degree-of-freedom model. Further, using a geometric model, we estimated the length of the plantar structures crossing the longitudinal arch while accounting for the first metatarsophalangeal wrapping length. We hypothesized that compared to level walking, downhill walking would increase negative and net-negative work magnitude, particularly at the early stance phase, and uphill walking would increase the positive work, particularly at the mid-to-late stance phase. We found that downhill walking increased the magnitude of the foot's negative and net-negative work, especially during early stance, highlighting its capacity to absorb impacts when locomotion demands excessive energy dissipation. Notably, the foot maintained its net dissipative behavior between slopes; however, the ankle, knee, and hip shifted from net energy dissipation to net energy generation when changing from downhill to uphill. Such results indicate that humans rely more on joints proximal to the foot to modulate the body's total mechanical energy. Uphill walking increased midtarsal's positive and distal-to-forefoot negative work in near-equal amounts. That coincided with the prolonged lengthening and delayed shortening of the plantar structures, resembling a spring-like function that possibly assists the energetic demands of locomotion during mid-to-late stance. These results broaden our understanding of the foot's mechanical function relative to the leg's joints and could inspire the design of wearable assistive devices that improve walking capacity.

Does human foot anthropometry relate to plantar flexor fascicle mechanics and metabolic energy cost across various walking speeds?

Foot structures define the leverage in which the ankle muscles push off against the ground during locomotion. While prior studies have indicated that inter-individual variation in anthropometry (e.g. heel and hallux lengths) can directly affect force production of ankle plantar flexor muscles, its effect on the metabolic energy cost of locomotion has been inconclusive. Here, we tested the hypotheses that shorter heels and longer halluces are associated with slower plantar flexor (soleus) shortening velocity and greater ankle plantar flexion moment, indicating enhanced force potential as a result of the force-velocity relationship. We also hypothesized that such anthropometry profiles would reduce the metabolic energy cost of walking at faster walking speeds. Healthy young adults (N=15) walked at three speeds (1.25, 1.75 and 2.00 m s-1), and we collected in vivo muscle mechanics (via ultrasound), activation (via electromyography) and whole-body metabolic energy cost of transport (via indirect calorimetry). Contrary to our hypotheses, shorter heels and longer halluces were not associated with slower soleus shortening velocity or greater plantar flexion moment. Additionally, longer heels were associated with reduced metabolic cost of transport, but only at the fastest speed (2.00 m s-1, R2=0.305, P=0.033). We also found that individuals with longer heels required less increase in plantar flexor (soleus and gastrocnemius) muscle activation to walk at faster speeds, potentially explaining the reduced metabolic cost.

Does the Heel's Dissipative Energetic Behavior Affect Its Thermodynamic Responses During Walking?

Most of the terrestrial legged locomotion gaits, like human walking, necessitate energy dissipation upon ground collision. In humans, the heel mostly performs net-negative work during collisions, and it is currently unclear how it dissipates that energy. Based on the laws of thermodynamics, one possibility is that the net-negative collision work may be dissipated as heat. If supported, such a finding would inform the thermoregulation capacity of human feet, which may have implications for understanding foot complications and tissue damage. Here, we examined the correlation between energy dissipation and thermal responses by experimentally increasing the heel's collisional forces. Twenty healthy young adults walked overground on force plates and for 10 min on a treadmill (both at 1.25 ms-1) while wearing a vest with three different levels of added mass (+0%, +15%, & +30% of their body mass). We estimated the heel's work using a unified deformable segment analysis during overground walking. We measured the heel's temperature immediately before and after each treadmill trial. We hypothesized that the heel's temperature and net-negative work would increase when walking with added mass, and the temperature change is correlated with the increased net-negative work. We found that walking with +30% added mass significantly increased the heel's temperature change by 0.72 ± 1.91   ℃ (p = 0.009) and the magnitude of net-negative work (extrapolated to 10 min of walking) by 326.94 ± 379.92 J (p = 0.005). However, we found no correlation between the heel's net-negative work and temperature changes (p = 0.277). While this result refuted our second hypothesis, our findings likely demonstrate the heel's dynamic thermoregulatory capacity. If all the negative work were dissipated as heat, we would expect excessive skin temperature elevation during prolonged walking, which may cause skin complications. Therefore, our results likely indicate that various heat dissipation mechanisms control the heel's thermodynamic responses, which may protect the health and integrity of the surrounding tissue. Also, our results indicate that additional mechanical factors, besides energy dissipation, explain the heel's temperature rise. Therefore, future experiments may explore alternative factors affecting thermodynamic responses, including mechanical (e.g., sound & shear-stress) and physiological mechanisms (e.g., sweating, local metabolic rate, & blood flow).

Walking with added mass magnifies salient features of human foot energetics.

The human foot serves numerous functional roles during walking, including shock absorption and energy return. Here, we investigated walking with added mass to determine how the foot would alter its mechanical work production in response to a greater force demand. Twenty-one healthy young adults walked with varying levels of added body mass: 0%, +15% and +30% (relative to their body mass). We quantified mechanical work performed by the foot using a unified deformable segment analysis and a multi-segment foot model. We found that walking with added mass tended to magnify certain features of the foot's functions. Magnitudes of both positive and negative mechanical work, during stance in the foot, increased when walking with added mass. Yet, the foot preserved similar amounts of net negative work, indicating that the foot dissipates energy overall. Furthermore, walking with added mass increased the foot's negative work during early stance phase, highlighting the foot's role as a shock-absorber. During mid to late stance, the foot produced greater positive work when walking with added mass, which coincided with greater work from the structures spanning the midtarsal joint (i.e. arch). While this study captured the overall behavior of the foot when walking with varying force demands, future studies are needed to further determine the relative contribution of active muscles and elastic tissues to the foot's overall energy.

Ankle and midtarsal joint quasi-stiffness during walking with added mass.

Examination of how the ankle and midtarsal joints modulate stiffness in response to increased force demand will aid understanding of overall limb function and inform the development of bio-inspired assistive and robotic devices. The purpose of this study is to identify how ankle and midtarsal joint quasi-stiffness are affected by added body mass during over-ground walking. Healthy participants walked barefoot over-ground at 1.25 m/s wearing a weighted vest with 0%, 15% and 30% additional body mass. The effect of added mass was investigated on ankle and midtarsal joint range of motion (ROM), peak moment and quasi-stiffness. Joint quasi-stiffness was broken into two phases, dorsiflexion (DF) and plantarflexion (PF), representing approximately linear regions of their moment-angle curve. Added mass significantly increased ankle joint quasi-stiffness in DF (p < 0.001) and PF (p < 0.001), as well as midtarsal joint quasi-stiffness in DF (p < 0.006) and PF (p < 0.001). Notably, the midtarsal joint quasi-stiffness during DF was ~2.5 times higher than that of the ankle joint. The increase in midtarsal quasi-stiffness when walking with added mass could not be explained by the windlass mechanism, as the ROM of the metatarsophalangeal joints was not correlated with midtarsal joint quasi-stiffness (r = -0.142, p = 0.540). The likely source for the quasi-stiffness modulation may be from active foot muscles, however, future research is needed to confirm which anatomical structures (passive or active) contribute to the overall joint quasi-stiffness across locomotor tasks.