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Gravitational forces can induce deviations in body posture from desired configurations in multi-legged arboreal robot locomotion with low leg stiffness, affecting the contact angle between the swing leg's end-effector and the climbing surface during the gait cycle. The relationship between desired and actual foot positions is investigated here in a leg-stiffness-enhanced model under external forces, focusing on the challenge of unreliable end-effector attachment on climbing surfaces in such robots. Inspired by the difference in ceiling attachment postures of dead and living geckos, feedforward compensation of the stance phase legs is the key to solving this problem. A feedforward gravity compensation (FGC) strategy, complemented by leg coordination, is proposed to correct gravity-influenced body posture and improve adhesion stability by reducing body inclination. The efficacy of this strategy is validated using a quadrupedal climbing robot, EF-I, as the experimental platform. Experimental validation on an inverted surface (ceiling walking) highlights the benefits of the FGC strategy, demonstrating its role in enhancing stability and ensuring reliable end-effector attachment without external assistance. In the experiment, robots without FGC only completed 3 out of 10 trials, while robots with FGC achieved a 100% success rate in the same trials. The speed was substantially greater with FGC, achieving 9.2 mm s -1 in the trot gait. This underscores the proposed potential of the FGC strategy in overcoming the challenges associated with inconsistent end-effector attachment in robots with low leg stiffness, thereby facilitating stable locomotion even at an inverted body attitude., (Creative Commons Attribution license.)

In this study we developed an analytical relationship between adhesive digit orientation and adhesive force capacity to describe the tendencies of climbing organisms that use adhesion for climbing to align their toes in the direction of loading, maximizing adhesive force capacity. We fabricated a multi-component adhesive device with multiple contact surfaces, or digits, to act as a model system mimicking the angular motion of a foot and found the synthetic experiments agree with the developed analytical relationship. In turn, we find that observations of gekkonid lizards climbing on vertical substrates correlate well with our analytical relationship; a reduction in toe spacing is seen on the forelimbs when the animals are facing up. Interestingly, the toes on the hindlimbs tend to have an increase in spacing, possibly a mechanism for stabilization rather than load-bearing., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

Anoline lamellae terminate in an epidermal free margin carrying the majority of its setae. How the free margin is extruded from the body of the scale is not well understood. Two hypotheses have been advanced to account for it, one advocating distal migration of the outer epidermal layers relative to the body of the lamella, and the other proposing regression of the dermal core. Available evidence provides partial support for both. We assembled a series of specimens of Anolis grahami representing all shedding cycle stages, and prepared histological sections of the toe pads to allow measurement of appropriate lamellar components through the shedding cycle. Through its proliferative phases the lamellae increase markedly in length, with the distance between the distal tip of the dermal core and that of the lamella accounting for most of this, indicating that epidermal extrusion is responsible for production of the new free margin. The dermal core showed no evidence of regression. Concomitant with epidermal extrusion, the lacunar cells on the inner lamellar face hypertrophy and keep pace with the increasing thickness of the outer lamellar face resulting from the lengthening of the replacement setae. The integrated changes observed are consistent with continuity of functioning and alignment of the exposed setal carpet of the outer epidermal generation while ensuring that the new setal carpet is fully aligned and functional immediately after shedding. At shedding the original proportions of the lamellae are restored. Development of the new free margin results from a combination of distal displacement of Oberhäutchen cells along with arrested maturation of the epidermis in this region. Changes in length of the lamellae during the proliferative stages may impact the overall size of the adhesive toe pad, which may have consequences for assessments of the relationship between whole animal clinging ability and adhesive pad area. J. Morphol. 278:360-368, 2017. © 2017 Wiley Periodicals, Inc., (© 2017 Wiley Periodicals, Inc.)

It has long been thought that quadrupedal primates successfully occupy arboreal environments, in part, by relying on their grasping feet to control balance and propulsion, which frees their hands to test unstable branches and forage. If this interlimb decoupling of function is real, there should be discernible differences in forelimb versus hind limb musculoskeletal control, specifically in how manual and pedal digital flexor muscles are recruited to grasp during arboreal locomotion. New electromyography data from extrinsic flexor muscles in red ruffed lemurs (Varecia rubra) walking on a simulated arboreal substrate reveal that toe flexors are activated at relatively higher levels and for longer durations than finger flexors during stance phase. This demonstrates that the extremities of primates indeed have different functional roles during arboreal locomotion, with the feet emphasizing maintenance of secure grips. When this dichotomous muscle activity pattern between the forelimbs and hind limbs is coupled with other features of primate quadrupedal locomotion, including greater hind limb weight support and the use of diagonal-sequence footfall patterns, a complex suite of biomechanical characters emerges in primates that allow for the co-option of hands toward non-locomotor roles. Early selection for limb functional differentiation in primates probably aided the evolution of fine manipulation capabilities in the hands of bipedal humans., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

When the adhesive toe pads of geckos become wet, they become ineffective in enabling geckos to stick to substrates. This result is puzzling given that many species of gecko are endemic to tropical environments where water covered surfaces are ubiquitous. We hypothesized that geckos can recover adhesive capabilities following exposure of their toe pads to water by walking on a dry surface, similar to the active self-cleaning of dirt particles. We measured the time it took to recover maximum shear adhesion after toe pads had become wet in two groups, those that were allowed to actively walk and those that were not. Keeping in mind the importance of substrate wettability to adhesion on wet surfaces, we also tested geckos on hydrophilic glass and an intermediately wetting substrate (polymethylmethacrylate; PMMA). We found that time to maximum shear adhesion recovery did not differ in the walking groups based on substrate wettability (22.7±5.1 min on glass and 15.4±0.3 min on PMMA) but did have a significant effect in the non-walking groups (54.3±3.9 min on glass and 27.8±2.5 min on PMMA). Overall, we found that by actively walking, geckos were able to self-dry their wet toe pads and regain maximum shear adhesion significantly faster than those that did not walk. Our results highlight a unexpected property of the gecko adhesive system, the ability to actively self-dry and recover adhesive performance after being rendered dysfunctional by water.

Foot and toe clearance (TC) are used regularly to describe locomotor control for both clinical and basic research. However, accuracy of TC during obstacle crossing can be compromised by typical sample frequencies, which do not capture the frame when the foot is over the obstacle due to high limb velocities. The purpose of this study was to decrease the error of TC measures by increasing the spatial resolution of the toe trajectory with interpolation. Five young subjects stepped over an obstacle in the middle of an 8 m walkway. Position data were captured at 600 Hz as a gold standard signal (GS-600-Hz). The GS-600-Hz signal was downsampled to 60 Hz (DS-60-Hz). The DS-60-Hz was then interpolated by either upsampling or an algorithm. Error was calculated as the absolute difference in TC between GS-600-Hz and each of the remaining signals, for both the leading limb and the trailing limb. All interpolation methods reduced the TC error to a similar extent. Interpolation reduced the median error of trail TC from 5.4 to 1.1 mm; the maximum error was reduced from 23.4 to 4.2 mm (16.6-3.8%). The median lead TC error improved from 1.6 to 0.5 mm, and the maximum error improved from 9.1 to 1.8 mm (5.3-0.9%). Therefore, interpolating a 60 Hz signal is a valid technique to decrease the error of TC during obstacle crossing., (Published by Elsevier Ltd.)

This study compared stiffness of the landing leg in ballet dancers performing grand jeté on a sprung floor to leg stiffness during the same movement on a hard floor (wood on concrete). Leg stiffness was calculated as the ratio of vertical ground reaction force (in Newtons) to compression of the lower limb (in meters). Thirteen female dancers were measured for five repetitions each at the point of maximum leg compression while landing grand jeté on both of the surfaces, such that 20 milliseconds of data were represented for each trial. The stiffness of the landing leg at the point of maximum compression was decreased by a mean difference score of 6168.0 N/m ± 11,519.5 N/m on the hard floor compared to the sprung floor. Paired t-test yielded a one-tailed probability of p = 0.038. This effect was seen in 11 of the 13 participants. The finding of increased stiffness of the landing leg in the sprung floor condition suggests that some of the force of landing the leap was absorbed by the surface, and therefore did not need to be absorbed by the landing leg itself. This in turn implies that a sprung dance floor may help to prevent dance-related injuries.

Assessment of the therapeutic potential of interventions to bridge-repair peripheral nerve defects heavily relies on the demonstration of improved functional outcome. In the present study we used CatWalk gait analysis (locomotor-test) and Static Sciatic Index (SSI) (static-toe-spread-test) to assess the behavioural benefits of autologous nerve transplantation (ANT) repair of 2-cm rat sciatic nerve defects (neurotmesis-lesion). A reproducible and standardised rat sciatic nerve crush lesion model (axonotmesis-lesion) was used to assess the extent of recovery supported by maximal axon regeneration (measured by SSI and CatWalk). Animals were behaviourally followed for a period of 10 weeks. SSI analysis showed that ANT induced a significant improvement in motor deficit from about -95 to -65, however, CatWalk analysis did not show any major indication of locomotor recovery. This discrepancy might suggest that improvements in static motor functions (such as toe spreading) could reflect an early indicator for the recovery of function. We also noted differences in axon regeneration including increased axon density, smaller axon diameters and thinner myelin sheaths in the distal region of the ANT in comparison to the equivalent region of crushed and normal nerves. This difference in axon regeneration may be related to the clearly improved toe spreading function. We conclude that SSI and CatWalk present different advantages and disadvantages for the assessment of motor recovery after bridge-repair of peripheral nerve defects., (Copyright © 2010 Elsevier B.V. All rights reserved.)

Geckos are super climbers: they can readily and rapidly stick to almost any surface, whether hydrophilic or hydrophobic, rough or smooth, in dry or wet conditions, and detach with equal rapidity within tens of milliseconds. In this paper, we discuss the rapid switching between the strong adhesion/friction (attached) state and zero adhesion/friction (detached) state, and present a finite element analysis of gecko setae in terms of their adhesion and friction forces. The analysis shows why the asymmetric, naturally curved setae with a directional tilt play a crucial role in the gecko's articulation mechanism, consistent with recent experimental studies of gecko setal arrays. We derive guidelines for designing synthetic versions of gecko adhesive pads, and propose a design for a "gecko-inspired" adhesive surface consisting of arrays of curved, asymmetric, and directionally oriented microfibrils, attached to a semirigid backing, and suggest a method for its actuation.

The evolution of ornithopod dinosaurs provides a well-documented example of the transition from digitigrady to subunguligrady. During this transition, the ornithopod pes was drastically altered from the plesiomorphic dinosaurian morphology (four digits, claw-shaped unguals, strongly concavo-convex joints, phalanges longer than wide, excavated collateral ligament fossae, presence of sagittal ridge, and prominent processes for the attachment of tendons) to a more derived condition (tridactyly, modification of the unguals into hooves, phalanges wider and thinner than long, lack of collateral ligament fossae, loss of sagittal ridge and tendon attachment processes, relatively flattened articular surfaces). These changes are particularly noteworthy given the overall conservatism in pedal morphology seen across Dinosauria. But what are the functional consequences of these specific morphological transitions? To study them, we examine a wide range of pedal morphologies in four non-avian dinosaurs and two birds. Our analyses of the external morphology, two-dimensional models (using Finite Element Analysis), and internal bone structure demonstrate that this evolutionary shift was accompanied by a loss of digit mobility and flexibility. In addition, pedal posture was modified to better align the pes with the main direction of the ground reaction force, thus becoming well suited to support high loads. These conclusions can be applied to other, parallel evolutionary changes (in both dinosaurs and mammals) that involved similar transitions to a subunguligrade posture.

Geckos can run rapidly on walls and ceilings, requiring high friction forces (on walls) and adhesion forces (on ceilings), with typical step intervals of approximately 20 ms. The rapid switching between gecko foot attachment and detachment is analyzed theoretically based on a tape model that incorporates the adhesion and friction forces originating from the van der Waals forces between the submicron-sized spatulae and the substrate, which are controlled by the (macroscopic) actions of the gecko toes. The pulling force of a spatula along its shaft with an angle between theta 0 and 90 degrees to the substrate, has a "normal adhesion force" contribution, produced at the spatula-substrate bifurcation zone, and a "lateral friction force" contribution from the part of spatula still in contact with the substrate. High net friction and adhesion forces on the whole gecko are obtained by rolling down and gripping the toes inward to realize small pulling angles between the large number of spatulae in contact with the substrate. To detach, the high adhesion/friction is rapidly reduced to a very low value by rolling the toes upward and backward, which, mediated by the lever function of the setal shaft, peels the spatulae off perpendicularly from the substrates. By these mechanisms, both the adhesion and friction forces of geckos can be changed over three orders of magnitude, allowing for the swift attachment and detachment during gecko motion. The results have obvious implications for the fabrication of dry adhesives and robotic systems inspired by the gecko's locomotion mechanism.

Geckos may represent the world's most demanding adhesives application. The adhesive setae on the toes of climbing geckos must adhere strongly yet avoid fouling or attachment at inappropriate times. We tested the hypothesis that gecko setae are non-adhesive in their unloaded default state by comparing the water droplet contact angle (theta) of isolated setal arrays to the smooth surface of eye spectacle scales of tokay geckos (Gekko gecko). At equilibrium, theta was 98.3 +/- 3.4 degrees in spectacle scales of live geckos and 93.3 +/- 3.5 degrees in isolated spectacles. Isolated setal arrays were ultrahydrophobic, with theta of 160.6 +/- 1.3 degrees (means +/- SD). The difference in theta of setal arrays and smooth spectacles indicates a very low contact fraction. Using Cassie's law of surface wettability, we infer that less than 6.6% of the surface of unloaded setae is solid and at least 93.4% is air space. We calculated that the contact fraction must increase from 6.6% in the unloaded state to 46% in the loaded state to account for previously measured values of adhesion. Thus gecko setae may be non-sticky by default because only a very small contact fraction is possible without mechanically deforming the setal array.

Adaptive human locomotion is dependent on safe clearance of obstacles encountered in the path of locomotion. When the terrain is uneven or compliant, stability along with safe obstacle clearance are competing demands presented to the central nervous system (CNS). To examine how the CNS deals with the two competing demands, six participants walked under four conditions: normal ground walking, normal ground walking with an obstacle in the travel path, compliant surface walking, and compliant surface walking with an obstacle in the travel path. Full body kinematics were measured and swing limb kinetics were derived from these measurements. Results showed that on a compliant surface, the CNS was able to decrease foot placement variability at foot contact when approaching an obstacle, similar to the normal ground terrain. Limb trajectory over the obstacle showed that toe elevation was maintained while clearance over the obstacle was lower in the compliant surface condition due to depression of the surface during push off. This illustrates that the CNS controls toe elevation, not toe clearance when stepping over an obstacle. Work done in the knee during elevation and hip during lowering was similar in the compliant and ground conditions even though a lower clearance over the obstacle was achieved in the complaint condition. This shows the inability of the CNS to account for compression of the surface prior to obstacle clearance and provides further evidence the CNS controls toe elevation, not clearance when stepping over an obstacle.

This study investigated whether a mirror mechanism exists for bipedal locomotion. We employed the soleus (plantar flexor) Hoffman reflex to investigate corticospinal excitability at the spinal level during observation of bipedal locomotion. The differential amplitude modulation of the left soleus Hoffman reflex during observation of bipedal heel-stepping (plantar dorsiflexion) (324+/-53 microV), standing still (383+/-60 microV), and bipedal toe-stepping (plantar flexion) (419+/-53 microV) reached significance (P < 0.05). The observation of bipedal toe-stepping produced a greater increase in spinal excitability than the observation of bipedal heel-stepping. These findings support the suggestion that there is a mirror mechanism for bipedal locomotion and they demonstrate that spinal excitability for observation of bipedal locomotion mirrors that for execution of bipedal locomotion.

Sole-floor reaction forces were recorded from five anatomically discrete points to analyze characteristics of human locomotion. Strain gauge of 14 mm diameter were firmly attached to the sole of bare-foot for recording force changes from the following five points: (1) medial process of calcaneus, (2) head of 1st metatarsal, (3) head of 3rd metatarsal, (4) head of 5th metatarsal and (5) great toe. Fifteen healthy adults were asked to walk at 2, 4, 6 and 8 km/h and to run at 8 km/h on the treadmill. Sole-floor reaction forces from 1st to 5th metatarsals show reciprocal changes during stance phase, while force from 1st metatarsal is strong 5th metatarsal shows weak reaction and vice versa. This phenomenon may be an expression of locomotor program to maintain vertical stability of the body during stance phase. There was a linear relation between walking speeds and sum of force from the five points, although sum of forces from three metatarsals did not change significantly during the walking speeds, indicating mainly calcaneus and great toe contribute to increasing walking speed. During running the sum of force from the three metatarsals increased sharply, joining the other two points to increase thrust.

The goal of the study was to examine the accuracy and precision of control of adaptive locomotion using haptic information in normally sighted humans before and after practice. Obstacle avoidance paradigm was used to study adaptive locomotion; individuals were required to approach and step over different sizes of obstacles placed in the travel path under three sensory conditions: full vision (FV); restricted lower visual field (RLVF) using blinders on custom glass frames; and no vision (NV) using haptic information only. In the NV condition, individuals were a given an appropriate-sized cane to guide their locomotion. Footfall patterns were recorded using the GAITRite system, and lead and trail limb trajectories were monitored using the OPTOTRAK system, which tracked infrared diodes placed on the toes and the cane. Approach step lengths were reduced for the haptic condition: this slowed the forward progression and allowed greater time for haptic exploration, which ranged from 2.5 to 4 s and consisted of horizontal cane movements (to detect the width and relative location of the obstacle) and vertical cane movements (to detect the height of the obstacle). Based on feed-forward and on-line sensory (under both vision and haptic conditions) information about location of the obstacle relative to the individual, variability of foot placement reduced as the individual came closer to the obstacle, as has been shown in the literature. The only difference was that the reduction in variability of foot placement under haptic condition occurred in the last step compared with earlier under vision. Considering that the obstacle is detected only when the cane comes in contact, as opposed to vision condition when it is visible earlier, this difference is understandable. Variability and magnitude of lead and trail limb elevation for the haptic condition was higher than the RLVF and FV conditions. In contrast, only the magnitude of lead and trail limb elevation was higher in the RLVF condition when compared with the FV condition. This suggests that it is the inability of the haptic sense to provide accurate information about obstacle characteristics compared with the visual system, and not simple caution that lead to higher limb elevation. In the haptic and RLVF condition when vision was unavailable for on-line monitoring of lead limb elevation, kinesthetic information from lead limb elevation was used to fine-tune trail limb elevation. Both the control of approach phase and limb elevation findings held up even after sufficient practice to learn haptic guidance of adaptive locomotion in the second experiment. These results provide a clear picture of the efficacy of the haptic sensory system to guide locomotion in a cluttered environment.

To investigate the specificity of fusimotor (gamma) drive during locomotion, gamma-efferents were recorded from the flexor digitorum longus (FDL) and flexor hallucis longus (FHL) nerves in a decerebrate cat preparation. These nerves innervate hindlimb muscles that differ in some aspects of their mechanical action. For both FHL and FDL two stereotyped patterns of gamma activity were distinguished. Tonic units fired throughout the step cycle and had less modulation, but higher minimum rates, than phasic units, which were mainly recruited with ankle extensor [soleus (SOL)] electromyogram (EMG) activity. Differences in the relative timing of these patterns were apparent. In FHL the activity of phasic and most tonic neurons peaked after EMG onset. With FDL, tonic units generally reached maximum rate before, while phasic units peaked after, the beginning of EMG activity. During locomotion FHL and FDL alpha activity were rhythmically recruited with SOL. However, consistent with previous reports, FHL and FDL differed in their patterns of alpha activity. FHL was stereotyped while FDL was variable. Both FHL and FDL had activity related to ankle extensor EMG, but only FDL exhibited a peak around the end of this phase. No corresponding gamma activity was observed in FDL. In conclusion, 1) FHL and FDL received tonic and phasic fusimotor drive; 2) there was no alpha/gamma linkage for the late FDL alpha burst; 3) phasic gamma-efferents in both muscles received similar inputs, linked to plantar flexor alpha activity; and 4) tonic gamma-efferents differed, to the extent that they were modulated at all. The FHL units peaked with the plantar flexor alphas. The FDL neurons generally peaked before alpha activity even began.

To examine the hydrodynamic propulsion mechanism of a diving great crested grebe (Podiceps cristatus), the three-dimensional kinematics was determined by digital analysis of sequential video images of dorsal and lateral views. During the acceleration phase of this foot-propelled bird, the feet move through an arc in a plane nearly normal to the bird's line of motion through the water, i.e. the toes move dorsally and medially but not caudally relative to the water. The kinematics of the grebe's lobed feet is different from that in anseriforms, whose feet move in a plane mostly parallel to the bird's line of progress through the water. Our results suggest that the foot-propelled locomotor mechanism of grebes is based primarily on a lift-producing leg and foot stroke, in contrast to the drag-based locomotion assumed previously. We suggest that the lift-based paddling of grebes considerably increases both maximum swimming speed and energetic efficiency over drag-based propulsion. Furthermore, the results implicate a new interpretation of the functional morphology of these birds, with the toes serving as a self-stabilizing multi-slotted hydrofoil during the power phase.

This study was designed to test the impact characteristics of the equine digit in vitro with the objective of providing a better understanding of the role of the digital structures in the attenuation of impact shock. Uni-axial accelerometers were mounted on cadaver digits on the distolateral hoof wall, the proximolateral hoof wall, the dorsal surface of the second phalanx, and the mid-lateral first phalanx. The hoof-mounted accelerometers were aligned with the hoof tubules while the bone-mounted accelerometers were oriented along the longitudinal axis of the bone. Each digit was mounted in a test apparatus designed to simulate impact of the hoof with the ground during locomotion. The digits were subjected to 3 impact trials against a barrier at each of 3 vertical impact velocities that simulated a forward trotting velocity in the range of 2.67 to 4.46 m/s. The impact deceleration tended to increase with impact velocity. Attenuation of the impact shock by the digital tissues resulted in a reduction in impact decleration in the more proximal measuring locations. The interphalangeal joints appeared to play a larger role in amplitude attenuation than the hoof wall or the soft tissue structures within the hoof wall. The signal frequency data showed that the soft tissues within the hoof acted as a 'lowpass' filter, attenuating the higher deceleration frequencies. The hoof wall and the interphalangeal joints showed little frequency attenuation.

Additional graphical tools are needed to better visualize the joint kinematics of human locomotion. Standard plots in which the joint displacements are plotted against time or percent gait cycle do not provide sufficient information about the dynamics of the system. In this article, a study based on the two graphical tools of nonlinear dynamics to visualize the steady-state kinematics of human gait is presented. An experimental setup was developed to acquire the necessary data for application of the techniques. Twenty young adults, whose medical histories are free of gait pathology, were tested. Computerized electrogoniometers and foot switches were used to measure the kinematic data of the lower extremities and capture four instants of the gait cycle: heel strike, foot flat, heel off, and toe off. Phase plane portraits of each joint were constructed for the sagittal plane by plotting angular velocity against angular displacement. Poincaré maps were obtained by periodically sampling the joint profiles at toe off and plotting the ith iterate against the (i + 1)th one. Phase plane portraits are useful in monitoring the variations of joint velocity and position on the same graph in a more compact form. Poincaré maps are effective in differentiating steady gait from transient locomotion.

Human feet and toes provide a mechanism for changing the gear ratio of the ankle extensor muscles during a running step. A variable gear ratio could enhance muscle performance during constant-speed running by applying a more effective prestretch during landing, while maintaining the muscles near the high-efficiency or high-power portion of the force-velocity curve during takeoff. Furthermore, during acceleration, variable gearing may allow muscle contractile properties to remain optimized despite rapid changes in running speed. Forceplate and kinematic analyses of running steps show low gear ratios at touchdown that increase throughout the contact phase.

The gait cycle is usually analyzed using one of two conventions: heel contact initiates the cycle and stance is followed by swing phases or toe-off initiates the cycle and swing then stance phases are analyzed. The purpose of the study is to demonstrate that based on a consideration of relevant biomechanical measures, a convention identifying toe-off as the beginning of the gait cycle and presenting swing followed by stance phases is the superior convention. Net joint torques and electromyographic (EMG) data from selected muscles in the lower extremity were obtained from four subjects while walking and running. Data were collected for consecutive swing, stance and swing phases to compare the variables at the swing-to-stance and stance-to-swing transitions. Larger joint torques were observed at the swing-to-stance transition at the hip and knee for both gaits compared to the other transition. EMG results showed greater activation levels for five of the six muscles at the swing-to-stance transition. Results indicated that the subjects needed to prepare for the initiation of stance and the application of relatively large external forces and moments. Further, the transition from stance to swing did not seem to be as critical a point in the gait cycle since the movements and EMG were relatively low. This being the case, the stance-to-swing transition should be used as the beginning and ending of the gait cycle (toe-off initiating the cycle) and the more meaningful transition of swing-to-stance phases should occur in the middle of the analysis.

The importance of well-functioning toes has long been recognised but has not previously been assessed in biomechanical studies. We have examined the weight-bearing function of the foot in 160 normal subjects by use of the pedobarograph. The function of the toes was assessed by reference to the time they were in contact with the ground and the peak pressures they exerted individually in comparison with other parts of the foot. The toes were in contact for about three-quarters of the stance phase of gait and exerted peak pressures similar to those of the metatarsal region. When the foot was bearing the second peak of total force, the area in contact with the ground (the metatarsal heads and toes) was decreasing.

The foot has been studied in walking, running and jogging using high speed cinema photography in the laboratory for gait analysis in the Shriner's Hospital for Crippled Children, San Francisco. During running, as well as in walking, the extrinsic toe flexors restrain progression of the tibia. The extrinsic extensors accelerate the tibia over the fixed foot. The intrinsic toe muscles function from mid-stance to lift-off when the subject runs on his toes. In sprinting, the intrinsics are active throughout the weight-bearing phase.