Influence of the structure of the support surface under the sole on vertical posture during standing with different body weight distributions between legs

The vertical posture was studied during standing with fееt on the support surfaces of different structures. The movements of the center of pressure (CP) of each leg and the common CP (CCP) were recorded while the subject stood with a support on a smooth floor and with the support of one foot on a spike mat (SM) with different load distributions between the legs. When the body weight was transferred to one leg during standing under ordinary conditions on a smooth floor, the CP of the loaded leg moved more than the CP of the unloaded leg; i.e., the posture sway was compensated mainly due to the activity of the loaded leg, which created a larger torque. When the subject stood with one foot on the SM, the CP movement of this leg did not depend on the leg load and was about 60% of the CP movement of the leg on the smooth floor. Apparently, the CP displacement of the unloaded leg on smooth support was larger than the CP displacement of the loaded leg creating the torque necessary for compensating the body sway. Thus, maintaining the vertical posture was carried out mainly by the leg standing on the smooth support. It is assumed that additional stimulation of different surface and deep receptors of the foot caused by foot support on the SM hampered the perception of its CP position, and the vertical posture was maintained mainly by the leg afferent signals from which more precisely reflected the CP position.     

Characteristics of the maintenance of the vertical posture during standing with an asymmetrical load on the legs

Movements of the common center of pressure (CP) and the CPs of the right and left legs separately were studied during the maintenance of the vertical posture by subjects standing with symmetrical load on their legs or with the shift of the load to the right or left leg. It was shown that standing with a symmetrical load on the legs was accompanied by the movement of the CP of an individual leg along the straight line with small deviations aside, whereas movement of the common CP represented the curve with frequent changes in direction and filling up some space. The shift of the load to one leg resulted in the movement of the CP of the loaded leg that was similar to that observed during a symmetrical load on the legs. The movement of the CP of the unloaded leg was chaotic. The shift of the load to one leg decreased the correlation between the movements of the CPs of the left and right legs compared to standing with a symmetrical load on the legs. The velocity of movement of the CP of the leg loaded increased in the sagittal direction but remained stable in the frontal direction. The velocity of movement of the CP of the unloaded leg remained stable in the sagittal direction but increased in the frontal direction. We suppose that during standing with an asymmetrical load on the legs the role of the single in the maintenance of the vertical posture depend on the load on the leg.     

Maintenance of Human Vertical Posture upon Asymmetric Leg Loading and Fixation of the Knee Joint of One Leg

Maintenance of a vertical posture was studied in standing subjects with a fixed knee joint of one leg and a different weight distribution between the legs. Knee fixation on one leg did not affect the speed of movements of the common center of pressure (CP) at any weight distribution between the legs, and the stability of vertical posture was therefore unchanged. However, the relative contributions of the legs to the posture control changed when knee movements of one leg were restricted. The speed of CP movements of the free leg was independent of the weight loading on the leg. The speed of CP movements of the leg with the knee fixed depended on the weight distribution and was higher when the leg was loaded. Thus, the leg with the fixed knee joint made a greater contribution to maintaining vertical posture when the leg was loaded. Yet its contribution was comparable with that of the unloaded free contralateral leg even in this case, as was evident from lack of differences in CP movements between the two legs. It was assumed that the leg with the free knee joint played a major role in maintaining equilibrium of vertical posture, while the leg with the fixed knee joint mostly acted to more finely adjust the body position.     

Influence of Achilles tendon vibration on the vertical posture during standing with asymmetrical leg loading

The shift of the common center of pressure (CCP) and the center of pressure (CP) of one leg was studied during the Achilles tendon vibration of one or both legs while the subject was standing with symmetrical load on the legs or with the load transferred to one leg. The CP shift of the standing subject during unilateral Achilles tendon vibration depended on both the side of application of vibration and on the distribution of the leg load. During standing with a asymmetrical load on the legs, the shift of the CCP was larger than when the vibration was applied to the loaded leg. The CP shift of one leg was greater if both vibration and the load were applied to it. Vibration of the unloaded leg caused a CP shift in the loaded contralateral leg. In this case, vibration of the left unloaded leg did not cause any noticeable CP shift of the left leg, while vibration of the unloaded right leg caused a CP shift of the right leg. Under the similar conditions of loading and vibration, the displacement of the CP of the right leg was larger than the displacement of the CP of the left leg. It may be suggested that postural asymmetry and unilateral vibration of the leg muscles change the internal representation of the position of the body axis in relation to the vertical, which affects the displacement of the CP of one leg in response to afferent stimulation of the leg muscles.     

Reaction to the perturbation of the vertical posture under different conditions of standing and in the case of additional support

The reaction of equilibrium restoration in response to the perturbation of the vertical posture of a subject standing on a stable or unstable support was studied. Perturbation was induced by a sudden forward or backward shift of the support surface. In some of the experiments, the subject was holding onto a handrail suspended on a long soft belt with a fixed upper end. The results of the study showed that the reaction to support movement depended on the direction of the perturbation. The soleus muscle was activated first upon a backward movement, and the anterior tibial muscle was activated first upon a forward movement, with a latency of about 50 ms. Within 30–70 ms, bursts of activity were also detected in the respective antagonist muscles. Sudden movement of the platform caused bursts of impulses in the arm muscles even in the absence of contact with the handrail. These impulse bursts had a longer latency (80–130 ms) and probably played an auxiliary role in the process of the restoration of balance. In the case of standing on an unstable support, the latency of leg muscle activation increased. When a subject was holding onto a handrail, the intensity of impulse bursts decreased in the leg muscles and increased in the arm muscles, while the latent periods of the bursts in the arm muscles decreased. This effect proved to be still more pronounced in the case of balance maintenance on an unstable support. Thus, the change in the response to external perturbations during maintenance of the vertical posture on an unstable support demonstrates that an additional contact of the hand changes the adjustment of the posture control system.     

Characteristics of sit-to-stand and gait coordination in patients with total hip replacement

Upright posture, standing up from a chair, and gait were analyzed in patients after one-sided total hip replacement and in healthy subjects (control). It was found that the patients predominantly loaded the unoperated leg when they stood quietly or rose from a chair. Subjects’ walking on a 10-m podograph treadmill showed that their walking speed was slower than that of healthy subjects and the swing phase on the side of hip replacement was longer than on the unoperated side. It was assumed that the unequal load on legs during walking, standing, and sit-to-stand performance in patients with total hip replacement was related to the sensory deficit of the artificial joint, leading to the overstrain of the unoperated leg and coxarthrosis in it.     

Locomotion of the Eurasian nuthatch on vertical and horizontal substrates

Legged locomotion of the Eurasian nuthatch Sitta europaea on horizontal and vertical substrates was examined using field observations and experiments. Although previous studies have reported that nuthatches use 'walking' on vertical substrates, we found that they usually used 'hopping' on both vertical and horizontal substrates. When climbing up a vertical substrate, the feet were staggered in position and small phase differences were observed between the left and right leg movements in the gait. In upward climbing, the body was inclined towards the substrate during the first stance phase similar to other tree-trunk climbers, but the tail was not used for helping body rotation unlike most tree-trunk climbers. The staggered position of the feet may allow the legs to play different roles in pulling towards and pushing away from the substrate. In downward climbing, the feet were staggered in position, but the phase difference was quite small. In field observations, the Eurasian nuthatch preferred to move vertically, rather than in an inclined direction.     

Common motor mechanisms support body load in serially homologous legs of cockroaches in posture and walking

We studied the mechanisms underlying support of body load in posture and walking in serially homologous legs of cockroaches. Activities of the trochanteral extensor muscle in the front or middle legs were recorded neurographically while animals were videotaped. Body load was increased via magnets attached to the thorax and varied through a coil below the substrate. In posture, tonic firing of the slow trochanteral extensor motoneuron (Ds) in each leg was strongly modulated by changing body load. Rapid load increases produced decreases in body height and sharp increments in extensor firing. The peak of extensor activity more closely approximated the maximum velocity of body displacement than the body position. In walking, extensor bursts in front and middle legs were initiated during swing and continued into the stance phase. Moderate tonic increases in body load elicited similar, specific, phase dependent changes in both legs: extensor firing was not altered in swing but was higher after foot placement in stance. These motor adjustments to load are not anticipatory but apparently depend upon sensory feedback. These data are consistent with previous findings in the hind legs and support the idea that body load is countered by common motor mechanisms in serially homologous legs.     

Sensing the effect of body load in legs: responses of tibial campaniform sensilla to forces applied to the thorax in freely standing cockroaches

Sense organs in the legs that detect body weight are an important component in the regulation of posture and locomotion. We tested the abilities of tibial campaniform sensilla, receptors that can monitor forces in the cockroach leg, to encode variations in body load in freely standing animals. Small magnets were attached to the thorax and currents were applied to a coil below the substrate. Sensory and motor activities were monitored neurographically. The tibial sensilla could show vigorous discharges to changing forces when animals stood upon their legs and actively supported the body weight. Firing of individual afferents depended upon the orientation of the receptors cuticular cap: proximal sensilla (oriented perpendicular to the leg axis) discharged to force increases while distal receptors (parallel to the leg) fired to decreasing forces. Proximal sensillum discharges were prolonged and could encode the level of load when increases were sustained. Firing of the trochanteral extensor motoneuron was also strongly modulated by changing load. In some postures, sensillum discharges paralleled changes in motor frequency consistent with a known interjoint reflex. These findings demonstrate that tibial campaniform sensilla can monitor the effects of body weight upon the legs and may aid in generating support of body load.     

Influences of overall thermal balance on local inputs for drive of evaporation in men

Three kinds of experiments were carried out in a climatic chamber: experiments with warm load on the whole body at 36 degrees C (4 subjects); experiments at 36 degrees C with reduction of thermal load (28 degrees C) on the left leg (right leg at 36 degrees C) (8 subjects); and experiments at 36 degrees C with antisymmetric thermal load on the legs of 44 degrees C (right leg) and 28 degrees C (left leg), which resulted in additional thermal loads of +/- 30 W/leg (8 subjects). The additional thermal loads, which were applied via two climatic boxes, produced measurable effects on sweat rate when applied to one leg only. In comparison to the experiment 1, experiment 2 brought about a significant reduction of local evaporation on the left leg. With antisymmetric thermal loads on both legs (experiment 3), which did not influence the overall thermal balance, there was no significant influence on local evaporation, although significant changes of local temperatures were measured. It is suggested that the well-known regulatory models, declaring local, mean skin, and core temperatures as local evaporation drive should be supplemented with an important additional feature: local control of evaporation by local skin temperature may be blocked by an overall thermal balance.     

Characteristic Features of Disorders of the Upright Posture in Patients with Poststroke Hemipareses

Characteristic features of upright posture maintenance and mechanisms of postural disorders in poststroke hemiparetic patients were studied using a bilateral force platform. The following features of postural disorders were revealed in the patients tested: an increase in the velocity and amplitude of the center-of-pressure (CP) sway as compared to in healthy subjects, an absolute decrease in the half-cycles of the CP sway, asymmetry of weight bearing by both feet, and a shift of the center of pressure of an affected foot towards the toe. The disturbance of stability of the vertical posture in such patients is to a greater extent associated with weight-bearing asymmetry. It was shown that the character of the CP sway is mainly determined by a disorder of the sensory motor control, whereas damage to the efferent pathways is responsible for the postural asymmetry. Increase in the muscle tone restricts the sway amplitude. Thus, several forms of postural instability are characteristic of hemiparetic patients. Predominantly sensory, motor, or tonic disorders are responsible for these disturbances of stability.     

On Heels and Toes: How Ants Climb with Adhesive Pads and Tarsal Friction Hair Arrays

Ants are able to climb effortlessly on vertical and inverted smooth surfaces. When climbing, their feet touch the substrate not only with their pretarsal adhesive pads but also with dense arrays of fine hairs on the ventral side of the 3^rd and 4^th tarsal segments. To understand what role these different attachment structures play during locomotion, we analysed leg kinematics and recorded single-leg ground reaction forces in Weaver ants (Oecophylla smaragdina) climbing vertically on a smooth glass substrate. We found that the ants engaged different attachment structures depending on whether their feet were above or below their Centre of Mass (CoM). Legs above the CoM pulled and engaged the arolia (‘toes’), whereas legs below the CoM pushed with the 3^rd and 4^th tarsomeres (‘heels’) in surface contact. Legs above the CoM carried a significantly larger proportion of the body weight than legs below the CoM. Force measurements on individual ant tarsi showed that friction increased with normal load as a result of the bending and increasing side contact of the tarsal hairs. On a rough sandpaper substrate, the tarsal hairs generated higher friction forces in the pushing than in the pulling direction, whereas the reverse effect was found on the smooth substrate. When the tarsal hairs were pushed, buckling was observed for forces exceeding the shear forces found in climbing ants. Adhesion forces were small but not negligible, and higher on the smooth substrate. Our results indicate that the dense tarsal hair arrays produce friction forces when pressed against the substrate, and help the ants to push outwards during horizontal and vertical walking.     

Responses of the lower limb to load carrying in walking man

Muscle activity patterns of several lower limb muscles were examined in the left leg of normal human subjects walking at comfortable speed on a treadmill. In addition knee angular changes and the durations of the swing and stance phases of the step cycle were recorded. Data were collected during a period of normal control walking and when the subject carried a load, either in his right or left hand or on his back. Load (up to 20% of body weight) carried in either hand caused minimal changes in the kinematic parameters investigated but evoked significant prolongation of the normal ongoing electromyographic activity in the contralateral Gluteus medius and in the ipsilateral Gastrocnemius, Vastus lateralis and Semimembranosus. Load (up to 50% of body weight) carried on the back significantly shortened the swing phase and prolonged the ongoing electromyographic activity of the Vastus lateralis. These findings would seem to indicate that the activity of the leg musculature during walking is so tightly controlled that deviation from the normal kinematic pattern of the legs is largely prevented even when body posture and balance are disturbed by carrying substantial additional load.     

The locomotion of the camel (Camelus dromedarius)

The feet and gaits of many camels Camelus dromedarius were studied and filmed in Mauritania, Africa. The camel has a digitigrade stance, large feet to support the animal in soft sand, and soles of flexible pads that step readily onto small stones where necessary. The walking stride is long and slow, with the body supported for much of each stride on the two right or two left legs. The pattern of supporting legs was significantly different in slow compared to fast walking camels, and in young compared to adult camels and compared to adults pulling water at the wells. There was no difference in pattern in one individual's walk, when it was either loaded or unloaded. The angles that the leg bones made with each other and with the horizon are depicted for the walk and the pace. The camel is the only animal which paces often and never trots. The pace is an unstable gait only suitable for flat terrain such as that in deserts. It may have evolved from the pace-like walk which is by far the dominant gait in this animal, which spends most of each day walking from plant to plant browsing or grazing. The pace is not used by all camelids, as one author has claimed. The pace and the gallop were only used by the camels at wells, when the animals were chased from the water by men.     

Intersegmental transfer of sensory signals in the stick insect leg muscle control system

Intersegmental coordination during locomotion in legged animals arises from mechanical couplings and the exchange of neuronal information between legs. Here, the information flow from a single leg sense organ of the stick insect Cuniculina impigra onto motoneurons and interneurons of other legs was investigated. The femoral chordotonal organ (fCO) of the right middle leg, which measures posture and movement of the femur-tibia joint, was stimulated, and the responses of the tibial motoneuron pools of the other legs were recorded. In resting animals, fCO signals did not affect motoneuronal activity in neighboring legs. When the locomotor system was activated and antagonistic motoneurons were bursting in alternation, fCO stimuli facilitated transitions from flexor to extensor activity and vice versa in the contralateral leg. Following pharmacological treatment with picrotoxin, a blocker of GABA-ergic inhibition, the tibial motoneurons of all legs showed specific responses to signals from the middle leg fCO. For the contralateral middle leg we show that fCO signals encoding velocity and position of the tibia were processed by those identified local premotor nonspiking interneurons known to contribute to posture and movement control during standing and voluntary leg movements. Interneurons received both excitatory and inhibitory inputs, so that the response of some interneurons supported the motoneuronal output, while others opposed it. Our results demonstrate that sensory information from the fCO specifically affects the motoneuronal activity of other legs and that the layer of premotor nonspiking interneurons is a site of interaction between local proprioceptive sensory signals and proprioceptive signals from other legs.     

Postural control in healthy young adults using a double seesaw device

Postural control on single and double seesaws was investigated in young healthy adults required to stand as still as possible on two side-by-side seesaws favoring pitch motion and lying on two separate force platforms. The device offers the possibility to get associated or dissociated seesaws and, if dissociated, to induce asymmetric patterns for the centers-of-pressure (CP) under both left and right feet by using different radii for the two seesaws. Substituting a parallelepiped volume to one seesaw offering a firm contact to one foot is also possible. The results indicated that dissociating the two seesaws led to increased resultant CP (CP_Res) and vertically projected center-of-gravity movements (CG_v) only along the mediolateral axis, whereas a slight decreasing tendency characterized these movements along the antero-posterior axis. When standing on two independent seesaws with different radii, significantly larger CP displacements were seen along the antero-posterior axis under the foot lying on the more stable support, i.e., the seesaw with the longer radius or the parallelepiped volume. In these two asymmetrical conditions, the CP_Res output results from a compensatory mechanism, i.e. larger movements under one foot to compensate for the decreased movements occurring under the opposite foot. This postural control strategy is aimed at allowing sufficient CP_Res displacements in order to appropriately secure balance. Because of the complex sensorimotor coordination induced, involving differentially in certain cases both legs, the double seesaw device can be viewed as a possible tool for challenging postural control by inducing asymmetrical patterns between left and right feet CP movements.     

Influence of an asymmetrical body weight distribution on the control of undisturbed upright stance

Postural asymmetry in humans is generally associated with different pathologies. However, its specific influence on undisturbed upright stance is poorly understood. To evaluate its separate effects on each support, the centre of pressure (CP) displacements were recorded through two force platforms. In a second step, the complex resultant centre of pressure trajectories (CP(Res)) were computed and decomposed into two elementary components: the horizontal displacements of the centre of gravity (CG(h)) and the difference in the plane of support between the vertical projection of CG(h) and CP(Res) (CP-CG(v)). These motions were then processed through a frequency analysis and modelled as fractional Brownian motion to gain some additional insight into their spatio-temporal organisation. Ten healthy adults were tested in three conditions consisting of various weight distributions. The quality of the mechanism involved in the control of the unloaded support CP motions appears to decrease as the asymmetry becomes more pronounced. To be precise, larger increases of the CP displacements are observed for the unloaded support compared to the loaded one. As a result, the CP(Res) motions are themselves augmented in the ML direction, inducing in turn larger CG(h) and CP-CG(v) motions. Postural asymmetry thus constitutes an important constraint on the control of upright undisturbed stance by generating changes in the control of both supports and by reducing the efficiency of the hip load/unload mechanisms. On the other hand, by inducing larger body sways, postural asymmetry necessitates higher energy expenditure and the setting of particular control mechanisms.     

A robotic device for understanding neuromechanical interactions during standing balance control

Postural stability in standing balance results from the mechanics of body dynamics as well as active neural feedback control processes. Even when an animal or human has multiple legs on the ground, active neural regulation of balance is required. When the postural configuration, or stance, changes, such as when the feet are placed further apart, the mechanical stability of the organism changes, but the degree to which this alters the demands on neural feedback control for postural stability is unknown. We developed a robotic system that mimics the neuromechanical postural control system of a cat in response to lateral perturbations. This simple robotic system allows us to study the interactions between various parameters that contribute to postural stability and cannot be independently varied in biological systems. The robot is a 'planar', two-legged device that maintains compliant balance control in a variety of stance widths when subject to perturbations of the support surface, and in this sense reveals principles of lateral balance control that are also applicable to bipeds. Here we demonstrate that independent variations in either stance width or delayed neural feedback gains can have profound and often surprisingly detrimental effects on the postural stability of the system. Moreover, we show through experimentation and analysis that changing stance width alters fundamental mechanical relationships important in standing balance control and requires a coordinated adjustment of delayed feedback control to maintain postural stability.     

Effect of an additional manual motor task on the maintenance of equilibrium in the frontal and sagittal planes in standing subjects

The postural oscillations of a standing subject during an additional manual motor task consisting in holding a movable ball in the center of a flat box were studied. The movements of the center of pressure (CP) in the frontal and sagittal planes were studied when subjects were standing on a stable rigid support and on a movable unstable support. The effect of the additional motor task on the movement of the CP depended on the stability of the support. When the additional task was performed, the sagittal movements of the CP increased in the case a movable support and did not increase when the support was stable. The additional task decreased the frontal movements of CP in the case of a stable support, and it did not change the frontal movements of CP when the support was unstable. Thus, the performance of an additional motor task led to a reduction of the efficiency of the postural control system in maintaining equilibrium on an unstable support. This decrease may be due to a greater cortical influence on the posture control system in subjects standing on a movable support in comparison with this influence in the case of a stable support.     

Kinematics of walking in the hermit crab,Pagurus pollicarus

Hermit crabs are decapod crustaceans that have adapted to life in gastropod shells. Among their adaptations are modifications to their thoracic appendages or pereopods. The 4th and 5th pairs are adapted for shell support; walking is performed with the 2nd and 3rd pereopods, with an alternation of diagonal pairs. During stance, the walking legs are rotated backwards in the pitch plane. Two patterns of walking were studied to compare them with walking patterns described for other decapods, a lateral gait, similar to that in many brachyurans, and a forward gait resembling macruran walking.Video sequences of free walking and restrained animals were used to obtain leg segment positions from which joint angles were calculated. Leading legs in a lateral walk generated a power stroke by flexion of MC and PD joints; CB angles often did not change during slow walks. Trailing legs exhibited extension of MC and PD with a slight levation of CB. The two joints, B/IM and CP, are aligned at 90° angles to CB, MC and PD, moving dorso-anteriorly during swing and ventro-posteriorly during stance. A forward step was more complex; during swing the leg was rotated forward (yaw) and vertically (pitch), due to the action of TC. At the beginning of stance, TC started to rotate posteriorly and laterally, CB was depressed, and MC flexed. As stance progressed and the leg was directed laterally, PD and MC extended, so that at the end of stance the dactyl tip was quite posterior. During walks of the animal out of its shell, the legs were extended more anterior-laterally and the animal often toppled over, indicating that during walking in a shell its weight stabilized the animal.An open chain kinematic model in which each segment was approximated as a rectangular solid, the dimensions of which were derived from measurements on animals, was developed to estimate the CM of the animal under different load conditions. CM was normally quite anterior; removal of the chelipeds shifted it caudally. Application of forces simulating the weight of the shell on the 5th pereopods moved CM just anterior to the thoracic-abdominal junction. However, lateral and vertical coordinates were not altered under these different load conditions. The interaction of the shell aperture with proximal leg joints and with the CM indicates that the oblique angles of the legs, due primarily to the rotation of the TC joints, is an adaptation that confers stability during walking.