Head-bobbing of walking birds

Many birds show a rhythmic forward and backward movement of their heads when they walk on the ground. This so-called “head-bobbing” is characterized by a rapid forward movement (thrust phase) which is followed by a phase where the head keeps its position with regard to the environment but moves backward with regard to the body (hold phase). These head movements are synchronized with the leg movements. The functional interpretations of head-bobbing are reviewed. Furthermore, it is discussed why some birds do bob their head and others do not.     

Segment and joint angles of hind limb during bipedal and quadrupedal walking of the bonobo (Pan paniscus)

We describe segment angles (trunk, thigh, shank, and foot) and joint angles (hip, knee, and ankle) for the hind limbs of bonobos walking bipedally ("bent-hip bent-knee walking," 17 sequences) and quadrupedally (33 sequences). Data were based on video recordings (50 Hz) of nine subjects in a lateral view, walking at voluntary speed. The major differences between bipedal and quadrupedal walking are found in the trunk, thigh, and hip angles. During bipedal walking, the trunk is approximately 33-41 degrees more erect than during quadrupedal locomotion, although it is considerably more bent forward than in normal human locomotion. Moreover, during bipedal walking, the hip has a smaller range of motion (by 12 degrees ) and is more extended (by 20-35 degrees ) than during quadrupedal walking. In general, angle profiles in bonobos are much more variable than in humans. Intralimb phase relationships of subsequent joint angles show that hip-knee coordination is similar for bipedal and quadrupedal walking, and resembles the human pattern. The coordination between knee and ankle differs much more from the human pattern. Based on joint angles observed throughout stance phase and on the estimation of functional leg length, an efficient inverted pendulum mechanism is not expected in bonobos.     

Effect of walking speed on inter-joint coordination differs between young and elderly adults

Investigating inter-joint coordination at different walking speeds in young and elderly adults could provide insights to age-related changes in neuromuscular control of gait. We examined effects of walking speed and age on the pattern and variability of inter-joint coordination. Gait analyses of 10 young and 10 elderly adults were performed with different self-selected speeds, including a preferred, faster, and slower speed. Continuous relative phase (CRP), derived from phase planes of two adjacent joints, was used to assess the inter-joint coordination. CRP patterns were examined with cross-correlation measures and root-mean-square (RMS) differences when comparing ensemble mean curves of the faster or slower speed to preferred speed walking. Variability of coordination for each participant was assessed with the average value of all standard deviations calculated for each data point over a gait cycle from all CRP curves, namely the deviation phase (DP). For hip-knee CRP pattern, RMS differences were significantly greater between the slower and preferred walking speeds than between the faster and preferred walking speeds in young adults, but this was not found in elderly adults. Significant group differences in RMS differences and cross-correlation measures were detected in hip-knee CRP patterns between the slower and preferred walking speeds. No significant walking speed or age effects were detected for the knee-ankle CRP. Significant walking speed effects were also detected in hip-knee DP values. However, no significant group differences were detected for all three speeds. These findings suggested that young and elder adults compromise changes of walking speed with different neuromuscular control strategies.     

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.     

The control of walking movements in the leg of the rock lobster

1. Experiments with rock lobsters walking on a treadmill were undertaken to obtain information upon the system controlling the movement of the legs. Results show that the position of the leg is an important parameter affecting the cyclic movement of the walking leg. Stepping can be interrupted when the geometrical conditions for terminating either a return stroke or a power stroke are not fullfilled. 2. The mean value of anterior and posterior extreme positions (AEP and PEP respectively) of the walking legs do not depend on the walking speed (Fig. 1). 3. When one leg is isolated from the other walking legs by placing it on a platform the AEPs and PEPs of the other legs show a broader distribution compared to controls (Figs. 2 and 3). 4. Force measurements (Fig. 4) are in agreement with the hypothesis that the movement of the leg is controlled by a position servomechanism. 5. When one leg stands on a stationary force transducer this leg develops forces which oscillate with the step rhythm of the other legs (Fig. 5). 6. A posteriorly directed influence is found, by which the return stroke of a leg can be started when the anterior leg performs a backward directed movement. 7. Results are compared with those obtained from stick insects. The systems controlling the movement of the individual leg are similar in both, lobster and stick insect but the influences between the legs seem to be considerably different.     

Leg stiffness increases with speed to modulate gait frequency and propulsion energy

Bipedal walking models with compliant legs have been employed to represent the ground reaction forces (GRFs) observed in human subjects. Quantification of the leg stiffness at varying gait speeds, therefore, would improve our understanding of the contributions of spring-like leg behavior to gait dynamics. In this study, we tuned a model of bipedal walking with damped compliant legs to match human GRFs at different gait speeds. Eight subjects walked at four different gait speeds, ranging from their self-selected speed to their maximum speed, in a random order. To examine the correlation between leg stiffness and the oscillatory behavior of the center of mass (CoM) during the single support phase, the damped natural frequency of the single compliant leg was compared with the duration of the single support phase. We observed that leg stiffness increased with speed and that the damping ratio was low and increased slightly with speed. The duration of the single support phase correlated well with the oscillation period of the damped complaint walking model, suggesting that CoM oscillations during single support may take advantage of resonance characteristics of the spring-like leg. The theoretical leg stiffness that maximizes the elastic energy stored in the compliant leg at the end of the single support phase is approximated by the empirical leg stiffness used to match model GRFs to human GRFs. This result implies that the CoM momentum change during the double support phase requires maximum forward propulsion and that an increase in leg stiffness with speed would beneficially increase the propulsion energy. Our results suggest that humans emulate, and may benefit from, spring-like leg mechanics.     

Effects of neck and circumoesophageal connective lesions on posture and locomotion in the cockroach

Few studies in arthropods have documented to what extent local control centers in the thorax can support locomotion in absence of inputs from head ganglia. Posture, walking, and leg motor activity was examined in cockroaches with lesions of neck or circumoesophageal connectives. Early in recovery, cockroaches with neck lesions had hyper-extended postures and did not walk. After recovery, posture was less hyper-extended and animals initiated slow leg movements for multiple cycles. Neck lesioned individuals showed an increase in walking after injection of either octopamine or pilocarpine. The phase of leg movement between segments was reduced in neck lesioned cockroaches from that seen in intact animals, while phases in the same segment remained constant. Neither octopamine nor pilocarpine initiated changes in coordination between segments in neck lesioned individuals. Animals with lesions of the circumoesophageal connectives had postures similar to intact individuals but walked in a tripod gait for extended periods of time. Changes in activity of slow tibial extensor and coxal depressor motor neurons and concomitant changes in leg joint angles were present after the lesions. This suggests that thoracic circuits are sufficient to produce leg movements but coordinated walking with normal motor patterns requires descending input from head ganglia.Electronic Supplementary Material Supplementary material is available for this article at     

Walking in Aretaon asperrimus

This article describes basic parameters characterizing walking of the stick insect Aretaon asperrimus to allow a comparative approach with other insects studied. As in many other animals, geometrical parameters such as step amplitude and leg extreme positions do not vary with walking velocity. However, the relation between swing duration and stance duration is quite constant, in contrast to most insects studied. Therefore, velocity profiles during swing vary with walking velocity whereas time course of leg trajectories and leg angle trajectories are independent of walking velocity. Nevertheless, A. asperrimus does not show a classical tripod gait, but performs a metachronal, or tetrapod, gait, showing phase values differing from 0.5 between ipsilateral neighbouring legs. As in Carausius morosus, the detailed shape of the swing trajectory may depend on the form of the substrate. Effects describing coordinating influences between legs have been found that prevent the start of a swing as long as the posterior leg performs a swing. Further, the treading on tarsus reflex can be observed in Aretaon. No hint to the existence of a targeting influence has been found. Control of rearward walking is easiest interpreted by maintaining the basic rules but an anterior-posterior reversal of the information flow.     

Insect rope-walkers: kinematics of walking on thin rods in a bug, Graphosoma italicum (Heteroptera, Pentatomidae)

Leg positions during walking on a plane and on thin rods were recorded by photography, videorecording, and videokymography. Joint angles were reconstructed from the tibia-ending position, using a 3-D model of the body. Participation of leg joints in propulsion was analysed by calculating the partial derivatives of tibia end-point position on different joint angles. Adjustment to walking with a narrow ground base is achieved by additional femur depression and flexion of the tibia in the stance phase. In the swing phase, the leg is raised by the same amount as when walking on a plane, but not to the same superior position, as on a plane. The contribution of the subcoxal joint to body propulsion is 64-94% in fore-and middle legs and 22-49% in hind legs. The oblique alignment of the coxal pivot within the thorax helps maintain a long stride for variable ground bases. In Graphosoma , it is close to the optimal position: according to several criteria, the angle between the coxal axis and the body vertical shall be arctan π/2, or ∼ 57.5°.     

The effects of sensory loss and walking speed on the orbital dynamic stability of human walking

Peripheral sensory feedback is believed to contribute significantly to maintaining walking stability. Patients with diabetic peripheral neuropathy have a greatly increased risk of falling. Previously, we demonstrated that slower walking speeds in neuropathic patients lead to improved local dynamic stability. However, all subjects exhibited significant local instability during walking, even though no subject fell or stumbled during testing. The present study was conducted to determine if and how significant changes in peripheral sensation and walking speed affect orbital stability during walking. Trunk and lower extremity kinematics were examined from two prior experiments that compared patients with significant neuropathy to healthy controls and walking at multiple different speeds in young healthy subjects. Maximum Floquet multipliers were computed for each time series to quantify the orbital stability of these movements. All subjects exhibited orbitally stable walking kinematics, even though these same kinematics were previously shown to be locally unstable. Differences in orbital stability between neuropathic and control subjects were small and, with the exception of knee joint movements (p=0.001), not statistically significant (0.380p0.946). Differences in knee orbital stability were not mediated by differences in walking speed. This was supported by our finding that although orbital stability improved slightly with slower walking speeds, the correlations between walking speed and orbital stability were generally weak (r(2)16.7%). Thus, neuropathic patients do not gain improved orbital stability as a result of slowing down and do not experience any loss of orbital stability because of their sensory deficits.     

Bipedal locomotion: effects of speed, size and limb posture in birds and humans

Seven species of ground-dwelling birds (body mass range: 0.045-90 kg) were filmed while walking and running on a treadmill. High-speed light films were also taken of humans to compare kinematic patterns of avian with human bipedalism. Consistent patterns of stride frequency, stride length, step length, duty factor and limb excursion were observed in all species, with most of the variation among species being due to differences in body size. In general, smaller bipeds have higher stride frequencies (α M ^−0.18), shorter stride lengths (α M ^0.38) and more limited ranges of speed within each gait than large bipeds. After normalizing for size (based on Froude number, after Alexander, 1977), remaining kinematic variation is largely due to interspecific differences in posture and relative limb segment lengths. For their size, smaller bipeds have greater step lengths, limb excursion angles and duty factors than large bipeds because of their more crouched posture and greater effective limb length. The most notable differences in limb kinematics between birds and humans occur at the walk-run transition and are maintained as running speed increases. Change of gait is smooth and difficult to discern in birds, but distinct in humans, involving abrupt decreases in step length and duty factor (time of contact) and a corresponding increase in limb swing time. These differences appear to reflect a spring-like run that is stiff in humans (favouring elastic energy recovery) but more compliant in birds (increasing time of ground contact). Differences between birds and humans in balance of the body's centre of mass not only affect femoral orientation and motion, but also affect pattern of limb excursion with speed.     

Stick insects walking on a wheel: Perturbations induced by obstruction of leg protraction

Summary Stick insects (Carausius morosus) walking on a wheel were perturbed by restricting the forward protraction of individual legs. A barrier placed before a single middle or rear leg prevented that leg from reaching its normal protraction endpoint but allowed it unimpeded retraction. Upon striking the barrier, the protracting leg attempted to get past it and thereby prolonged protraction. This prolongation increased with the extent to which the obstruction infringed upon the leg's normal step range. Barriers placed near the midpoint of this range elicited large perturbations: the blocked leg often continued its protraction throughout many step cycles of the other legs (Fig. 1 E, F). For the most part walking was irregular and smooth forward progression was disrupted. Nevertheless, the infrequent steps by the affected leg usually were coordinated with those of the adjacent ipsilateral legs.More rostral barrier positions elicited smaller perturbations: the blocked leg usually made one step in each step cycle of the other legs (Fig. 1 B, C, D, G). Measurements for these regular step sequences showed quantitatively that protraction duration increased in proportion to the severity of the infringement on normal leg movement (Figs. 3, 4). The fraction of the step period occupied by protraction increased from ca. 10% for normal walking to ca. 50% for caudal barrier positions. This proportionality is interpreted to show the importance of spatial components of the walking program.When one leg was obstructed, its extended protraction influenced the stepping of the three adjacent legs as follows. First, the ipsilateral rostral leg showed the largest change: its protraction onset was regularly delayed for the duration of the extended protraction (Figs. 4, 7, 8), demonstrating a strong, centrally mediated inhibition. The presence of a further delay of up to 100 to 140 ms suggests that peripheral input from the protracting leg may be important for releasing this inhibition. Second, steps by an adjacent caudal leg were not measurably affected. However, the method may not have sufficed to reveal such effects because during regular walking middle leg protractions rarely lasted long enough to conflict with subsequent steps by the ipsilateral rear leg. Third, contralateral effects differed between middle and rear leg obstructions. If the obstructed leg was a middle leg, its extended protraction had little effect upon stepping by the contralateral middle leg: the latter leg frequently protracted while the blocked leg continued its protraction and there was no consistent change in the phase relation of these two legs (Table 1). In contrast, if the obstructed leg was a rear leg, protractions by the contralateral rear leg tended to be delayed (Table 1).     

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.     

Motor output to the protractor and retractor coxae muscles in stick insects walking on a treadwheel

ABSTRACT. The motor output to the protractor and retractor mucles moving the coxa of the middle leg of Carausius morosus was recorded from the thoracic nerves during walking on a treadwheel. The leg movements on the wheel were generally similar to those found in free-walking animals, but tripod coordination was relatively independent of period, and the coordination of the adult animal on the wheel was most closely related to that found in free-walking first instars. The activity of a common inhibitor and four excitatory axons of the retractor and an excitatory axon of the protractor were followed for 850 steps (in six animals) to give a summary of the behaviour of the different units. The motor activity is less stereotyped than that previously reported for insects. There was strong reciprocity between the antagonists, but this was not directly correlated with the forward and backward movements of the legs. The first part of the stance phase of the leg was accompanied by a strong burst in the protractor nerve and relatively little retractor activity. This was followed by the main retractor burst which occupied the last 60% of the stance phase. The results are compared with motor output records of the locust and with earlier force-plate measurements on the stick insect. It must be concluded that the mesothoracic leg initially resists forward movement of the body by the other legs during a typical walking step.     

The influence of locomotor training on dynamic balance during steady-state walking post-stroke

Slow walking speed and lack of balance control are common impairments post-stroke. While locomotor training often improves walking speed, its influence on dynamic balance is unclear. The goal of this study was to assess the influence of a locomotor training program on dynamic balance in individuals post-stroke during steady-state walking and determine if improvements in walking speed are associated with improved balance control. Kinematic and kinetic data were collected pre- and post-training from seventeen participants who completed a 12-week locomotor training program. Dynamic balance was quantified biomechanically (peak-to-peak range of frontal plane whole-body angular-momentum) and clinically (Berg-Balance-Scale and Dynamic-Gait-Index). To understand the underlying biomechanical mechanisms associated with changes in angular-momentum, foot placement and ground-reaction-forces were quantified. As a group, biomechanical assessments of dynamic balance did not reveal any improvements after locomotor training. However, improved dynamic balance post-training, observed in a sub-group of 10 participants (i.e., Responders), was associated with a narrowed paretic foot placement and higher paretic leg vertical ground-reaction-force impulse during late stance. Dynamic balance was not improved post-training in the remaining seven participants (i.e., Non-responders), who did not alter their foot placement and had an increased reliance on their nonparetic leg during weight-bearing. As a group, increased walking speed was not correlated with improved dynamic balance. However, a higher pre-training walking speed was associated with higher gains in dynamic balance post-training. These findings highlight the importance of the paretic leg weight bearing and mediolateral foot placement in improving frontal plane dynamic balance post-stroke.     

IMU-based ambulatory walking speed estimation in constrained treadmill and overground walking

This study evaluated the performance of a walking speed estimation system based on using an inertial measurement unit (IMU), a combination of accelerometers and gyroscopes. The walking speed estimation algorithm segments the walking sequence into individual stride cycles (two steps) based on the inverted pendulum-like behaviour of the stance leg during walking and it integrates the angular velocity and linear accelerations of the shank to determine the displacement of each stride. The evaluation was performed in both treadmill and overground walking experiments with various constraints on walking speed, step length and step frequency to provide a relatively comprehensive assessment of the system. Promising results were obtained in providing accurate and consistent walking speed/step length estimation in different walking conditions. An overall percentage root mean squared error (%RMSE) of 4.2 and 4.0% was achieved in treadmill and overground walking experiments, respectively. With an increasing interest in understanding human walking biomechanics, the IMU-based ambulatory system could provide a useful walking speed/step length measurement/control tool for constrained walking studies.     

IMU-based ambulatory walking speed estimation in constrained treadmill and overground walking

This study evaluated the performance of a walking speed estimation system based on using an inertial measurement unit (IMU), a combination of accelerometers and gyroscopes. The walking speed estimation algorithm segments the walking sequence into individual stride cycles (two steps) based on the inverted pendulum-like behaviour of the stance leg during walking and it integrates the angular velocity and linear accelerations of the shank to determine the displacement of each stride. The evaluation was performed in both treadmill and overground walking experiments with various constraints on walking speed, step length and step frequency to provide a relatively comprehensive assessment of the system. Promising results were obtained in providing accurate and consistent walking speed/step length estimation in different walking conditions. An overall percentage root mean squared error (%RMSE) of 4.2 and 4.0% was achieved in treadmill and overground walking experiments, respectively. With an increasing interest in understanding human walking biomechanics, the IMU-based ambulatory system could provide a useful walking speed/step length measurement/control tool for constrained walking studies.     

Evaluation of gait and slip parameters for adults with intellectual disability

Adults with intellectual disability (ID) experience more falls than their non-disabled peers. A gait analysis was conducted to quantify normal walking, and an additional slip trial was performed to measure slip response characteristics for adults with ID as well as a group of age- and gender-matched controls. Variables relating to gait pattern, slip propensity, and slip severity were assessed to compare the differences between groups. The ID group was found to have significantly slower walking speed, shorter step lengths, and increased knee flexion angles at heel contact. These gait characteristics are known to reduce the likelihood of slip initiation in adults without ID. Despite a more cautious gait pattern, however, the ID group exhibited greater slip distances indicating greater slip severity. This study suggests that falls in this population may be due to deficient slip detection or insufficient recovery response.     

What are the relations between mechanics, gait parameters, and energetics in terrestrial locomotion?

Are the different energy-conserving mechanics (i.e., pendulum and spring) used in different gaits reflected in differences in energetics and/or stride parameters? The analysis included published data from several species and new data from horses. When changing from pendulum to spring mechanics, there is a change in the slope of metabolic rate (MR) vs. speed in all species, in birds and quadrupeds there is no step increase, and in humans there are conflicting reports. At the trot-gallop transition, where quadrupeds are hypothesized to change from spring mechanics to some combination of spring and pendulum mechanics, there is a change in slope of MR vs. speed in horses but not in other species. Stride frequency (SF) is a logarithmic function of walking speed in all species, a linear function of trotting/running speed, and nearly independent of speed in galloping. In humans and horses there is a discontinuity in SF at the walk-trot (run) transition but not in birds. The slope of time of contact vs. speed does not change with mechanics in most species, but it does in humans. In horses and humans, there is a discontinuity at the walk-trot (run) transition and data for other species do not permit generalization. Duty factor (DF) in humans is greater than 0.5 in walking (pendulum mechanics) and less than 0.5 when running (spring mechanics). However, this is not true in many species that have DF>0.5 at the lowest speeds where they use spring mechanics. When trotting at low speeds, horses use forelimb DF>0.5 and hind limb DF<0.5. Thus, it is confusing to distinguish between walking and running by DF.     

Voluntary bipedal walking of infant chimpanzees

The voluntary bipedal walking of infant chimpanzees was studied by the analysis of foot force and by motion analysis. The infants were trained to locomote on a level platform without any restrictions on the locomotor pattern. The voluntary bipedal walking was compared with the other types of locomotion at the same age and with the trained bipedal walking performed by other chimpanzees, including adult chimpanzees. The characteristics of voluntary bipedal walking in the infant until one year of age were: (1) high-speed walking with short cycle duration; (2) short stance phase duration; (3) small braking component of the preceding leg and large acceleration of the following leg; (4) one downward peak in the vertical component; and (5) a relatively small transverse component. Bipedal walking usually continued for less than one second and ended in quadrupedal locomotion. During walking, the preceding foot touched the floor, heel first, as in the case of older chimpanzees and humans. At this age, bipedal walking was similar to high-speed locomotion. The voluntary bipedal walking of the two-year-old and frour-yearold chimpanzees was characterized as follows: (1) slower speed than during quadrupedal locomotion, (2) relatively long periods and distances; (3) well balanced accelerating and braking components; and (4) a vertical component showing two downward peaks and a trough in between during numerous trials. The last characteristic means that the body center of gravity is higher in the single stance phase, just as in the bipedal walkinbg of the adult chimpanzees and humans. The bipedal walking of infant chimpanzees was discussed in comparison with the walking of humans, including infants.