Biomechanical mechanism for transitions in phase and frequency of arm and leg swing during walking

As humans increase walking speed, there are concurrent transitions in the frequency ratio between arm and leg movements from 2:1 to 1:1 and in the phase relationship between the movements of the two arms from in-phase to out-of-phase. Superharmonic resonance of a pendulum with monofrequency excitation had been proposed as a potential model for this phenomenon. In this study, an alternative model of paired pendulums with multiple-frequency excitations is explored. It was predicted that the occurrence of the concurrent transitions was a function of (1) changes in the magnitude ratio of shoulder accelerations at step and stride frequencies that accompany changes in walking speed and (2) proximity of these frequencies to the natural resonance frequencies of the arms modeled as a pair of passive pendulums. Model predictions were compared with data collected from 14 healthy young subjects who were instructed to walk on a treadmill. Walking speeds were manipulated between 0.18 and 1.52 m/s in steps of 0.22 m/s. Kinematic data for the arms and shoulders were collected using a 3D motion analysis system, and simulations were conducted in which the movements of a double-pendulum system excited by the accelerations at the suspension point were analyzed to determine the extent to which the arms acted as passive pendulums. It was confirmed that the acceleration waveforms at the shoulder are composed primarily of stride and step frequency components. Between the shoulders, the stride frequency components were out-of-phase, while the step frequency components were in-phase. The amplitude ratio of the acceleration waveform components at the step and stride frequencies changed as a function of walking speed and were associated with the occurrence of the transitions. Simulation results using these summed components as excitatory inputs to the double-pendulum system were in agreement with actual transitions in 80% of the cases. The potential role of state-dependent active muscle contraction at shoulder joints on the occurrence of the transitions was discussed. Due to the tendency of arm movements to stay in the vicinity of their primary resonance frequency, these active muscle forces were hypothesized to function as escapements that created limit cycle oscillations at the shoulders resonant frequency.     

Are interlimb interactions disturbed in patients with Parkinson’s disease? A study under unloading conditions

During natural human locomotion, neural connections are activated that are typical of regulation of the quadrupedal walking. The interaction between the neural networks generating rhythmic movements of the upper and lower limbs depends on tonic state of each of these networks regulated by motor signals from the brain. Distortion of these signals in patients with Parkinson’s disease (PD) may lead to disruption of the interlimb interactions. We examined the effect of movements of the limbs of one girdle on the parameters of the motor activity of another limb girdle at their joint cyclic movements under the conditions of arm and leg unloading in 17 patients with PD and 16 healthy subjects. We have shown that, in patients, the effect of voluntary and passive movements of arms, as well as the active movement of the distal parts of arms, on the voluntary movement of legs is weak, while in healthy subjects, the effect of arm movements on the parameters of voluntary stepping is significant. The effect of arm movements on the activation of the involuntary stepping by vibrational stimulation of-legs in patients was absent, while in healthy subjects, the motor activity of arms increased the possibility of involuntary rhythmic movements activation. Differences in the effect of leg movements on the rhythmic movements of arms were found in both patients and healthy subjects. The interlimb interaction appeared after drug administration. However, the effect of the drug was not sufficient for the recovery of normal state of the neural networks in patients. In PD patients, neural networks generating stepping rhythm have an increased tonic activity, which prevents the activation and appearance of involuntary rhythmic movements facilitating the effects of arms on legs.     

Interlimb interactions during cyclic in-phase and antiphase movements of arms and legs and their dependence on afferent influences

Coordinated arm and leg movements imply neural interactions between the rhythmic generators of the upper and lower extremities. In ten healthy subjects in the lying position, activity of the muscles of the upper and lower extremities was recorded during separate and joint cyclic movements of the arms and legs with different phase relationships between the movements of the limbs and under various conditions of the motor task. Antiphase active arm movements were characterized by higher muscle activity than during the inphase mode. The muscle activity during passive arm movements imposed by the experimentalist was significantly lower than muscle activity during passive arm movements imposed by the other arm. When loading one arm, the muscle activity in the other, passively moving, arm increased independently from the synergy of arm movements. During a motor task implementing joint antiphase movements of both upper and lower extremities, compared to a motor task implementing their joint in-phase movements, we observed a significant increase in activity in the biceps brahii muscle, the tibialis anterior muscle, and the biceps femoris muscle. Loading of arms in these motor tasks has been accompanied by increased activity in some leg muscles. An increase in the frequency of rhythmic movements resulted in a significant growth of the muscle activity of the arms and legs during their cooperative movements with a greater rate of rise in the flexor muscle activity of the arms and legs during joint antiphase movements. Thus, both the spatial organization of movements and the type of afferent influences are significant factors of interlimb interactions, which, in turn, determine the type of neural interconnections that are involved in movement regulation.     

Mutual influences of upper and lower extremities during cyclic movements

The possibility for the activation of muscles in a passive arm during its cyclic movements imposed by active movements of the contralateral arm or by an experimenter and the effect that the movements of lower extremities have on the activity of the arm muscles have been studied. In addition, the activity of the leg muscles was studied as dependent on the motor task performed by the arms. Ten healthy subjects performed antiphase arm movements with and without stepping-like movements of both legs in the supine position. The experiment was performed under three conditions for the arm movements: (1) both arms performed active movements; (2) one arm performed active movements, and the contralateral arm, being entirely passive, was forced to participate in movements; (3) the movement of the passive arm was caused by an experimenter. Under condition (2), additional loadings of 30 and 60 N were applied to the active arm. Under all conditions, the arm movements were performed with and without leg movements. The possibility for the activation of muscles in the arm performing passive movements has been demonstrated. To a large extent, this is possible due to an increase in the afferent inflow from the muscles of the contralateral arm. The electrical activity was modulated during cyclic arm movements and depended on the level of loading of the active arm. During the combined active movements of the arms and legs, the reduction in the activity of the flexor muscles of the shoulder and forearm was observed. In the case of passive stepping-like movements, the concomitant arm movements increased the magnitude of electromyographic bursts in most of the examined leg muscles. During active leg movements, a similar increase in electromyographic bursts was observed only in the m. biceps femoris (BF) and the anterior tibial muscle. An increase in the loading of one arm caused a significant increase in the EMG activity in most examined muscles of the legs. The data obtained provide additional proof for the existence of a functionally significant neuronal interaction between the arms, as well as between the upper and lower extremities, which is probably due to intraspinal neuronal connections.     

Abnormalities in mutual influences of upper and lower limbs in patients with stroke

Previously, in healthy subjects the common pattern of muscle activation and specifics of interlimb neuron connections during performance of rhythmic separate and simultaneous movements of arms and legs in the lying position, which reflect functional meaningful of interlimb interactions, were shown. The aim of this research was to investigate such mutual influences of upper and lower limbs during the execution of similar motor tasks by patients with stroke. In sixteen poststroke patients with different stage of hemiparesis arms movements together with or without legs movements were performed, while lying supine. It was demonstrated that the common pattern of muscle activity distribution under the execution of voluntary cyclic movements by both arms was disordered. Passive rhythmic movements of each arm caused the phased EMG activity in shoulder muscles in patients with mild hemiparesis, but no activation was observed in patients with severe paresis. The loading of nonparetic arm resulted in an increasing of activity in shoulder flexor muscles of paretic arm in patients with weak paresis (which was typical for healthy subjects), while it not exerted essential influences in patients with severe paresis. Under connecting the cyclic movements of arms with stepping movements of legs in diagonal synergy the activity in proximal muscles of both arms was decreased irrespective of the paresis degree, as it was seeing in healthy subjects. Simultaneous arms and legs movements did not change the muscle activity in non-paretic leg in both groups of patients, but in some muscles of paretic leg the activity even decreased. The results obtained revealed important features of poststroke motor disturbances, which caused the changes of interlimb interaction and in great degree depended on the level of paresis. The data of investigation can be of a great importance for developing the new methods for rehabilitative procedure in patients with stroke.     

Gender differences in active musculoskeletal stiffness. Part II. Quantification of leg stiffness during functional hopping tasks.

Leg stiffness was compared between age-matched males and females during hopping at preferred and controlled frequencies. Stiffness was defined as the linear regression slope between the vertical center of mass (COM) displacement and ground-reaction forces recorded from a force plate during the stance phase of the hopping task. Results demonstrate that subjects modulated the vertical displacement of the COM during ground contact in relation to the square of hopping frequency. This supports the accuracy of the spring-mass oscillator as a representative model of hopping. It also maintained peak vertical ground-reaction load at approximately three times body weight. Leg stiffness values in males (33.9+/-8.7 kN/m) were significantly (p<0.01) greater than in females (26.3+/-6.5 kN/m) at each of three hopping frequencies, 3.0, 2.5 Hz, and a preferred hopping rate. In the spring-mass oscillator model leg stiffness and body mass are related to the frequency of motion. Thus male subjects necessarily recruited greater leg stiffness to drive their heavier body mass at the same frequency as the lighter female subjects during the controlled frequency trials. However, in the preferred hopping condition the stiffness was not constrained by the task because frequency was self-selected. Nonetheless, both male and female subjects hopped at statistically similar preferred frequencies (2.34+/-0.22 Hz), therefore, the females continued to demonstrate less leg stiffness. Recognizing the active muscle stiffness contributes to biomechanical stability as well as leg stiffness, these results may provide insight into the gender bias in risk of musculoskeletal knee injury.     

Abnormalities of mutual influences of the upper and lower limbs after a stroke

The common pattern of muscle activation and specifics of interlimb neuronal connections during the performance of rhythmic separate and simultaneous arm and leg movements in the lying position in healthy subjects, which reflected functionally significant interlimb neuronal interactions, were shown. The study was designed to investigate these mutual influences of the upper and lower limbs during the performance of similar motor tasks by stroke patients. Sixteen poststroke patients with different degrees of hemiparesis performed active and passive arm movements simultaneously with stepping leg movements or without them while lying supine. It was demonstrated that the patients had a disordered common pattern of distribution of muscle activity when they performed voluntary cyclic movements with both arms. Passive movements of both paretic and nonparetic arms led to different degrees of activation of their muscles, depending on the degree of paresis: in patients with mild paresis, muscle activation was similar to that in healthy subjects; in patients with severe paresis, it was insignificant. The loading of the nonparetic arm resulted in an increase in the activity in the paretic arm shoulder flexor muscles in patients with mild paresis (which was typical of healthy subjects), while loading did not influence significantly patients with severe paresis. The combination of cyclic arm movements and stepping leg movements in diagonal synergy decreased the activity in the proximal muscles of both arms, irrespective of the degree of paresis, as it was observed in healthy subjects. Simultaneous arm and leg movements did not change the muscle activity in nonparetic legs in either groups of patients, but the activity in the paretic leg muscles even decreased. The results obtained revealed important features of poststroke motor disturbances, which caused changes in interlimb interactions and largely depended on the degree of paresis. The data could be useful for developing new methods for the performance of rehabilitative procedures in poststroke patients.     

Activation of interlimb interactions increases the motor output in legs of healthy subjects: Study under the conditions of arm and leg unloading

The effect of arm movements and movements of individual arm joints on the electrophysiological and kinematic characteristics of voluntary and vibration-triggered stepping-like leg movements was studied under the conditions of horizontal support of the upper and lower limbs. The horizontal support of arms provided a significant increase in the rate of activation of locomotor automatism by noninvasive impact on tonic sensory inputs. The addition of active arm movements during involuntary stepping-like leg movements led to an increase in the EMG activity of hip muscles and was accompanied by an increase in the amplitude of hip and shin movements. The movement of the shoulder joints led to an increase in the activity of hip muscles and was accompanied by an increase in the amplitude of hip and shin movements. Passive arm movements had the same effect on induced leg movements. The movement of the shoulder joints led to an increase in the activity of hip muscles and an increase in the amplitude of movements of knee and hip joints. At the same time, the movement of forearms and wrists had a similar facilitating effect on the physiological and kinematic characteristics of rhythmic stepping-like movements, but influenced the distal segments of legs to a greater extent. Under the conditions of subthreshold vibration of leg muscles, voluntary arm movements led to activation of involuntary rhythmic stepping movements. During voluntary leg movements, the addition of arm movements had a significantly smaller impact on the parameters of rhythmic stepping than during involuntary leg movements. Thus, the simultaneous movements of the upper and lower limbs are an effective method of activation of neural networks connecting the rhythm generators of arms and legs. Under the conditions of arm and leg unloading, the interactions between the cervical and lumbosacral segments of the spinal cord seem to play the major role in the impact of arm movements on the patterns of leg movements. The described methods of activation of interlimb interactions can be used in the rehabilitation of post-stroke patients and patients with spinal cord injuries, Parkinson’s disease, and other neurological diseases.     

Organization of a complex movement: fixed and variable components of the cockroach escape behavior

The escape behavior of the cockroach Periplaneta americana was studied by means of high speed filming (250 frames/s) and a computer-graphical analysis of the body and leg movements. The results are as follows: 1. The behavior begins with pure rotation of the body about the posteriorly located cerci, followed by rotation plus forward translation, and finally pure translation (Figs. 1, 2). 2. A consistent inter-leg coordination is used for the entire duration of the turn (Fig. 3A). At the start of the movement, five or all six legs execute their first stance phase (i.e. leg on the ground during locomotion) simultaneously. By the end of the turn the pattern has changed to the alternate 'tripod' coordination characteristic of insect walking. The change-over from all legs working together, to working alternately, occurs by means of a consistent pattern of delays in the stepping of certain legs. 3. The movements made by each leg during its initial stance phase are carried out using consistent movement components in the anterior-posterior (A-P) and the medial-lateral (M-L) axes (Fig. 4A). The movement at a particular joint in each middle leg is found to be diagnostic for the direction of turn. 4. The size and direction of a given leg's M-L movement in its initial stance phase depends on the same leg's prior A-P position (Fig. 5). No such feedback effects were seen among different legs. 5. Animals that are fixed to a slick surface on which they make slipping leg movements show the same inter-leg coordination (Fig. 3B), direction of initial stance movement (Fig. 4B) and dependence of the leg's initial M-L movement on its prior A-P position (Fig. 6), as did free-ranging animals. 6. Cockroaches that are walking at the moment they begin their escape reverse those ongoing leg movements that are contrary to escape movements. 7. These results are discussed in terms of the overall coordination of the complex movements, and in terms of the known properties of the neural circuitry for escape. Possibilities for neurobiological follow-up of certain of the findings presented here are also addressed.     

Mutual influences of upper and lower extrimities during cyclic movements

The possibility of muscle activation of passive arm during its cyclic movements, imposed by active movements of contralateral arm or by experimenter was studied, as well as the influence of lower extremities cyclic movements onto arm muscles activity. In addition to that the activity of legs muscles was estimated in dependence on motor task condition for arms. Ten healthy supine subjects carried out opposite movements of arms with and without stepping-like movements of both legs. The experiment included three conditions for arm movements: 1) the active movements of both arms; 2) the active movements of one arm, when other entirely passive arm participated in the movement by force; 3) passive arm movement caused by experimenter. In the condition 2) additional load on active arm was applied (30 N and 60 N). In all three conditions the experiment was carried out with arms movements only or together with legs movements. The capability of passive moving arm muscles activation depended on increasing afferent inflow from muscles of contralateral arm was demonstrated. Emerging electrical activity was modulated in the arms movements cycle and depended on the degree of active arm loading. During combined active movements of arms and legs the reduction of activity in the flexor muscles of shoulder and forearm was observed. Concomitant arms movements increased the magnitude ofelectromiographic bursts during passive stepping-like movements in the most of recorded muscles, and the same increasing was only observed in biceps femoris and tibialis anterior muscles during active legs movement. The increasing of loading of one arm caused essential augmentation of EMG-activity in the majority of recording legs muscles. The data obtained are the additional proof of existence of functionally significant neuronal interaction both between arms and between upper and lower extremities, which is evidently depend on the intraspinal neuronal connections.     

Rotational locomotion by the cockroach Blattella germanica

Locomotion on complex substrata can be expressed in a plane by two geometric components of body movement: linear locomotion and rotational locomotion. This study examined pure rotation by analysing the geometry of leg movements and stepping patterns during the courtship turns of male Blattella germanica. Strict rotation or translation by an insect requires that each side of the body cover equal distance with respect to the substrate. There are three mechanisms by which the legs can maintain this equality: frequency of stepping, magnitude of the leg arcs relative to the body and the degree to which legs flex and extend during locomotion. During the courtship behaviour of Blattella germanica selected males executed turns involving body rotation along with leg movements in which the legs on the outside of the turn swung through greater average arcs than those on the inside of the turn. This difference should have resulted in a translation component. However, legs on the inside of the turn compensated by flexion and extension movements which were greater than those of opposing legs. The net effect was that both sides of the body covered equal average ground. These cockroaches used a wide variety of stepping combinations to effect rotation. The frequency of these combinations was compared to an expected frequency distribution of stepping combinations and further to an expected frequency of these stepping combinations used for straight walking. These comparisons demonstrated a similarity between interleg coordination during straight walking and that during turning in place.     

Frequency tuning in animal locomotion

In locomotion that involves repetitive motion of propulsive structures (arms, legs, fins, wings) there are resonant frequencies f(*) at which the energy consumption is a minimum. As animals need to change their speed, they can maintain this energy minimum by tuning their body resonances. We discuss the physical principles of frequency tuning, and how it relates to forces, damping, and oscillation amplitude. The resonant frequency of pendulum-type oscillators (e.g. swinging arms and legs) may be changed by varying the mass moment of inertia, or the vertical acceleration of the pendulum pivot. The frequency of elastic vibrations (e.g. the bell of a jellyfish) can be tuned with a non-linear modulus of elasticity: soft for low deflection amplitudes (low resonant frequency), and stiff for large displacements (high resonant frequency). Tuning of elastic oscillations can also be achieved by changing the effective length or cross-sectional area of the elastic members, or by allowing springs in parallel or in series to become active. We propose that swimming and flying animals generate oscillating propulsive forces from precisely placed shed vortices and that these tuned motions can only occur when vortex shedding and the simple harmonic motion of the elastic elements of the propulsive structures are in resonance.     

Hemiplegic gait: a kinematic analysis using walking speed as a basis.

The kinematics of treadmill ambulation of stroke patients (N = 9) and healthy subjects (N = 4) was studied at a wide range of different velocities (i.e. 0.25-1.5 m s-1), with a focus on the transverse rotations of the trunk. Video recordings revealed, for both stroke patients and healthy subjects, similar relations between walking speed and stride length as well as stride frequency. The phase difference between pelvic and thoracic rotations (i.e. trunk rotation) and the total range of trunk rotation were almost linearly related to the walking speed. Healthy subjects showed a marked increase in pelvic rotation from 1 to 1.5 m s-1. Using dimensional analysis in a comparison between stroke patients and healthy subjects, invariances in the coordination of gait were found for stride length, stride frequency, pelvic rotation, and trunk rotation. Constant relations were obtained between, on the one hand, dimensionless velocity and, on the other, dimensionless stride length as well as stride frequency. Transitions were found between the velocities 0.75 and 1 m s-1 for dimensionless pelvic rotation and trunk rotation, indicating that, from this velocity range onwards, pelvic swing lengthens the stride: rotations of pelvis, thorax and trunk become tightly coordinated. On the basis of the dimensionless stride length, stride frequency, pelvic rotation and trunk rotation, deficits in the gait of stroke patients could be quantified. It is concluded that walking speed is an important control parameter, which should be used as a basic variable in the evaluation of the gait of stroke patients.     

Mechanical stimulation of the support zones of soles: The method of noninvasive activation of the stepping movement generators in humans

The effects of mechanical stimulation of the soles’ support zones in the modes of slow and fast walking (75 and 120 steps per minute) were studied using the model of supportlessness (legs suspension). 20 healthy subjects participated in the study. EMG activity of hip and shin muscles was recorded. Kinematics of leg movements was assessed with the use of videoanalysis system. In 80% of cases support stimulation was followed by leg movements, in 69% of which they had characteristics of locomotions being accompanied by the burst-like electromyographic activities. The order of involvement of leg muscles and organization of antagonistic muscles activities were analogous to those of voluntary walking. The latencies of electromyographic activity in hip and shin muscles composed 5.17 ± 1.08 and 14.01 ± 2.82 s, respectively, the frequencies of bursts differed significantly depending on stimulation frequency. In 31% of cases the electromyographical activity following the stimulation of the soles’ support zones had not burst-like but uninterrupted pattern. Its amplitude rose smoothly reaching a certain level that was subsequently maintained. Results of the study showed that soles’ support zones stimulation in the mode of locomotion could activate a locomotor generator provoking the appearance of locomotion-like activity and that effect evoked by this stimulation includes not only rhythmical but also non-rhythmical (probably postural) components of walking.     

Bionic ankle-foot prosthesis normalizes walking gait for persons with leg amputation

Over time, leg prostheses have improved in design, but have been incapable of actively adapting to different walking velocities in a manner comparable to a biological limb. People with a leg amputation using such commercially available passive-elastic prostheses require significantly more metabolic energy to walk at the same velocities, prefer to walk slower and have abnormal biomechanics compared with non-amputees. A bionic prosthesis has been developed that emulates the function of a biological ankle during level-ground walking, specifically providing the net positive work required for a range of walking velocities. We compared metabolic energy costs, preferred velocities and biomechanical patterns of seven people with a unilateral transtibial amputation using the bionic prosthesis and using their own passive-elastic prosthesis to those of seven non-amputees during level-ground walking. Compared with using a passive-elastic prosthesis, using the bionic prosthesis decreased metabolic cost by 8 per cent, increased trailing prosthetic leg mechanical work by 57 per cent and decreased the leading biological leg mechanical work by 10 per cent, on average, across walking velocities of 0.75-1.75 m s(-1) and increased preferred walking velocity by 23 per cent. Using the bionic prosthesis resulted in metabolic energy costs, preferred walking velocities and biomechanical patterns that were not significantly different from people without an amputation.     

Kinematics and motor activity during tethered walking and turning in the cockroach, <Emphasis Type="Italic">Blaberus discoidalis</Emphasis>

When insects turn from walking straight, their legs have to follow different motor patterns. In order to examine such pattern change precisely, we stimulated single antenna of an insect, thereby initiating its turning behavior, tethered over a lightly oiled glass plate. The resulting behavior included asymmetrical movements of prothoracic and mesothoracic legs. The mesothoracic leg on the inside of the turn (in the apparent direction of turning) extended the coxa-trochanter and femur-tibia joints during swing rather than during stance as in walking, while the outside mesothoracic leg kept a slow walking pattern. Electromyograms in mesothoracic legs revealed consistent changes in the motor neuron activity controlling extension of the coxa-trochanter and femur-tibia joints. In tethered walking, depressor trochanter activity consistently preceded slow extensor tibia activity. This pattern was reversed in the inside mesothoracic leg during turning. Also for turning, extensor and depressor motor neurons of the inside legs were activated in swing phase instead of stance. Turning was also examined in free ranging animals. Although more variable, some trials resembled the pattern generated by tethered animals. The distinct inter-joint and inter-leg coordination between tethered turning and walking, therefore, provides a good model to further study the neural control of changing locomotion patterns.     

Mechanisms of stick insect locomotion in a gap-crossing paradigm

Locomotion of stick insects climbing over gaps of more than twice their step length has proved to be a useful paradigm to investigate how locomotor behaviour is adapted to external conditions. In this study, swing amplitudes and extreme positions of single steps from gap-crossing sequences have been analysed and compared to corresponding parameters of undisturbed walking. We show that adaptations of the basic mechanisms concern movements of single legs as well as the coordination between the legs. Slowing down of stance velocity, searching movements of legs in protraction and the generation of short steps are crucial prerequisites in the gap-crossing task. The rules of leg coordination described for stick insect walking seem to be modified, and load on the supporting legs is assumed to have a major effect on coordination especially in slow walking. Stepping into the gap with a front leg and antennal contact with the far edge of the gap provide information, as both events influence the following leg movements, whereas antennal non-contact seems not to contain information. Integration of these results into the model of the walking controller can improve our understanding of insect locomotion in highly irregular environments.Abbreviations AEP anterior extreme position - fAEP fictive anterior extreme position - PEP posterior extreme position - TOT treading-on-tarsus     

Phase-Dependent Effects of Transcutaneous Spinal Cord Stimulation on Regulation of Kinematics of Human Stepping Motions

The effect of transcutaneous electrical spinal cord stimulation on the kinematic parameters of movement of the ipsilateral and contralateral legs in healthy subjects during treadmill walking at speeds of 1.5 to 1.7 km/h has been studied. The stimulation electrodes were placed 2.5 cm lateral from the right and left sides of the spinal midline at L1 and T11 levels. During the stance phase, stimulation was administered at L1 level at a frequency of 15 Hz; during the swing phase the stimuli was delivered to T11 at a frequency of 30 Hz, followed by alternating stimulation at L1 and T11. The stimulation during the swing phase (T11) was more effective than that during the stance phase (L1); the most impressive changes in kinematic parameters were observed when combined delivery of stimulations to L1 and T11 was performed. With unilateral spinal stimulation, the amplitude of the angles in the hip, knee and/or ankle joints, the length of the transfer, and the height of the leg elevation increased in the ipsilateral leg. Similar but less pronounced changes were observed in the contralateral leg. A 10% increase in the duration of stimulation in the swing phase caused a change in the kinematic stepping parameters both in ipsilateral and contralateral legs. The maximum effect was observed when bilateral alternating stimulation was used. These data show that phasic transcutaneous electrical spinal cord stimulation, using a wide range of natural walking speeds, can be applied to control kinematic movement parameters.

     

Righting kinematics in beetles (Insecta: Coleoptera)

Twenty modes of stereotyped righting motions were observed in 116 representative species of coleoptera. Methods included cine and stereocine recording with further frame by frame analysis, stereogrammetry, inverse kinematic reconstruction of joint angles, stroboscopic photography, recording of electromyograms, 3D measurements of the articulations, etc. The basic mode consists of a search phase, ending up with grasping the substrate, and a righting, overturning phase. Leg coordination within the search cycle differs from the walking cycle with respect to phasing of certain muscle groups. Search movements of all legs appear chaotic, but the tendency to move in antiphase is still present in adjacent ipsilateral and contralateral leg pairs. The system of leg coordination might be split: legs of one side might search, while contralateral legs walk, or fore and middle legs walk while hind legs search. Elaborated types of righting include somersaults with the aid of contralateral or diagonal legs, pitch on elytra, jumps with previous energy storage with the aid of unbending between thoracic segments (well-known for Elateridae), or quick folding of elytra (originally described in Histeridae). Righting in beetles is compared with righting modes known in locusts and cockroaches. Search in a righting beetle is directed dorsad, while a walking insect searches for the ground downwards. Main righting modes were schematized for possible application to robotics.     

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.