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  • 1
    Electronic Resource
    Electronic Resource
    Springer
    Experimental brain research 90 (1992), S. 404-414 
    ISSN: 1432-1106
    Keywords: Human gait ; Sural nerve ; Ankle angle ; Reflex reversal ; Phase-dependent modulation ; Human
    Source: Springer Online Journal Archives 1860-2000
    Topics: Medicine
    Notes: Summary To investigate whether phase-dependent reversals in reflex responses on electromyography (EMG) are accompanied by movement reversals, a series of human volunteers were studied for their behavioural responses to sural nerve stimulation during running or walking on a treadmill. Low-intensity stimulation (〈 2.5 x perception threshold, T) of the sural nerve yielded facilitatory responses in the tibialis anterior muscle (TA), correlated with an induced ankle dorsiflexion (mean maximum 4°) in early swing. The same stimuli yielded primarily TA suppression and weak ankle plantar flexion (mean maximum 1°) at end swing. The correlated induced knee angle changes did not precede the ankle changes, and they were relatively small. Mean maximum flexion in early swing was 6.2°, while mean maximum extension was 3.7°. High-intensity stimulation of the sural nerve (〉 2.5 x T) always gave rise to suppression of the ongoing activity. This resulted in a second type of movement reversal. During late stance and early swing the responses in TA were suppressive (i.e. below background activity) and related to ankle plantar flexion. In contrast, the responses during early and middle stance consisted of suppression in extensor activity (gastrocnemius medialis and soleus) and ankle dorsiflexion. The data are discussed in terms of a new hypothesis, which states that the responses to electrical stimulation of cutaneous nerves during locomotion do not correspond directly to corrections for stumbling following mechanical perturbations during the step cycle. Instead, the data invite a reinterpretation in terms of the opening and closing of reflex pathways, presumably by a central pattern generator for locomotion.
    Type of Medium: Electronic Resource
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  • 2
    ISSN: 1432-1106
    Keywords: Key words Human gait ; Transcranial magnetic stimulation ; Motor cortex ; Leg flexor/extensor muscle ; Corticospinal input ; Visual control
    Source: Springer Online Journal Archives 1860-2000
    Topics: Medicine
    Notes: Abstract  The aim of this study was to investigate visuomotor control during human gait. It was assumed that visual input should modulate transcranially evoked motor potentials (EMPs) during walking. The effect of transcranial magnetic stimulation (TMS) in a visually guided precision stepping task was compared with that during normal gait. EMPs were studied in tibialis anterior (TA), gastrocnemius (GM), and abductor digiti minimi (AD) muscles during treadmill walking. In both stepping tasks, a facilitation of EMPs was observed prior to activation of the respective leg muscle. EMP facilitation proved to be modulated throughout the stride cycle when normalising EMP with respect to the underlying electromyogram (EMG). Facilitation was strongest in TA prior to the swing phase. Significant differences of EMP facilitation between the visual and control tasks were present. In the visual task, maximal facilitation of TA EMPs prior to and during the swing phase was decreased compared to the control task. Conversely, there was increased facilitation of GM EMPs during swing phase of the visual task, prior to the heel strike and prior to the plantarflexion, which was the moment when the target was hit. Thus, the effect of visual input upon EMPs in TA and GM was differential and reciprocal according to the respective functional state. The results support the hypothesis of a conditioning effect of visual or, alternatively volitional, drive on EMPs during stepping.
    Type of Medium: Electronic Resource
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  • 3
    ISSN: 1432-1106
    Keywords: Split-belt locomotion ; Interlimb coordination ; Adaptation ; Motor learning ; Human
    Source: Springer Online Journal Archives 1860-2000
    Topics: Medicine
    Notes: Abstract Split-belt locomotion (i.e., walking with unequal leg speeds) requires a rapid adaptation of biome-chanical parameters and therefore of leg muscle electromyographic (EMG) activity. This adaptational process during the first strides of asymmetric gait as well as learning effects induced by repetition were studied in 11 healthy volunteers. Subjects were switched from slow (0.5 m/s) symmetric gait to split-belt locomotion with speeds of 0.5 m/s and 1.5 m/s, respectively. All subjects were observed to adapt in a similar way: (1) during the first trial, adaptation required about 12–15 strides. This was achieved by an increase in stride cycle duration, i.e., an increase in swing duration on the fast side and an increase in support duration on the slow side. (2) Adaptation of leg extensor and flexor EMG activity paralleled the changes of biomechanical parameters. During the first strides, muscle activity was enhanced with no increase in coactivity of antagonistic leg muscles. (3) A motor learning effect was seen when the same paradigm was repeated a few minutes later — interrupted by symmetric locomotion — as adaptation to the split-belt speeds was achieved within 1–3 strides. (4) This short-time learning effect did not occur in the “mirror” condition when the slow and fast sides were inverted. In this case adaptation again required 12–15 strides. A close link between central and proprioceptive mechanisms of interlimb coordination is suggested to underlie the adaptational processes during split-belt conditions. It can be assumed that, as in quadrupedal locomotion of the cat, human bipedal locomotion involves separate locomotor generators to provide the flexibility demanded. The present results suggest that side-specific proprioceptive information regarding the dynamics of the movement is necessary to adjust the centrally generated locomotor activity for both legs to the actual needs for controlled locomotion. Although the required pattern is quickly learned, this learning effect cannot be transferred to the contralateral side.
    Type of Medium: Electronic Resource
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  • 4
    Electronic Resource
    Electronic Resource
    Springer
    Experimental brain research 40 (1980), S. 89-95 
    ISSN: 1432-1106
    Keywords: Human balancing ; Spinal stretch reflex ; Leg muscle EMG ; Proprioceptive posture control
    Source: Springer Online Journal Archives 1860-2000
    Topics: Medicine
    Notes: Summary While subjects balanced on a seesaw consisting of a platform with a curved base, the antero-posterior sway of head and body as well as changes in the angle of the ankle joint were recorded and analysed for their frequency power spectrum. The EMG of leg muscles and the position of the resultant force exerted by the seesaw on a force-measuring platform were simultaneously registered and analysed. Balancing oscillations of 4–5 Hz were observed under this condition. They were accompanied by short, reciprocally organized bursts of EMG activity in the leg muscles. When stimulating the tibialis nerves to produce a displacement, the delay until the counterbalancing EMG activity started (about 40 ms) was in the time range of a fast-conducting segmental reflex. After partial ischaemic blocking of group I afferents from the leg muscles or fixation of the ankle joints, the predominant sway frequency was lacking, bursts of EMG activity became longer and stronger, and body balance was more unstable. Altering the height of the seesaw showed that a threshold change in the ankle angle was the determining factor in the production of spinal stretch reflex activity for fast regulation of balance.
    Type of Medium: Electronic Resource
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  • 5
    Electronic Resource
    Electronic Resource
    Springer
    Experimental brain research 109 (1996), S. 441-449 
    ISSN: 1432-1106
    Keywords: Presynaptic inhibition ; Ia fibres ; Spinal reflexes ; Gait ; Human
    Source: Springer Online Journal Archives 1860-2000
    Topics: Medicine
    Notes: Abstract Modulation of presynaptic inhibition of Ia afferents projecting monosynaptically to soleus motoneurones was investigated during human gait. Changes in presynaptic inhibition of Ia afferents were deduced from alterations in the amount of heteronymous soleus H-reflex facilitation evoked by a constant femoral nerve stimulation. It has been shown that this facilitation is mediated through a monosynaptic Ia pathway and that during its first 0.5 ms it is still uncontaminated by any polysynaptic effect and can be used to assess ongoing presynaptic inhibition of Ia terminals to soleus motoneurones. During gait, heteronymous facilitation was reduced with respect to its control value (rest during sitting) and modulated during the step cycle: it reached its maximum at mid-stance and decreased to near zero by the end of stance. At the same time the H-reflex amplitude was to some extent similarly modulated. It is argued that this decrease in heteronymous Ia facilitation and in H-reflex amplitude reflects an increased, ongoing presynaptic inhibition of Ia terminals projecting onto soleus motoneurones, which could be from central and/or peripheral origin. D1 inhibition, i.e. the late and long-lasting inhibition of the soleus H-reflex evoked by a train of stimuli to the common peroneal nerve, was used as another method to assess presynaptic inhibition. This D1 inhibition was decreased during gait, and it is argued that this decrease might reflect an occlusion in presynaptic pathways or increased presynaptic inhibition of pathways mediating the conditioning volley.
    Type of Medium: Electronic Resource
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  • 6
    Electronic Resource
    Electronic Resource
    Springer
    Experimental brain research 115 (1997), S. 234-246 
    ISSN: 1432-1106
    Keywords: Key words Human gait ; Transcranial magnetic stimulation ; Motor cortex ; Leg flexor/extensor muscle ; Corticospinal input
    Source: Springer Online Journal Archives 1860-2000
    Topics: Medicine
    Notes: Abstract  Transcranial magnetic stimulation (TMS) of the motor cortex was applied during locomotion to investigate the significance of corticospinal input upon the gait pattern. Evoked motor responses (EMR) were studied in the electromyogram (EMG) of tibialis anterior (TA), gastrocnemius (GM) and, for reference, abductor digiti minimi (AD) muscles by applying below-threshold magnetic stimuli during treadmill walking in healthy adults. Averages of 15 stimuli introduced randomly at each of 16 phases of the stride cycle were analysed. Phase-dependent amplitude modulation of EMR was present in TA and GM which did not always parallel the gait-associated modulation of the EMG activity. No variation of onset latency of the EMR was observed. The net modulatory response was calculated by comparing EMR amplitudes during gait with EMR amplitudes obtained (at corresponding background EMG activities) during tonic voluntary muscle contraction. Large net responses in both muscles occurred prior to or during phasic changes of EMG activity in the locomotor pattern. This facilitation of EMR was significantly higher in leg flexor than extensor muscles, with maxima in TA prior to and during late swing phase. A comparison of this facilitation of TA EMR prior to swing phase and prior to a phasic voluntary foot dorsiflexion revealed a similar onset but an increased amount of early facilitation in the gait condition. The modulated facilitation of EMR during locomotion could in part be explained by spinal effects which are different under dynamic and static motor conditions. However, we suggest that changes in corticospinal excitability during gait are also reflected in this facilitation. This suggestion is based on: (1) the similar onset yet dissimilar size of facilitatory effects in TA EMR prior to the swing phase of the stride cycle and during a voluntary dynamic activation, (2) the inverse variation of EMR and EMG amplitudes during this phase, and (3) the occurrence of this inversion at stimulation strengths below motor threshold (motor threshold was determined during weak tonic contraction and EMR were facilitated during gait). It is hypothesized that the facilitation is phase linked to ensure postural stability and is most effective during the phases prior to and during rhythmical activation of the leg muscles resulting in anticipatory adjustment of the locomotor pattern.
    Type of Medium: Electronic Resource
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  • 7
    Electronic Resource
    Electronic Resource
    Springer
    Experimental brain research 38 (1980), S. 117-119 
    ISSN: 1432-1106
    Keywords: Vestibular postural control ; Ischemic blocking ; Spindle afferents ; Tabes dorsalis ; Man
    Source: Springer Online Journal Archives 1860-2000
    Topics: Medicine
    Notes: Summary After minimizing proprioceptive input from the legs by ischemia without degradation of muscle force and excluding visual stabilization by eye closure, a characteristic anterior-posterior postural sway around 1 Hz was observed in three normal subjects. This is similar to the instability seen in two tabes dorsalis patients. From the spectral analysis of head and hip movements, displacements of the center of force and of ankle angle as well as from EMG recordings of the anterior tibial and gastrocnemius muscle it is concluded that the oscillations around 1 Hz are due to the long latency and high threshold of vestibularly induced leg muscle discharges (200–300 ms) arriving in the counterbalancing phase of the trunk, which causes an overshoot in body sway.
    Type of Medium: Electronic Resource
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  • 8
    Electronic Resource
    Electronic Resource
    Springer
    Experimental brain research 47 (1982), S. 172-176 
    ISSN: 1432-1106
    Keywords: Spinal EMG coordination ; EMG in balancing ; Leg muscle activity
    Source: Springer Online Journal Archives 1860-2000
    Topics: Medicine
    Notes: Summary While subjects were standing and balancing on two separate seesaws, the EMG of the leg muscles and the positions of the two seesaws were recorded. The spontaneous balancing movements with predominant oscillations of 4–5 Hz, and the accompanying bursts of EMG activity in the leg muscles occurred quite symmetrically on the two sides. After a displacement, induced either by stimulating the tibial nerves, or by a brisk anterior tilt of one seesaw, the EMG responses of the tibialis anterior muscles started with the same latency (about 50 ms) on both sides, and with similar amplitudes, even when only one side was displaced. It is concluded that this symmetrical leg muscle activation is mediated by a spinal coordinating mechanism the function of which depends on the actual motor task.
    Type of Medium: Electronic Resource
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  • 9
    Electronic Resource
    Electronic Resource
    Springer
    Experimental brain research 100 (1994), S. 93-106 
    ISSN: 1432-1106
    Keywords: Vestibular ; Posture ; Head stabilization ; Somatosensory ; Human
    Source: Springer Online Journal Archives 1860-2000
    Topics: Medicine
    Notes: Abstract The relative contribution of vestibular and somatosensory information to triggering postural responses to external body displacements may depend on the task and on the availability of sensory information in each system. To separate the contribution of vestibular and neck mechanisms to the stabilization of upright stance from that of lower body somatosensory mechanisms, responses to displacements of the head alone were compared with responses to displacements of the head and body, in both healthy subjects and in patients with profound bilateral vestibular loss. Head displacements were induced by translating two 1-kg weights suspended on either side of the head at the level of the mastoid bone, and body displacements were induced translating the support surface. Head displacements resulted in maximum forward and backward head accelerations similar to those resulting from body displacements, but were not accompanied by significant center of body mass, ankle, knee, or hip motions. We tested the effect of disrupting somatosensory information from the legs on postural responses to head or body displacements by sway-referencing the support surface. The subjects' eyes were closed during all testing to eliminate the effects of vision. Results showed that head displacements alone can trigger medium latency (48–84 ms) responses in the same leg and trunk muscles as body displacements. Nevertheless, it is unlikely that vestibular signals alone normally trigger directionally specific postural responses to support surface translations in standing humans because: (1) initial head accelerations resulting from body and head displacements were in opposite directions, but were associated with activation of the same leg and trunk postural muscles; (2) muscle responses to displacements of the head alone were only one third of the amplitude of responses to body displacements with equivalent maximum head accelerations; and (3) patients with profound bilateral vestibular loss showed patterns and latencies of leg and trunk muscle responses to body displacements similar to those of healthy subjects. Altering somatosensory information, by sway-referencing the support surface, increased the amplitude of ankle muscle activation to head displacements and reduced the amplitude of ankle muscle activation to body displacements, suggesting context-specific reweighting of vestibular and somatosensory inputs for posture. In contrast to responses to body displacements, responses to direct head displacements appear to depend upon a vestibulospinal trigger, since trunk and leg muscle responses to head displacements were absent in patients who had lost vestibular function as adults. Patients who lost vestibular function as infants, however, had near normal trunk and leg response to head displacements, suggesting a substitution of upper trunk and neck somatosensory inputs for missing vestibular inputs during development.
    Type of Medium: Electronic Resource
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  • 10
    Electronic Resource
    Electronic Resource
    Springer
    Experimental brain research 118 (1998), S. 126-130 
    ISSN: 1432-1106
    Keywords: Key words Split-belt locomotion ; Afferent input ; Body loading ; Locomotor adaptation ; Human gait
    Source: Springer Online Journal Archives 1860-2000
    Topics: Medicine
    Notes: Abstract  The modification of the normal locomotor pattern of humans was investigated using a split-belt locomotion protocol (treadmill belt speeds of 4.5 km/h and 1.5 km/h for the right and left legs, respectively) and also by changing afferent input from the legs (30% reduction or increase in body weight by suspending subjects in a parachute harness or by wearing a lead-filled vest). After a control-speed training period (10 min) of symmetrical walking (3 km/h each leg) and a period (10 min) of split-belt walking, the adjustment back to the control speed resulted in a mean speed difference between the right leg and the left leg of 0.85 km/h. Adjustment of belt speed on either side was performed by the hands using a potentiometer. For comparison, also speed adjustment by the feet via feedback derived from changes in the treadmill drive current was studied. No significant difference was obtained when both modes of adjustment were compared. Body unloading or loading during the training period resulted in an improved adjustment of treadmill belt speed. This suggests that load receptor information plays a major role in the programming of a new walking pattern.
    Type of Medium: Electronic Resource
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