G.A. Horstmann

Gating and reversal of reflexes in ankle muscles during human walking

SummaryPhase-dependent reflex modulation was studied by recording the electromyographic (EMG) responses in ankle flexors (Tibialis Anterior, TA) and extensors (Gastrocnemius Medialis, GM and Soleus, SOL) to a 20 ms train of electrical pulses, applied to the tibial or sural nerve at the ankle, in human volunteers walking on a treadmill at 4 km/h. For low intensity stimuli (i.e. 1.6 times perception threshold), given during the swing phase, the most common response was a suppression of the TA activity with a latency of 67 to 118 ms. With high intensity of stimulation (i.e. 2.8 × T), a facilitatory response appeared in TA with a latency of 74 ms. This latter response was largest during the middle of the swing phase, when it was correlated with exaggerated ankle dorsiflexion. The TA reflex amplitude was not a simple function of the level of spontaneous ongoing activity. During stance, TA responses were small or absent and accompanied by a suppression of the GM activity with a latency ranging from 62 to 101 ms. A few subjects showed an early facilitatory, instead of a suppressive, GM response (88 to 136 ms latency). They showed a phase-dependent reflex reversal from a dominant TA response during swing to a facilitatory GM response with an equivalent latency during stance. The GM facilitation occurred exclusively during the early stance phase and habituated more than the TA responses. It is concluded that phase-dependent gating of reflexes occurs in ankle muscles of man, but only when vigorous extensor reflexes are present. More commonly, a phase-dependent modulation is seen, both of facilitatory and suppressive responses.

Human postural reflexes and gravity — An under water simulation

This study represents the first attempt to investigate the influence of gravity on postural adjustments. Subjects were displaced while standing under water on a movable platform, while the buoyancy of the body was adjusted by using a variety of lead vests. Under water, an approximately linear relationship was found between body weight and impulse directed electromyographic response amplitudes in the leg and thigh muscles. Loading of the subjects out of water resulted in a saturation of the response amplitude. The biomechanical signals recorded during the displacements indicated that neither vestibulospinal nor muscle proprioceptive reflex mechanisms can account for the effect observed under water. It is suggested that the EMG responses are mediated by reflexes which are activated by pressure receptors within the body in order to hold the centre of gravity over the feet.

Fast head tilt has only a minor effect on quick compensatory reactions during the regulation of stance and gait.

SummarySudden tilts of the head to the front or rear were induced during stance, balancing, gait and during perturbations of gait. The most prominent response in the leg muscle electromyogram (e.m.g.) to head tilt occurred in the tibialis anterior muscle (latency about 55 ms) following a backward tilt induced during balancing. During stance and gait, the e.m.g. activity related to head tilt was only a minor component of the leg muscle activity normally occurring during gait. When the head tilt was induced shortly after a perturbation of gait (treadmill acceleration impulse), the compensatory reaction in the leg muscles did not significantly differ from that seen after the gait perturbation alone. In addition, the rate of acceleration of the head was tested against the compensatory e.m.g. responses: No correlation of influence could be discerned. The results indicate that sudden head tilts and the resulting head acceleration have little influence on the e.m.g. patterns that occur during gait and perturbations of gait. It is assumed that these patterns are regulated by central programs, and that the compensation for leg perturbation is achieved mainly by spinal reflex mechanisms. It is discussed whether the lack of head tilt responses is the result of an antagonistic vestibularneck interaction, or whether it indicates a reduced effectiveness of vestibulo- and cervico-spinal reflexes during gait.

Compensation of human stance perturbations: selection of the appropriate electromyographic pattern.

Perturbations of stance evoke purposive EMG patterns which are directed to hold the bodys centre of gravity over the feet. Dorsiflexing rotation of the feet is followed by a monosynaptic stretch reflex response in the gastrocnemius muscle, succeeded by a late compensatory tibialis anterior activation. Backward translation of the feet elicits only a compensatory polysynaptic EMG response in the gastrocnemius muscle, while an early gastrocnemius response is absent. The amplitude modulation of the gastrocnemius H-reflex has been investigated during the early part of the two modes of perturbation. Only during translational perturbation a progressive decrease in gastrocnemius H-reflex amplitude started within 5 ms after onset of displacement. The degree of the reduction in amplitude in the former perturbation was dependent on the displacement velocity. Only the contact forces (torques) differed between the two modes of perturbations within the first 10 ms after onset of perturbations. It is suggested that signals from pressure receptors within the body are responsible for the early change in H-reflex amplitude during translational perturbations and it is concluded that the simplest spinal reflex is under very rapid and powerful moment-to-moment control by changes in peripheral feedback. In view of a strong reciprocal modulation of monosynaptic and polysynaptic reflex responses, the later purposive EMG responses may be determined by early changes in presynaptic inhibition of group I afferents.

The contribution of vestibular input to the stabilization of human posture: A new experimental approach

An experiment was designed to evaluate the vestibular contribution to the stabilization of upright stance in normals and in two patients with loss of vestibular function. A forward or backward displacement of a load (2 kg) by a torque motor attached to the subject induced opposing movements in the head and trunk. The small linear acceleration of the head in space of about 0.1 g was followed, with a latency of 50-65 ms, by EMG responses in the tibialis anterior and rectus femoris (backward acceleration) or gastrocnemius muscles (forward acceleration). These responses were absent in patients with a vestibular deficit. It is suggested that the observed EMG responses are due to fast acting vestibulospinal reflexes involved in the regulation of upright stance. For comparable head accelerations the integrated EMG responses induced by the vestibulospinal mechanism are about ten times smaller than those induced by spinal stretch reflexes during displacement of the feet. Vestibulospinal reflexes would appear, therefore, to play only a minor role in the compensation of stumbling.

Involvement of different receptors in the regulation of human posture

Compensatory electromyographic (EMG) responses and several biomechanical parameters were studied following impulsive disturbance of the lower limbs during stance on a treadmill. Treadmill acceleration impulses were backwards or forwards directed, or their direction was inverted after 30 ms. Backwards directed impulses were followed by gastrocnemius and forwards directed ones by tibialis anterior EMG responses (latency 65-75 ms) whose duration depended on impulse duration. When the direction of the impulse was inverted, the respective antagonistic leg muscles were activated, with a delay of 68 to 75 ms after onset of stretch of these muscles. The behaviour of the EMG responses could best be correlated to the displacement at the ankle joint and may be described in terms of a stretch reflex response. The function of this stretch reflex mechanism is suggested to be connected with the control of the bodys centre of gravity in order to prevent falling. Head movements induced by the impulses showed little correlation with the appearance of the EMG responses, suggesting that the vestibular system is unlikely to be significantly involved in the generation of these responses.

Chapter 34 Significance of proprioceptive mechanisms in the regulation of stance

Compensatory electromyographic (EMG) responses and several biomechanical parameters were studied following impulsive disturbance of the limbs during stance of human volunteers on a treadmill. Treadmill acceleration impulses were backwards or forwards directed, or their initial direction was reversed after 30 ms. Backwards directed impulses were followed by gastrocnemius, forwards directed ones by tibialis anterior EMG responses (latency 65 to 75 ms) whose durations depended on impulse duration. When the direction of the impulse was reversed, the respective antagonistic leg muscles were activated again with a delay of 68 to 75 ms after onset of stretch of these muscles. The behaviour of the EMG responses could best be correlated to the displacement at the ankle joint and may be described in terms of a stretch reflex response. The results indicate that these stretch reflex responses help control of the bodys centre of gravity thereby preventing falling. Head movements induced by the impulses showed little correlation with the appearance of the EMG responses, suggesting that the vestibular system is unlikely to be directly involved in the generation of these responses. Vestibular signals may, however, significantly contribute to slow body sway stabilization.

Reproducibility and adaptation of the EMG responses of the lower leg following perturbations of upright stance.

The electromyographic (EMG) response of gastrocnemius, soleus and anterior tibialis muscles to a backwardly directed perturbation of stance was recorded in 12 normal subjects using surface electrodes and studied with regard to its reproducibility (test-retest reliability coefficients, variability coefficients) and to adaptational effects. (1) Reproducibility was shown to be uniformly high and can be interpreted as an index for the high intraindividual constancy of the results. (2) Adaptational effects have been found and should be circumvented, either by pre-adapting the subjects to the motor task, or by restriction of the period of measurement. (3) Variation of the position of the electrodes produced only small effects. The impedance of the surface electrodes was not critical if kept below 5 k omega. EMG investigations with surface electrodes during stance and perturbations of stance provide highly reliable results with respect to intraindividual changes but interindividual variability of the results clearly marks the limits of this method. The interindividual variability observed with surface electrodes is of the same order as that reported in the literature for inserted needle recording.

Body Sway Stabilization in Human Posture

Electromyographic (emg) responses and joint movements of the leg were analysed in subjects standing with eyes closed on a sinusoidally moving treadmill (0.16 Hz or 0.33 Hz, amplitude 33 cm). Activity in antagonistic leg muscles was reciprocally modulated, with a predominant gastrocnemius activation during deceleration of forward movement and tibialis anterior activation during deceleration of backward movement of the treadmill. In these phases, it was necessary to compensate for sway induced by body inertia. The match between treadmill movement and emg activity was better for the gastrocnemius than for the tibialis anterior muscle. The characteristic pattern of leg muscle emg activity is suggested to be modulated predominantly by vestibulo-spinal reflexes partly because treadmill movements did not evoke muscle strength, and partly because patients with loss of vestibular function showed basic alterations in the emg pattern and could only compensate for the slow sinus while standing unsupported.

Interlimb coordination of stance in children: Divergent modulation of spinal reflex responses and cerebral evoked potentials in terms of age

EMG responses in the gastrocnemius (GM) and tibialis anterior muscles (TA) of both legs together with cerebral evoked potentials (CP), were recorded following perturbations of stance on a treadmill with split belts, in two age groups of children. Unilateral displacements were followed by ipsilateral short latency and bilateral long latency EMG responses. The CP was similar in both tasks. When displacements were simultaneously induced in opposite directions, a significant reduction in the long latency components of EMG responses occurred, while the amplitude of the CP was maximal in this condition. In the older children the CP and long latency EMG responses were larger and the short latency reflex potentials smaller in all conditions compared to the younger children. It is concluded that (1) CP and EMG responses reflect a divergent modulation of a given somatosensory input; (2) developmental changes are reflected in alterations in the amplitude of CP and EMG responses; (3) there is no evidence of transcortically mediated muscle responses.