Proactive and reactive mechanisms play a role in stepping on inverting surfaces during gait.

Ankle inversions have been studied extensively during standing conditions. However, inversion traumas occur during more dynamic conditions, like walking. Therefore in this study sudden ankle inversions were elicited in 12 healthy subjects who stepped on a trap door while walking on a treadmill. First, 10 control trials (0 degrees of rotation) were presented. Then, 20 stimulus (25 degrees of rotation) and control trials were presented randomly. EMG recordings were made of six lower leg muscles. All muscles showed a short-latency response (SLR) of about 40 ms and a late-latency response (LLR) of about 90 ms. The peroneal muscles showed the largest amplitudes in both responses. The functionally more important, larger, and more consistent LLR response was too late to resist the induced stretch during the inversion. The functional relevance of this response must lie after the inversion. During the first trial a widespread "startle-like" coactivation of the LLR was observed. The last trials showed only a recruitment of the peroneal muscles and, to a lesser extent, the gastrocnemius lateralis, which is seen as a switch from reactive control to more proactive adaptive strategies. These proactive strategies were investigated separately by comparing trials in which inversion was expected (but did not occur) with those in which subjects knew that no inversion would occur. Anticipation of a possible inversion was expressed in decreased tibialis anterior activity before initial contact, consistent with a more cautious and stable foot placement. Furthermore, immediately after landing, the peroneal muscles were activated to counteract the upcoming stretch.

Cutaneous reflexes evoked during human walking are reduced when self-induced.

Reflex responses are often less pronounced when they are self-induced, but this question has barely been investigated quantitatively. The issue is particularly relevant for locomotion since it has been shown that reflexes elicited during normal gait are important for the regulation of locomotion. The cortex is thought to be involved in the control of reflexes during gait, but it is unclear whether it plays a role in the modulation of these reflexes during the step cycle. During gait, weak electrical stimulation of the sural nerve elicits reflexes in various leg muscles. Are these reflexes different when subjects themselves trigger the stimuli instead of being randomly released by the computer? Cutaneous reflexes were elicited by sural nerve stimulation in 16 phases of the gait cycle in healthy subjects. The stimuli were triggered either by computer or by the subjects themselves. In 6 out of 7 subjects it was observed that the facilitatory responses in leg muscles were smaller and the suppressive responses were more suppressive following self-generated stimuli. In some muscles such as tibialis anterior (TA) both effects were seen (reduced facilitation at end stance and exaggerated suppression at end swing). In all subjects the modulation of anticipatory influences was muscle specific. In the main group of six subjects, the mean reduction in reflex responses was strongest in the TA (max. 30.7%; mean over 16 phases was 12.5%) and weakest in peroneus longus (PL, max. 10.1%; mean over 16 phases was 2.6%). The observation that facilitation is reduced and suppression enhanced in several muscles is taken as evidence that anticipation of self-induced reflex responses reduces the excitatory drive to motoneurones, for example through presynaptic inhibition of facilitatory reflex pathways.

Startle responses in Parkinson patients during human gait.

Falls frequently occur in patients with Parkinson's disease (Bloem et al. 2001). One potential source for such falls during walking might be caused by the reaction to loud noises. In normal subjects startle reactions are well integrated in the locomotor activity (Nieuwenhuijzen et al. 2000), but whether this is also achieved in Parkinson patients is unknown. Therefore, in the present study, the startle response during walking was studied in eight patients with Parkinson's disease and in eight healthy subjects. To examine how startle reactions are incorporated in an ongoing gait pattern of these patients, unexpected auditory stimuli were presented in six phases of the step cycle during walking on a treadmill. For both legs electromyographic activity was recorded from biceps femoris and tibialis anterior. In addition, we measured the stance and swing phases of both legs, along with the knee angles of both legs and the left ankle angle. In all subjects and all muscles, responses were detected. The pattern of the responses, latency, duration, and phase-dependent modulation was similar in both groups. However, the mean response amplitude was larger in patients due to a smaller habituation rate. No correlation was found between the degree of habituation and disease severity. Moreover, a decreased habituation was already observed in mildly affected patients, indicating that habituation of the startle response is a sensitive measure of Parkinson's disease. The results complement the earlier findings of reduced habituation of blink responses in Parkinson's disease. With respect to behavioral changes in healthy subjects we observed that startle stimuli induced a shortening of the step cycle and a decrease in range of motion. In the patient group, less shortening of the subsequent step cycle and no decrease in range of motion of the knee and ankle was seen. It is argued that the observed changes might contribute to the high incidence of falls in patients with Parkinson's disease.

Reflex responses in the lower leg following landing impact on an inverting and non-inverting platform.

In the lower leg, landing after a jump induces reflexes, the role of which is not well understood. This is even more so for reflexes following landing on inverting surfaces. The latter condition is of special interest since ankle inversion traumata are one of the most common injuries during sport. Most studies have investigated ankle inversions during a static standing condition. However, ankle injuries occur during more dynamic activities such as jumping. Therefore, the present study aimed at reproducing these situations but in a completely safe setting. EMG responses were recorded after landing on an inverting surface, which caused a mild ankle inversion of 25 deg of rotation (in a range sufficient to elicit reflexes but safe enough to exclude sprains). The results are compared with data from landing on a non-inverting surface to understand the effect of the inversion. In general, landing on the platform resulted in short and long latency responses (SLR and LLR) in triceps surae (soleus, gastrocnemius medialis and lateralis) and peroneal muscles (long and short peroneal) but not in the tibialis anterior muscle. Landing on the inverting platform caused significant LLRs in the peroneal muscles (which underwent the largest stretch) but not in the triceps muscles. Conversely, landing on a non-inverting platform induced larger SLRs in triceps than in the peroneal muscles. Although the peroneal LLRs thus appeared to be selectively recruited in an inverting perturbation, their role during such perturbations should be limited since the latency of these responses was about 90 ms while the inversion lasts only 42 ms. The SLRs, if present, had an onset latency of around 44 ms. In the period following the inversion, however, the responses may be important in preventing further stretch of these muscles.

Mechanically induced ankle inversion during human walking and jumping.

A new method to study sudden ankle inversions during human walking and jumping is presented. Ankle inversions of 25 degrees were elicited using a box containing a trap door. During the gait task, subjects walked at a speed of 4 km/h. At a pre-programmed delay after left heel strike, an electromagnet released the box on the treadmill. This delay enabled the subject to step on the box without having to change the walking cadence. During the jumping task, subjects jumped from a 30 cm high platform on the box in a standardised way. In both tasks 20 stimulus and 20 control trials were presented randomly. The average tilting velocity of the trap door during the stimulus trials was 403 degrees /s during the walking task and 595 degrees /s during the jumping task. For the control trials a tilting of 0 degrees was used. With this method it is possible to evoke reproducible ankle inversions causing characteristic EMG responses in six lower leg muscles.

Dynamic posturography using a new movable multidirectional platform driven by gravity.

Human upright balance control can be quantified using movable platforms driven by servo-controlled torque motors (dynamic posturography). We introduce a new movable platform driven by the force of gravity acting upon the platform and the subject standing on it. The platform consists of a 1 m2 metal plate, supported at each of its four corners by a cable and two magnets. Sudden release of the magnets on three sides of the platform (leaving one side attached) induces rotational perturbations in either the pitch or roll plane. Release of all magnets causes a purely vertical displacement. By varying the slack in the supporting cables, the platform can generate small (0.5 degrees ) to very destabilising (19 degrees ) rotations. Experiments in healthy subjects showed that the platform generated standardised and reproducible perturbations. The peak rotation velocity well exceeded the threshold required to elicit postural responses in the leg muscles. Onset latencies were comparable to those evoked by torque motor-driven platforms. Randomly mixed multidirectional perturbations of large amplitude forced the subject to use compensatory steps (easily possible on the large support surface), with little confounding influence of habituation. We conclude that this gravity-driven multidirectional platform provides a useful and versatile tool for dynamic posturography.