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The Journal of Neurophysiology Vol. 84 No. 1 July 2000, pp. 65-74
Copyright ©2000 by the American Physiological Society
1Departement of Medical Physics and Biophysics, University of Nijmegen, 6525 EZ Nijmegen; and 2Nijmegen Institute for Cognition and Information (NICI), Motor Behavior and Rehabilitation Research, University of Nijmegen, 6500 HE Nijmegen, The Netherlands
Nieuwenhuijzen, P.H.J.A.,
A. M. Schillings,
G. P. Van Galen, and
J. Duysens.
Modulation of the Startle Response During Human
Gait. J. Neurophysiol. 84: 65-74, 2000. While
many studies have shown that there is a phase-dependent modulation of
proprioceptive and exteroceptive reflexes during gait, little is known
about such modulation for auditory reflexes. To examine how startle
reactions are incorporated in an ongoing gait pattern, unexpected
auditory stimuli were presented to eight healthy subjects in six phases
of the step cycle during walking on a treadmill at 4 km/h. For both
legs, electromyographic activity (EMG) was recorded in the biceps
femoris (BF), the rectus femoris (RF), the tibialis anterior (TA), and
the soleus (SO). In addition, stance and swing phases of both legs,
along with knee angles of both legs and the left ankle angle, were
measured. All subjects showed various response peaks. Responses with
latencies of ~60 ms (F1), ~85 ms (F2), and ~145 ms (F3) were
found. The amplitude of the reflex responses was dependent on the
timing of the startle stimulus in the step cycle. Although the startle
response habituated rapidly, the phase-dependent modulation pattern
generally remained the same. The phase-dependent amplitude modulations
were not strictly correlated with the modulation of the background
activity. The TA even showed a transition from facilitatory F2
responses during stance to suppressive responses during midswing.
Responses were observed in both flexors and extensors, often in
coactivation, especially during stance. Furthermore the gait
characteristics showed a shortening of the subsequent step cycle and a
small decrease in the range of motion of ankle and knees. These results
suggest that the responses are adapted to achieve extra stability
dependent on the phase of the step cycle. However, even in the first
trials, the changes in kinematics were small allowing a smooth
progression of gait.
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