Muscular responses and movement strategies during stumbling over obstacles.
J. Neurophysiol. 83: 2093-2102, 2000. Although many studies have investiga
ted reflexes after stimulation of either cutaneous or proprioceptive affere
nts, much less is known about responses after more natural perturbations, s
uch as stumbling over an obstacle, in particular, the phase dependency of t
hese responses and their relation to the stumbling behavior has received li
ttle attention. Hence response strategies during stumbling reactions after
perturbations at different times in the swing phase of gait were studied. W
hile subjects walked on a treadmill, a rigid obstacle unexpectedly obstruct
ed the forward sway of the foot. All subjects showed an "elevating strategy
" after early swing perturbations and a "lowering strategy" after late swin
g perturbations. During the elevating strategy, the foot was directly lifte
d over the obstacle through extra knee flexion assisted by ipsilateral bice
ps femoris (iBF) responses and ankle dorsiflexion assisted by tibialis ante
rior (iTA) responses. Later, large rectus femoris (iRF) activations induced
knee extension to place the foot on the treadmill. During the lowering str
ategy, the foot was quickly placed on the treadmill and was lifted over the
obstacle in the subsequent swing. Foot placement was actively controlled b
y iRF and iBF responses related to knee extension and deceleration of the f
orward sway. Activations of iTA mostly preceded the main ipsilateral soleus
(iSO) responses. For both strategies, four response peaks could be disting
uished with latencies of similar to 40 ms (RP1), similar to 75 ms (RP2), si
milar to 110 ms (RP3), and similar to 160 ms (RP4). The amplitudes of these
response peaks depended on the phase in the step cycle. The phase-dependen
t modulation of the responses could not be accounted for by differences in
stimulation or in background activity and therefore is assumed to be premot
oneuronal in origin. In mid swing. both the elevating and lowering strategy
could occur. For this phase, the responses of the two strategies could be
compared in the absence of phase-dependent response modulation. Both strate
gies had the same initial electromyographic responses till similar to 100 m
s (RP1-RP2) after perturbation. The earliest response (RP1) is assumed to b
e a short-latency stretch reflex evoked by the considerable impact of the c
ollision, whereas the second (RP2) has features reminiscent of cutaneous an
d proprioceptive responses. Both these responses did not determine the beha
vioral response strategy. The functionally important response strategies de
pended on later responses (RP3-RP3). These data suggest that during stumbli
ng reactions, as a first line of defense, the CNS releases a relatively asp
ecific response, which is Followed by an appropriate behavioral response to
avoid the obstacle.