Jl. Smith et al., FORMS OF FORWARD QUADRUPEDAL LOCOMOTION - III - A COMPARISON OF POSTURE, HINDLIMB KINEMATICS, AND MOTOR PATTERNS FOR DOWNSLOPE AND LEVEL WALKING, Journal of neurophysiology, 79(4), 1998, pp. 1702-1716
To gain further insight into the neural mechanisms for different forms
of quadrupedal walking, data on postural orientation, hindlimb kinema
tics, and motor patterns were assessed for four grades of downslope wa
lking, from 25% (14 degrees slope) to 100% (45 degrees), and compared
with data from level and downslope walking at five grades (5-25%) on t
he treadmill (0.6 m/s). Kinematic data were obtained by digitizing cin
e film, and electromyograms (EMGs) synchronized with kinematic records
were taken from 13 different hindlimb muscles. At grades from 25 to 7
5%, cycle periods were similar, but at the steepest grade the cycle wa
s shorter because of a reduced stance phase. Paw-contact sequences at
all grades were consistent with lateral-sequence walking, but pace wal
king often occurred at the steepest grades. The cats crouched at the s
teeper grades, and crouching was associated with changes in fore-and h
indlimb orientation that were consistent with increasing braking force
s and decreasing propulsive forces during stance. The average ranges o
f motion at the hindlimb joints, except at the hip, were often differe
nt at the two steepest slopes. During swing, the range of knee- and an
kle-joint flexion decreased, and the range and duration of extension i
ncreased at the ankle joint to lower the paw downward for contact. Dur
ing stance the range of flexion during yield increased at the ankle jo
int, and the range of extension decreased at the knee and metatarsopha
langeal joints. Downslope walking was also associated with EMG changes
for several muscles. The hip extensors were not active during stance;
instead, hip flexors were active, presumably to slow the rate of hip
extension. Although ankle extensors were active during stance, their b
urst durations were truncated and centered around paw contact. Ankle f
lexors were active after midstance at the steeper slopes before the ne
ed to initiate swing, whereas flexor and extensor digit muscles were c
oactive throughout stance. Overall the changes in posture, hindlimb ki
nematics, and activity patterns of hindlimb muscles during stance refl
ected a need to counteract external forces that would accelerate angul
ar displacements at some joints. Implications of these changes are dis
cussed by using current models for the neural control of walking.