Mg. Carpenter et al., Directional sensitivity of stretch reflexes and balance corrections for normal subjects in the roll and pitch planes, EXP BRAIN R, 129(1), 1999, pp. 93-113
A large body of evidence has been collected which describes the response pa
rameters associated with automatic balance corrections in man to perturbati
ons in the pitch plane. However, perturbations to human stance can be expec
ted from multiple directions. The purpose of the present study was to descr
ibe the directional sensitivities of muscle responses re-establishing distu
rbed stance equilibrium in normal subjects. The contributions of stretch re
flex and automatic balance-correcting responses to balance control, and con
comitant biomechanical reactions, were examined for combinations of pitch a
nd roll perturbations of the support surface. More specifically, muscle res
ponses, initial head accelerations and trunk velocities were analyzed with
the intention of identifying possible origins of directionally specific tri
ggering signals and to examine how sensory information is used to modulate
triggered balance corrections with respect to direction. Fourteen healthy a
dults were required to stand on a dual-axis rotating platform capable of de
livering rotational perturbations with constant amplitude (7.5 degrees) and
velocity (50 degrees/s) through multiple directions in the pitch and roll
planes. Each subject was randomly presented with 44 support surface rotatio
ns through 16 different directions separated by 22.5 degrees first under ey
es-open, and then, for a second identical set of rotations, under eyes-clos
ed conditions. Bilateral muscle activities from tibialis anterior, soleus,
lateral quadriceps and paraspinals, were recorded, averaged across directio
n, and areas, calculated over intervals with significant bursts of activity
. Trunk angular velocity and ankle torque data were averaged over intervals
corresponding to significant biomechanical events. Stretch reflex (interva
ls of 40-100, 80-120 ms) and automatic balance-correcting responses (120-22
0, 240-340 ms) in the same muscle were sensitive to distinctly different di
rections. The directions of the maximum amplitude of balance-correcting act
ivity in leg muscles were oriented along the pitch plane, approximately 180
degrees from the maximum amplitude of stretch responses. Ankle torques for
almost all perturbation directions were also aligned along the pitch plane
. Stretch reflexes in paraspinal muscles were tuned along the 45 degrees pl
ane but at 90 degrees to automatic balance corrections and 180 degrees to u
nloading responses in the same muscle. Stretch reflex onsets in paraspinal
muscles were observed at 60 ms, as early as those of soleus muscles. In con
trast, unloading reflexes in released paraspinal muscles were observed at 4
0 ms for perturbations which caused roll of the trunk towards the recorded
muscle. Onsets of trunk roll velocities were earlier and more rapid than th
ose: observed for pitch velocities. Trunk pitch occurred for pure roll dire
ctions but not vice versa. When considered together, early stretch and unlo
ading of paraspinals,;md concomitant roll and pitch velocities of the trunk
requiring a roll-and-pitch-based hip torque strategy, bring into question
previous hypotheses of an ankle-based trigger signal or ankle-based movemen
t strategies for postural balance reactions. These findings are compatible
with the hypothesis that stretch-, force- and joint-related proprioceptive
receptors at the level of the trunk provide a directionally sensitive trigg
ering mechanism underlying a minimally two-stage (pitch-based leg and pitch
-and-roll-based trunk) balance-correcting strategy. Accelerometer recording
s from the head identified large vertical linear accelerations only for pit
ch movements and angular roll accelerations only during roll perturbations
with latencies as early as 15 ms.
Thus, it appears that balance corrections in leg and trunk muscles may rece
ive strong, receptor-dependent (otolith or vertical canal) and directionall
y sensitive amplitude-modulating input from vestibulospinal signals.