J Physiol 565.

2 (2005) pp 675684 675

Reversals of anticipatory postural adjustments
during voluntary sway in humans
Vijaya Krishnamoorthy
1,2
and Mark L. Latash
1
1
Department of Kinesiology, The Pennsylvania State University, University Park, PA 16802, USA
2
Department of Physical Therapy, University of Delaware, Newark, DE 19716, USA
We describe reversals of anticipatory postural adjustments (APAs) with the phase of a voluntary
cyclic whole-body sway movement. Subjects (n =9) held a standard load in extended arms and
released it by a bilateral shoulder abduction motion in a self-paced manner at different phases
of the sway. The load release task was also performed during quiet stance in three positions: in
the middle of the sway range and close to its extreme forward and backward positions. Larger
APAs were seen during the sway task as compared to quiet stance. Although the direction of
postural perturbation associated with the load release was always the same, the direction of the
APAs in the leg muscles reversed when the subjects were close to the extreme forward position
as compared to the APAs in other phases and during quiet stance. The trunk muscles showed
smaller APA modulation at the extreme positions but larger modulation when passing through
the middle position, depending onthe directionof sway, forwardor backward. The phenomenon
of APAreversals emphasizes theimportant roleof safetyinthegenerationof postural adjustments
associatedwithvoluntary movements. Basedonthese ndings, APAs couldbe denedas changes
in the activity of postural muscles associated with a predictable perturbation that act to provide
maximal safety of the postural task component.
(Resubmitted 8 February 2005; accepted after revision 21 March 2005; rst published online 24 March 2005)
Corresponding author M. L. Latash: Rec. Hall - 267L, Department of Kinesiology, Penn State University, University
Park, PA 16802, USA. Email: mll11@psu.edu
Upright human posture is inherently unstable with the
challenge of balancing the body, whichhas a relatively high
centre of mass (COM) and several joints along the body
axis, on a narrow base of support (Hayes, 1982; Winter
et al. 1996, 1998). When voluntary movements are made
while standing, the redistribution of mass and inertial
forces threatenbalance. Anticipatory postural adjustments
(APAs) are changes in the activity of postural muscles and
associated shifts in the centre of pressure (COP; point
of application of ground reaction forces) prior to the
initiation of action (Belenkii et al. 1967; Marsden et al.
1979; Cordo & Nashner, 1982; Bouisset & Zattara, 1987;
for a review, see Massion, 1992).
It has been assumed that APAs are based on an estimate
of the effects of an expected perturbation on standing
balance. It has also been assumed that the purpose of
APAs is to generate forces and moments opposing the
mechanical effects of the expected postural perturbation
(Bouisset & Zattara, 1987, 1990; Friedli et al. 1988).
Humans frequently perform fast arm movements and
manipulate objects while moving the body, e.g. during
walking, standing up, or leaning. In particular, subjects
pulling on a handle while walking exhibit APAs that differ
across the gait cycle (Nashner &Forssberg, 1986; Forssberg
& Hirschfeld, 1988; Hirschfeld & Forssberg, 1991). In
general, these ndings have supported the idea that APAs
act to oppose the effects of forthcoming perturbations.
A load release task has been used in many APA studies
(Aruin & Latash, 1996; Slijper et al. 2002; Aruin &
Shiratori, 2004; Slijper & Latash, 2004). This task leads
to reproducible APAs in most postural muscles. It has a
certain advantage as compared to voluntary movement
tasks since the magnitude of the perturbationdoes not vary
with the natural variability in the voluntary movement
that triggers the load release. We used the load release task
to examine APAs when subjects stood at different angles
of body lean and while subjects voluntarily swayed. The
results of our study show that APAs can reverse direction
according to the lean of the body during the sway task.
It therefore appears that APAs are not always organized
to oppose the mechanical effects of expected postural
perturbations.
Methods
Subjects
Nine healthy right-handed subjects, without any known
neurological or motor disorder, participated in the
C
The Physiological Society 2005 DOI: 10.1113/jphysiol.2005.084772
676 V. Krishnamoorthy and M. L. Latash J Physiol 565.2
experiment. Of these, ve were males and four
were females, with the mean age of 29.5 4.6 years,
mean weight of 63.6 10.2 kg and mean height of
167.1 9.6 cm (means s.d.). The subjects gave written
informedconsent accordingtothe procedures approvedby
the Ofce for Regulatory Compliance of the Pennsylvania
State University. The study was performed according to
the Declaration of Helsinki.
Apparatus
A force platform (AMTI, OR-6) was used to record the
moment around a frontal axis (M
y
) and the vertical
component of the reaction force (F
z
). An oscilloscope
(Tektronics TDS 210) showed the time pattern of M
y
to the subject and the experimenter. A uni-directional
accelerometer (Sensotec) was taped to the dorsal aspect
of the subjects hand over the third metacarpal joint,
such that the axis of sensitivity of the accelerometer was
directedalongthe requiredmotionof the hand. Disposable
self-adhesive electrodes (3MRedDot, 3M Health Care, St.
Paul, MN) were used to record the surface EMG activity
of the following leg and trunk muscles: tibialis anterior
(TA), soleus (SOL), rectus femoris (RF), biceps femoris
(BF), rectus abdominis (RA) and erector spinae (ES). The
electrodes were placed on both sides of the subjects body
on the muscle bellies, with their centres approximately
3 cm apart. Data were recorded at the sampling frequency
of 1000 Hz with 12-bit resolution. A Gateway 450 MHz
PC with customized software based on the LabVIEW4
package (National Instruments) was used to control the
experiment and collect the data.
Procedure
During all experiments, the subjects held a 3 kg load
(20 cm20 cm10 cm; approximately 4% of subjects
body mass) in front of him/her by pressing on the sides of
the load with both hands. Pilot studies showed that this
load was large enough to bring about reproducible APAs
without causing fatigue over repeated trials. The shoulders
were at 90 deg exion and the elbows were fully extended.
The load was released under two task conditions: quiet
stance (QS) and voluntary sway (VS).
In the QS condition, subjects initially stood on the force
platform with their feet side-by-side, a hip width apart.
They were requiredtorelease the loadwitha quickbilateral
shoulder abduction movement. Subjects were instructed
to stand as quietly as possible in the initial position before
the beginning of the trial. Then, the computer generated a
beep 500 ms after data collection began, which indicated
that the subject should initiate the required action in a
self-paced manner. Subjects were reminded not to initiate
their actions immediately after the beep, but to wait
for about a second, so that reaction time did not lead
to APA modication (cf. De Wolf et al. 1998; Slijper
et al. 2002). Data were collected over 3 s for each trial.
Subjects performed three series, six repetitions each, at the
QS condition in three positions: leaning forward (QS
F
),
neutral standing upright (QS
M
) and leaning backward
(QS
B
). The degree of body lean in QS
F
and QS
B
was
such that the COP shift from the average value obtained
during QS
M
was between 1 and 2 cm corresponding to
typical COP shifts during the voluntary sway series (see
below). For a given subject and for a given direction
of the lean, the lean magnitude was reproduced across
trials by marking the M
y
position on the oscilloscope.
Subjects occupied this initial position prior to every
trial.
In the VS condition, the initial position of the sub-
ject was the same as in the QS
M
condition (vertical
neutral stance). M
y
at the neutral body position was
marked on the oscilloscope, and the subject was asked
to occupy this position prior to each trial. Then, subjects
were instructed to shift their body weight towards their
toes and heels rhythmically, thereby swaying forward and
backward. They were asked to match the times of the peak
forward and backward positions with the beats of the
metronome. The metronome frequency was set at 1 Hz,
so that the subject completed one full cycle of forward
and backward sway in 2 s or at the frequency of 0.5 Hz.
Subjects were asked to watch the oscilloscope, which
showed them the current value of M
y
. The required M
y
shift was also marked on the oscilloscope (approximately
10 Nm in each direction, which corresponded to COP
shifts of about 12 cm depending on the subjects body
weight). Thus, the degree of lean in the peak forward and
backward M
y
positions of the VS task corresponded to the
degree of lean in the QS
F
and QS
B
tasks.
For each trial, data were collected over 8 s. On hearing
the computer-generated beep, the subject was instructed
to begin swaying to the beat of the metronome. After
completing 23 full sway cycles, the subject released the
load at a self-selected time. The subjects were instructed to
release the load at different phases of the sway, randomly
varying across the trials to have approximately equal
numbers of trials withloadrelease close tothe twoextreme
positions (forwardandbackwardmaximal lean) andinthe
middle position while moving forward and backwards.
The total of 32 trials were collected in three blocks of
8 trials with a rest period of at least a minute between the
blocks. Pilot series showed that this instruction resulted in
subjects releasing the load at different phases of sway while
not forcing them to attend to particular cues. We wanted
to keep the load release self-paced and did not provide
specic cues at different phases of the cycle, as APAs are
known to be modied in reaction time tasks (e.g. De Wolf
et al. 1998; Slijper et al. 2002).
The order of the QS and VS conditions was balanced
across subjects. Rest periods of 8 s betweentrials and2 min
between the two conditions were given. Fatigue was not
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The Physiological Society 2005
J Physiol 565.2 APA reversals during voluntary sway 677
an issue. Prior to each condition, two practice trials were
given.
In addition to the main trials, two control trials were
performed. The subject was asked to hold the load of 3 kg
and sway to the beat of the metronome without releasing
the load (VS
bl
). These data were used to compute the EMG
activity of the leg and trunk muscles in the absence of the
perturbationcaused by the load release, as described inthe
next subsection.
Data processing
All signals were processed off-line and ltered with a
50 Hz low-pass, fourth-order, zero-lag Butterworth lter
using LabVIEW 4 to avoid phase distortions in the signal.
All EMG signals were rectied. Individual trials were
viewed on the monitor screen and aligned according to
the rst visible change in the signal of the accelerometer
(movement initiation). This moment will be referred
to as time zero (t
0
). Changes in the muscle activity
associated with APAs were quantied as follows. In the
QS trials, rectied EMG signals were integrated from
100 ms prior to t
0
to t
0
(

EMG). Although in some

conditions APAs were partly outside this time window,
it was not practical to adjust the windowof integration for
different muscles, subjects, and conditions. So we used a
conservative approach of EMG analysis within a standard
100 ms time window. Such an approach has been used in
many earlier studies of APAs (e.g. Shiratori &Latash, 2000;
Slijper & Latash, 2000). These integrals were corrected by
subtracting integrated activity at rest from 500 to
450 ms prior to t
0
multiplied by two to match the
intervals of EMG integration (the baseline EMG activity,

EMG
blqs
).

EMG
QS
=

t
0
100
EMG 2

t
450
500
EMG
blqs
(1A)
Inthe VStrials, the most forwardpositions of the M
y
signal
were identied and marked. Then all the data were cut
into cycles such that each cycle began at the most forward
position of M
y
and ended one sample before the next
most forward M
y
position. The time of load release within
each trial was computed as a percentage of the sway cycle,
with 0% representing the most forward position and 50%
representing the most backward position. The following
four ranges of the cycle time were used to assign a trial to
one of the four groups:
(1) Forward (VS
F
): 87100% and 05%
(2) Backward (VS
B
): 4055%
(3) Middle position while swaying forward: (VS
MF
):
7078%
(4) Middle position while swaying backward: (VS
MB
):
2028%
In general, subjects were able to release the load close
to the middle position far more often than close to the
extreme positions. In order to get a reasonable number of
trials from each subject for statistical purposes, we used
a wider range of percentages for the extreme positions.
These intervals were also slightly skewed in the direction
leading to an extreme position rather than away fromit, as
subjects dropped the load more often moving towards
these extreme positions than when moving away from
them.
Similar to the data processing of QS trials, for the
VS trials, rectied EMG signals were integrated from
100 ms prior to t
0
to t
0
. These integrals were corrected by
subtracting baseline EMG, which was computed from the
unperturbed VS (VS
bl
) trial over a 100 ms interval prior to
the phase of the sway cycle when the load was released
(

EMG
blvs
), that is, corresponding to the same time
interval as in the VS trial.

EMG
VS
=

t
0
100
EMG

t
0
100
EMG
blvs
(1B)
In order to compare the integrated EMG indices (

EMG)
across muscles and subjects, we normalized them by the
highest absolute value of

EMG values for each muscle,

across trials for each subject (

EMG
control
) as follows:

EMGindices for a muscle froma given trial were divided

by

EMG
control
, suchthat all EMGs were between1and1.
Negative values corresponded to a decrease in the EMG
activity during APAs.
IEMG =

EMG
QS,VS

EMG
control
(2)
To illustrate the change in pattern of IEMGs as a function
of sway cycle, IEMGs were calculated at different phases
of the VS task and arranged in order of their appearance
in the sway cycle expressed as a percentage of the cycle
duration. In order to minimize the effects of outliers, we
used a 5-point moving average window, i.e. the averages
over 5 consecutive data points within the sway cycle.
Coordinates of the centre of pressure (COP) in the
anteriorposterior directions were calculated using the
following approximation:
COP = M
Y
/F
Z
(3)
Statistics
Statistical methods included repeated-measures
multivariate analysis of variance (MANOVA). To
determine the effects of TASK (QS versus VS) and
POSITION (leaning forward, F or backward, B) on IEMG
indices of APAs, we performed two-way MANOVAs on
the IEMG indices for each pair of muscles acting at
each joint (TA/SOL, RF/BF, or RA/ES). Three separate
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The Physiological Society 2005
678 V. Krishnamoorthy and M. L. Latash J Physiol 565.2
two-way MANOVAs on APAs of muscle pairs from each
joint were run: MANOVA-Ia, on IEMG of TA and SOL;
MANOVA-Ib on IEMG of RF and BF and MANOVA-Ic
on IEMG of RA and ES.
In order to test if there was an effect of different task
conditions on the APAs when the load was released at
the middle position, a one-way MANOVA (MANOVA-II)
was performed with CONDITIONas a factor having three
levels: QS
M
, VS
MB
and VS
MF
. Again, we performed three
MANOVAs on the IEMG indices for each pair of muscles
acting at the three joints (TA/SOL, RF/BF or RA/ES).
Planned comparisons were performed to compare the
effects of QS and VS tasks. To test specic differences
-80
0
80
160
240
320
-60
-30
0
30
60
90
-0.4 -0.2 0 0.2 0.4
-50
-25
0
25
50
-0.4 -0.2 0 0.2 0.4
Time (s)
-0.4 -0.2 0 0.2 0.4
Quiet standing
A Forward (QS
F
) B Middle (QS
M
) C Backward (QS
B
)
Tibialis anterior
Soleus
Rectus femoris
Biceps femoris
Rectus abdominis
Erector spinae
E
M
G

(
m
V
)
E
M
G

(
m
V
)
E
M
G

(
m
V
)
Figure 1. General EMG patterns in a representative subject during the quiet stance (QS) task
The load was released in the forward (A: QS
F
), middle (B: QS
M
) and backward (C: QS
B
) positions. Time zero
corresponds to the time of load release, and the two vertical lines indicate the 100 ms time interval over which
indices of anticipatory postural activity (APA) were quantied. EMG signals for soleus, biceps femoris and erector
spinae are inverted for better visualization.
between pairs of conditions, Tukeys honest signicant
difference (HSD) tests were used.
Results
Muscles on the left and right sides of the body showed
qualitatively similar results in all analyses. Therefore for
simplicity, only the results of analyses onthe muscles of the
left side of the body are presented and discussed further.
General EMG patterns: quiet stance (QS)
When a person stood in a neutral upright position and
held the load in extended arms, typically there was an
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The Physiological Society 2005
J Physiol 565.2 APA reversals during voluntary sway 679
increase in the activity of the dorsal muscles (SOL, BF
and ES). When the person released the load with a quick
bilateral shoulder abduction motion, prior to the load
release, there was adropinthe activityof the dorsal muscles
accompanied sometimes by an increase in the activity of
the ventral muscles (TA, RF and RA). A typical pattern
is illustrated for a representative subject in Fig. 1B. Time
zero corresponds to the initiation of load release. APAs
were quantied within a 100 ms time interval prior to this
time (the time period between the two vertical lines in
Fig. 1). Not all subjects showed changes in muscle activity
in all muscles.
Asimilar patternwas seenwhenthesubject stoodquietly
while leaning forward or backward, i.e. on the toes or
the heels (Fig. 1A and C, respectively), but the early EMG
changes were usually less marked and more variable across
subjects. In particular, Fig. 1A shows no visible RF burst.
In Fig. 1C, the RF burst was smaller than while standing in
the neutral position (Fig. 1B) and there is no suppression
of the SOL activity.
General EMG patterns: voluntary sway (VS)
EMGpatterns associated with anunperturbed cycle (VS
bl
)
of voluntary sway when holding a load with extended
arms are presented in Fig. 2 for a representative subject. A
cycle beginning from the most forward position followed
by sway backward towards the heels and a return to the
forward position is illustrated (see M
y
in the top panel).
Since the frequency of sway was 0.5 Hz, half a cycle took
about 1 s in time. Therefore, times 0 and 2 s in Fig. 2
correspond to the extreme forward position, and time 1 s
corresponds to the extreme backward position. Note the
increase in the activity of the ventral muscles (TA, RF)
when approaching the extreme backward position and a
simultaneous drop in the activity of the dorsal muscles
(SOL, BF, ES). Conversely, when approaching the extreme
forward position, there is an increase in the activity of
the dorsal muscles, accompanied by a drop in the ventral
muscle activity.
Changes in the muscle activity associated with release
of the load (APAs) at four points during the sway cycle are
illustrated in Fig. 3 for a representative subject. Similarly
to Fig. 1, time zero corresponds to the initiation of load
release, and APAs were quantied within a 100 ms time
interval prior to this time, by subtracting the integrated
activity at the same phase and over the same time
interval during an unperturbed sway trail. Figure 3B and
D shows changes in the activity of muscles when the load
was released close to the neutral position during sway,
while passing the neutral position during swaying in the
backward and forward directions, respectively. Figure 3A
and C shows EMGs when the load was released close to
the extreme forward and backward positions, respectively,
during sway. Note the anticipatory increase in the activity
of ventral muscles (TA, RF, and to a lesser extent RA)
when the load was released while moving towards the
extreme backward position (Fig. 3B; VS
MB
) and while
passing through this position (Fig. 3C; VS
B
). This was
accompanied by an anticipatory drop in the activity of
the dorsal muscles (SOL, BF and ES). In contrast, if the
load was released while moving towards (Fig. 3D; VS
MF
)
or passing through the extreme forward position (Fig. 3A;
Baseline EMG during voluntary sway
- 5
- 2 . 5
0
2 . 5
5
F M B B M F F
- 6 0
- 4 0
- 2 0
0
2 0
4 0
6 0
8 0
E
M
G

(
m
V
)
- 1 2 0
- 1 0 0
- 8 0
- 6 0
- 4 0
- 2 0
0
2 0
E
M
G

(
m
V
)
- 6 0
- 5 0
- 4 0
- 3 0
- 2 0
- 1 0
0
1 0
0 0 . 5 1 1 . 5 2
Time (s)
E
M
G

(
m
V
)
Soleus
Erector spinae
Biceps femoris
Rectus abdominis
Rectus femoris
Tibialis anterior
M
y

(
N

m
)
Figure 2. EMG activity associated with an unperturbed cycle of
voluntary sway (VS) in a representative subject
The top panel shows the moment about the frontal axis (M
y
), where
the highest positive value corresponds to the extreme forward position
and the smallest negative value corresponds to the extreme backward
position during VS. The remaining three panels show EMG patterns in
the lower leg, thigh and trunk muscles, respectively. EMG signals for
soleus, biceps femoris and erector spinae muscles are inverted for
better visualization. Note that the ventral muscles (tibialis anterior,
rectus femoris, rectus abdominis) show an increase in the activity
during motion towards the backward position and the dorsal muscles
(soleus, biceps femoris, erector spinae) show an increase in activity
during motion towards the forward position.
C
The Physiological Society 2005
680 V. Krishnamoorthy and M. L. Latash J Physiol 565.2
VS
F
), there was an anticipatory increase in the activity of
the dorsal muscles and an anticipatory drop in the activity
of the ventral muscles. These APAs also varied in different
muscles across subjects.
Figure 4 shows smoothed IEMG indices (see eqn (2)
in the Methods) plotted as a function of phase of the
sway cycle for a representative subject. Recall that 0 and
100% correspond to the most forward position and 50%
to the most backward position. A third-order polynomial
was tted to the data in Fig. 4 and the squared regression
coefcient (R
2
) values are presented. In general, the ankle
(TA, SOL) and knee muscles (RF, BF) showed similar
directions of changes in APAs over the sway cycle. The
anterior muscles, TA and RF, showed an increase in
-120
0
120
240
360
480
-100
0
100
200
300
-80
-40
0
40
80
-0.4 -0.2 0 0.2 0.4 -0.4 -0.2 0 0.2 0.4 -0.4 -0.2 0 0.2 0.4 -0.4 -0.2 0 0.2 0.4
Voluntary sway
E
M
G

(
m
V
)
E
M
G

(
m
V
)
E
M
G

(
m
V
)
Tibialis anterior
Soleus
Rectus fermoris
Biceps femoris
Rectus abdominis
Erector spinae
A Forward
(VS
F
)
C Backward
(VS
B
)
D Middle-
Forward
(VS
MF
)
B Middle-
backward
(VS
MB
)
Time (s)
Figure 3. General EMG patterns in a representative subject during the voluntary sway (VS) task
The load was released in the forward (A: VS
F
), middle when moving backward (B: VS
MB
), backward (C: VS
B
) and
middle when moving forward (D: VS
MF
) positions. Time zero corresponds to time of load release and the vertical
lines indicate the 100 ms time interval before load release over which indices of anticipatory postural activity (APA)
were quantied. EMG signals for soleus, biceps femoris and erector spinae are inverted for better visualization.
APAs in the mid-backward position (passing through the
middle position while moving backwards, 25% of the
cycle), reaching the highest values at the extreme back-
ward position (50% of the cycle) followed by a drop
in APAs in the mid-forward position. (passing through
the middle position while moving forward, 75% of
the cycle) and the smallest APAs in the extreme forward
positions (0/100% of the cycle). The posterior muscles,
SOL and BF, showed out-of-phase APA changes with
respect to APA changes in TA and RF. The trunk muscles,
RA and ES, showed a different pattern of activity. Both
RA and ES showed larger APAs in the mid-forward and
mid-backward positions, and smaller APAs at the extreme
positions.
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The Physiological Society 2005
J Physiol 565.2 APA reversals during voluntary sway 681
Effect of task on APAs in extreme body positions
Figure 5 shows IEMGs of individual muscles averaged
across subjects with standard error bars for the QS and
VS tasks in the extreme forward and backward positions.
Note that the direction of APAs in the forward and back-
wardpositions is opposite inall the muscles. This effect was
small for the QS task, but was more striking in the VS task.
The lower leg and thigh muscles showed signicant main
effects of TASK (QS and VS) and POSITION (extreme
forward and extreme backward; both main effects Wilks
lambda <0.4; F
2,7
>6; P <0.05). SOL and RF muscles,
in particular showed signicantly larger APAs in the VS
condition as compared to QS (P <0.05). The ventral
muscles (TA and RF) showed an increase in IEMGs in the
backward position (P <0.05), while the dorsal muscles
(particularly SOL) showed an increase in IEMGs in the
forward position (P <0.05; see also Figs 1 and 3). There
R
2
= 0.9078
-1
-0.5
0
0.5
1
1.5
2
0 20 40 60 80 100
R
2
= 0.5976
-0.05
0
0.05
0.1
0.15
0.2
0.25
0 20 40 60 80 100
R
2
= 0.8304
-3
-2
-1
0
1
2
3
4
5
6
0 20 40 60 80 100
T
A

I
E
M
G

I
n
d
e
x
R
F

I
E
M
G

I
n
d
e
x
R
A

I
E
M
G

I
n
d
e
x
Percentage of VS cycle
80 20
R
2
= 0.9375
-1.5
-1
-0.5
0
0.5
1
1.5
2
0 20 40 60 80 100
R
2
= 0.851
-0.8
-0.7
-0.6
-0.5
-0.4
-0.3
-0.2
-0.1
0
0.1
0.2
0.3
0 40 60 100
R
2
= 0.8644
-0.8
-0.7
-0.6
-0.5
-0.4
-0.3
-0.2
-0.1
0
0 20 40 60 80 100
S
O
L

I
E
M
G

I
n
d
e
x
B
F

I
E
M
G

I
n
d
e
x
E
S

I
E
M
G

I
n
d
e
x
Percentage of VS cycle
F MB B MF F F MB B MF F
Figure 4. Pattern of IEMG change for APAs of individual muscles of a representative subject over a cycle
of voluntary sway (VS)
Ventral muscles: TA, tibialis anterior; RF, rectus femoris; RA, rectus abdominis. Dorsal muscles: SOL, soleus; BF, biceps
femoris; ES, erector spinae. 0 and 100%, extreme forward position; 50%, extreme backward position; 25%, middle
position when moving backward; and 75%, middle position when moving forward. Data were smoothed with a
5-point moving average window and tted with a third-order polynomial. R
2
values are presented.
was a signicant TASKPOSITION interaction effect for
the TA/SOL muscle pair (Wilks lambda <3; F
2,7
>10;
P <0.01), while this interaction effect was just below
signicance for the RF/BF pair (P =0.064). Even though
the trunk muscles, RA and ES, showed a difference in the
magnitudes of APAs in the different phases of the VS cycle
(see Fig. 4), these effects varied across subjects, and these
muscles did not show any signicant TASK or POSITION
effects.
Effect of task on APAs in the middle body position
To analyse the effects of sway on APAs when the subjects
were close to the middle position, IEMG indices for the
APAs were compared across three conditions: (1) in the
middle position during quiet stance (QS
M
), (2) while
passing the middle position in the forward direction
during VS (VS
MF
), and (3) while passing the middle
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682 V. Krishnamoorthy and M. L. Latash J Physiol 565.2
position in the backward direction (VS
MB
). Averaged
across subjects normalized IEMG indices are shown in
Fig. 6 with standard error bars. Note the pronounced
suppression of the muscle activity (negative values) in the
dorsal muscles for QS
M
(see also Fig. 1). During voluntary
sway, this suppression became less pronounced. In the
ventral muscles, a tendency towards higher APAs was
observed. MANOVA on IEMG over all pairs of muscles
(TA/SOL, RF/BF and RA/ES) showed a signicant main
effect of CONDITION (QS
M
, VS
MB
and VS
MF
; Wilks
lambda <0.2; F
4,5
>6; P <0.05). Planned comparisons
between the QS
M
and the two VS conditions revealed that
APAs in SOL, BF, RA and ES were signicantly larger in
-0.8
-0.6
-0.4
-0.2
0.0
0.2
0.4
0.6
0.8
QS VS
R
F

I
E
M
G

i
n
d
e
x
-0.6
-0.4
-0.2
0.0
0.2
0.4
0.6
QS VS
B
F

I
E
M
G

i
n
d
e
x
-0.6
-0.4
-0.2
0.0
0.2
0.4
0.6
QS VS
Backward
Forward
T
A

I
E
M
G

i
n
d
e
x
-0.8
-0.6
-0.4
-0.2
0.0
0.2
0.4
0.6
0.8
QS VS
Backward
Forward
S
O
L

I
E
M
G

i
n
d
e
x
-0.6
-0.4
-0.2
0.0
0.2
0.4
0.6
QS VS
R
A

I
E
M
G

i
n
d
e
x
-0.6
-0.4
-0.2
0.0
0.2
0.4
0.6
QS VS
E
S

I
E
M
G

i
n
d
e
x
*
*
*

TASK
TASK
Figure 5. IEMG indices of individual muscles in the quiet stance (QS) and voluntary sway (VS) tasks in
the extreme positions
Average across subjects IEMG indices in the extreme forward (open columns) and backward (lled columns)
positions are shown with standard error bars. Muscle names are abbreviated as in Fig. 4. Signicant differences
between the QS and VS tasks;

signicant differences between the extreme forward and backward positions.
the VS conditions as compared to QS (F
1,8
>5; P <0.05;
indicated by in Fig. 6). Further, SOL and ES APAs were
signicantly higher during VS
MF
as compared to QS
M
.
ES also showed signicantly higher APAs during VS
MF
as
compared to VS
MB
(indicated by

in Fig. 6).
Discussion
The magnitude of anticipatory postural adjustments
(APAs) is known to depend on the magnitude of the
perturbation, its direction and postural stability (Nouillot
et al. 1992; Aruin & Latash, 1995; Aruin et al. 1998; Dietz
et al. 2000). In a series of studies, Forssberg and colleagues
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The Physiological Society 2005
J Physiol 565.2 APA reversals during voluntary sway 683
(Nashner & Forssberg, 1986; Forssberg & Hirschfeld,
1988; Hirschfeld & Forssberg, 1991) have shown that
APA magnitude is modulated within the gait cycle. Their
subjects were asked to respond to an audio signal that was
presented in different phases of the step cycle with a strong
pull on the handle. Lateral gastrocnemius and hamstring
muscles were activated during APAs occurring in the early
support phase whereas tibialis anterior and quadriceps
muscles were activatedwhenthe pull was exertedjust prior
to and during the swing phase.
Interpretationof these ndings is complicated, however,
by the fact that the leg muscles were involved in two
activities, locomotion and APAs. In particular, in the
swing phase, the muscles of the leg could not contribute
to postural stabilization since they had no contact with
the support. It is hard therefore to assess the functional
signicance of the apparently reversed APAs seen in
the swing phase. Our experiments have demonstrated
reversals of APA action when both legs could contribute
to postural stabilization in all phases of the sway cycle.
APA reversals in such conditions represent a major novel
nding of our study.
The role of postural stability: rening
the APA denition
Typically, the magnitude of APAs decreases for both very
stable conditions (e.g. standing while leaning against
a wall, Friedli et al. 1984) and unstable conditions
(e.g. standing on a narrowsupport area, Aruin et al. 1998).
Our experiments conrmed observations by Aruin et al.
(1998) that shiftingthe bodyweight forwards or backwards
leads to a decrease in the magnitude of APAs associated
with releasing a standard load from the extended arms.
Releasing a load held in the extended arms always tends
to tilt the body backwards. Hence, in the most backward
positionintheswaycycle, this perturbationwas morelikely
to push the body out of the limits of postural stability. In
contrast, the same perturbation tended to move the body
away fromthe dangerously close edge of stability when the
subjects approached the most forward body posture.
The typical APA pattern associated with load release
(Aruin et al. 1998; Fig. 1) always tends to tilt the trunk
forward. When the body is close to the most forward
position, this APA pattern would move the body closer to
the limit of stability and, as such, might by itself violate the
balance. In such a situation, the CNS reverses the direction
of the APAs (Fig. 5) and moves the body away from the
edge of the support area. Mechanical action of reversed
APAs does not counteract the mechanical effects of the
perturbation associated with the load release but acts in
the same direction.
Traditionally APAs have been dened as changes in
the patterns of muscle activity that produce forces and
moments acting against the mechanical effect of an
expected perturbation (Bouisset & Zattara, 1987; Friedli
et al. 1988; Massion, 1992). Our experiments have shown
that APAs can produce mechanical effects on posture
acting together with the perturbation as long as such
action is consistent with maximizing the safety of the
posture. We therefore suggest that APAs couldbe denedas
changes inthe activityof postural muscles associatedwitha
predictable perturbation that act to provide maximal
safety of the postural task component. This denition
is consistent with the ndings of decreased APAs under
postural threat or fear of falling (Adkin et al. 2002).
One view on APAs is that they result from an internal
forward model that takes into account dynamic
consequences of an expected perturbation and generates
responses to counteract these consequences (Wing et al.
1997). However, we donot see howaforward-model-based
APA would be able to account for the reversal of APA
direction, whichis basedonsafety concerns. Analternative
interpretationis that predictivemotor control mechanisms
such as APAs are not based on computations of forces but
ensure stability of balance using equilibrium-point control
(Feldman & Latash, 2005); shifts in equilibrium position
of the body during APAs may be based on a combination
of factors, including closeness to the edge of the support.
The role of the magnitude of the perturbation
In our experiments, the actual perturbation associated
with the load release could be expected to vary with
the phase of the sway because of the modulation of
-0.6
-0.4
-0.2
0.0
0.2
0.4
0.6
TA SOL RF BF RA ES
QS
M
VS
MB
VS
MF
I
E
M
G

I
n
d
e
x

*
Muscle
Figure 6. Average across subjects IEMG indices of individual
muscles in the middle positions
Quiet stance middle position (QS
M
; lled columns), voluntary sway
middle position when swaying backwards (VS
MB
; open columns) and
when swaying forward (VS
MF
; hatched columns) are shown with
standard error bars. Muscle names are abbreviated as in Fig. 4.
Signicant differences between the QS and VS tasks;

signicant
differences between VS
MB
and VS
MF
.
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The Physiological Society 2005
684 V. Krishnamoorthy and M. L. Latash J Physiol 565.2
inertial forces. This modulation was relatively small. Peak
horizontal load acceleration during a sine sway at 2 Hz
with the amplitude of load motion of about 0.5 m may be
assessed as close to 0.5 ms
2
. With respect to the ankle
joint, the lever arm of the load was about 1 m. For the
3 kg load, the peak inertial torques at the ankle joints
related to the load motion were of the order of 1.5 Nm.
The gravitational force acting on the load produced the
moment about the ankle joint of about 20 Nm. Hence,
the motion-related modulation of the magnitude of the
perturbation at the two extreme positions was under 10%.
Large differences in the APAmagnitude were seen in the
phases of moving through the middle posture in different
directions (Fig. 6), particularly in the RA/ES muscle pair
that has been viewed as providing a basic pattern of the
APAs (Shiratori &Latash, 2000). At those phases, the body
movedat the highest speedwhile the accelerationwas close
to zero. Hence, this modulation could not be related to
the motion-related modulation of the magnitude of the
inertial forces.
The observed differences in the APAs while moving
through the mid-posture in opposite directions could be
related to preparing a response while taking into account
the fact that the body was moving towards one of the
extreme positions. This makes it possible to relate changes
in the APAs in all muscle pairs (cf. Fig. 5) to providing
maximal safety for the postural task.
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