Neurophysiologie Clinique/Clinical Neurophysiology (2008) 38, 345—362

REVIEW/MISE AU POINT

Posture, dynamic stability, and voluntary movement

Posture, stabilité dynamique et mouvement
volontaire
S. Bouisset ∗, M-C. Do

UFR-STAPS, université Paris-Sud-11, rue Langevin, 91405 Orsay, France

Received 1st October 2008; accepted 1st October 2008

Available online 18 October 2008

KEYWORDS Summary This paper addresses the question of why voluntary movement, which induces a
Voluntary movement; perturbation to balance, is possible without falling down. It proceeds from a joint biomechanical
Postural stabilization; and physiological approach, and consists of three parts. The first one introduces some basic
Postural chain; concepts that constitute a theoretical framework for experimental studies. The second part
Newton’s laws; considers the various categories of ‘‘postural adjustments’’ (PAs) and presents major data on
Dynamic stability; ‘‘anticipatory postural adjustments’’ (APA). The last part explores the concept of ‘‘posturo-
Anticipatory postural kinetic capacity’’ (PKC) and its possible applications.
adjustments (APAs); © 2008 Elsevier Masson SAS. All rights reserved.
Postural
programming;
Posturo-kinetic
capacity (PKC);
Foco-kinetic capacity
(FKC);
paraplegics;
Lower limb amputees

Résumé Cet article aborde la question de savoir pourquoi le mouvement, qui induit une per-
MOTS CLÉS turbation de l’équilibre corporel, s’avère possible sans entraîner de chute. On se situera dans
Mouvement une double perspective associant approche biomécanique et physiologique. La première partie
volontaire ; introduit quelques concepts de base servant de cadre théorique aux études expérimentales. La
Stabilisation seconde considère les diverses catégories d’« ajustements posturaux » et présente les princi-
pales données concernant les « ajustements posturaux anticipateurs » (APA). Enfin, la troisième

∗ Corresponding author.
E-mail address: simon.bouisset@u-psud.fr (S. Bouisset).

0987-7053/$ – see front matter © 2008 Elsevier Masson SAS. All rights reserved.
doi:10.1016/j.neucli.2008.10.001
346 S. Bouisset, M.-C. Do

posturale ; partie explore le concept de « capacité posturo-cinétique » (CPC, ou PKC en anglais) et ses
Chaîne posturale ; possibles applications.
Lois de Newton ; © 2008 Elsevier Masson SAS. All rights reserved.
Stabilité dynamique ;
Ajustements
posturaux
anticipateurs (APAs) ;
Programmation
posturale ;
Capacité
posturocinétique
(CPC) ;
Capacité
fococinétique (CFC) ;
Paraplégiques ;
Amputés de membre
inférieur

Conceptual framework maintenance’’ (which is presented as ‘‘static’’), occur-

ring in response to gravity and ground reaction. The
Physiological concepts other is ‘‘postural stabilization’’ (which is presented as
‘‘dynamic’’), occurring in response to a perturbation. Motor
development helps explain the difference: infants learn
Both the concept of voluntary movement and the distinction
first to sit, then to stand, and then to walk with increas-
between posture and movement underlie the approach.
ing efficiency: ‘‘static’’ equilibration is acquired before
‘‘dynamic’’ equilibration. Both constitute ‘‘equilibration
Motor task and voluntary movement
reactions’’, i.e. the physiological response to balance per-
A movement is called ‘‘voluntary’’ (‘‘intentional’’ or ‘‘goal
turbation.
directed’’) when its intent is to perform a given ‘‘task’’. Its
This chapter focuses on postural stabilization. More
achievement requires the mobilization and immobilization
precisely, neurophysiological control mechanisms differ
of body segments.
depending on whether the perturbation origin is exogenous
A system analysis, limited to the motor side of the pro-
(‘‘reaction equilibration’’) or endogenous (‘‘action equili-
cess, is useful in presenting the main steps leading to task
bration’’) [1]. In other words, recovery from an unexpected
achievement (Fig. 1). The ‘‘task to be performed’’, which is
fall and the execution of a voluntary movement do not pro-
the ‘‘intended task’’, can be characterized by the general
ceed from the same control mechanisms. The roles played by
category to which it belongs, such as pointing, throwing,
the postural phenomena associated to voluntary movement
walking, etc. . . It is also characterized by its specific param-
are illustrated in Fig. 2, which summarizes Hess’ views [44]:
eters, such as speed, amplitude, orientation or precision
(i) postural maintenance is different from postural stabiliza-
(including the initial and final conditions), and its envi-
tion; (ii) voluntary movement is a perturbation to posture,
ronmental context (ground elasticity or friction, wind or
and consequently to body balance; and (iii) voluntary move-
water resistance, etc.). The task to be performed is the
ment and related posture cooperate in the organization of
input to the sensory-motor system. The ‘‘real task’’, i.e.
the motor act.
the task that is actually performed, is the output of the
More recently, in accordance with Bernstein’s ideas [9],
system. It results from a hierarchical process associating
Gelfand et al. [38] hypothesized that motor control includes
three different levels. The real task fulfills the intended
two components: a ‘‘focal’’ and a ‘‘postural’’ one. The for-
task requirements with varying degrees of efficiency. Effi-
mer refers to the body segments that are mobilized in order
ciency is evaluated by ‘‘performance’’, which is measured
to perform voluntary movement, and the latter to the rest of
by the actual parameter values (speed, precision, etc.) with
the body, which is involved in the stabilizing reactions. The
respect to the intended ones.
body partitioning between a ‘‘focal chain’’ and a ‘‘postural
To summarize, a voluntary movement is part of a more
chain’’ [15] is a direct consequence of this assumption.
general process, called the ‘‘motor act’’. In other words, a
voluntary movement is the means to complete a motor task.
Biomechanical concepts
Equilibration function in posture and movement Newton’s laws
According to classical views, the motor system includes two ‘‘Despite the great variability in bodily form and general
types of organization: one referring to posture and the other pattern of propulsive movements, the number of basic
to movement. This duality has been stressed as far back as principles encountered during the study of vertebrate loco-
Babinski [3] and André-Thomas [1]. motion are relatively few. . . These basic principles are all
Postural activity, or better, ‘‘equilibration’’, is known incorporated in Newton’s laws of motion’’ [39]. More pre-
to present two different aspects. One is ‘‘postural cisely, Newton’s laws apply to all categories of mechanical
Posture, dynamic stability, and voluntary movement
Figure 1 From the task to perform to the real task: outline
of a system analysis. The ‘‘task to perform’’ is the task that the
subject intends to execute. It can be characterized by three fac-
tors: category, specific parameters and environmental context.
These different factors are taken into account at the program-
ming level. The central nervous system issues a command that
triggers inputs to the muscular system. The muscles mobilize
the bone and joint system, which produces movement. The
‘‘real task’’ is the task, which is actually performed. In order
to characterize motor activity in a non invasive manner, periph-
eral variables are measured at the muscular (electromyographic
activity: EMG) and movement (biomechanical quantities: kine-
matic and kinetic ones) levels. Movement efficiency is evaluated
by the actual specific parameters values (speed, precision, etc,
from [11]).
structures, both inanimate and animate ones, such as the
human body.
Newton’s laws are expressed by two equations (Fig. 3):
one (the dynamic resultant law) refers to linear movements,
and the other (the kinetic torque law), to rotations. They •
relate forces to movement at every instant. Indeed, on the
left side of the equal sign, there is an acceleration (to within
a constant coefficient) and on the right side, an expression •
of external forces.
More precisely, Eq. (1) relates the external forces resul-
tant (˙ Fe ) to the dynamic resultant (mG ), with G being •
the centre of gravity (CoG) linear acceleration and m, body
mass. Eq. (2) relates the resultant torque of external forces
347
→ →
 G (Fe )) to the kinetic momentum (JG −−
(˙ M  ), with −− being
angular acceleration and JG the body moment of inertia with
respect to the CoG1 . The first terms of Eqs. (1) and (2),
preceded by the minus sign, define the ‘‘inertial
→ forces’’ sys-
tem: −mG is the ‘‘inertial force’’, and −JG −−
 , the ‘‘inertial
torque’’.
In order to present some consequences of Newton’s laws
simply, the argument will use primarily the first equation.
The human body, as a whole
A direct consequence of Newton’s laws is that the movement
of the CoG (or around it) only results from forces, which are
‘‘external’’ to the structure under consideration. On the
ground track, the forces, which are external to the human
body considered as a whole, are often limited to gravity and
support reaction forces. As emphasized by Gray [39], ‘‘if the
body of an animal is at rest relative to its environment, it
can only be set in motion by the application of an external
force . . . Similarly, an external environmental force must be
brought into play if an animal is to change speed or direction
of motion’’.
External and internal forces. Two consequences can be
derived from the above comments.
First, when the human body is considered as a whole,
muscular forces are ‘‘internal’’ to the mechanical system.
As such, they cannot determine the overall movement of the
body; they can only provoke segmental movements.
Second, the global centre of gravity can only move
if the body encounters resistance originating from the
environment, usually a physical support, which offers the
appropriate reaction, in addition to gravity (Fig. 3). Pro-
vided that there is an external reaction to their action, the
rotation of body segments produced by the muscles leads, in
turn, to movements of the centre of gravity (or around it).
In other words, if a muscular contraction provokes a joint
movement—–upper limb flexion for instance—–the rest of the
body tends to move in the opposite direction. As expressed
by Gray [39]: ‘‘In order to subject the body to a propul-
sive force, it is necessary to exert an exactly equal but
opposite force against his external environment. The reac-
tion is equal to the subject’s action («action and reaction
principle»)’’.
In other words, voluntary movement occurs as the result
of a combination of muscle torques and ground reactions.
However, even though they are internal, muscular forces
play a major role. As they are under the control of the
CNS, they are the only ones that can be used to manage
the fatal effects of the external forces, and adjust coopera-
→
1
 = ˙M
Eq. (2) (JG −−  G (Fe )) refers to a solid, rotating about a fixed
axis. In this equation:
→ →
the kinetic momentum (JG −−  ) is the angular acceleration (−−
)
times the body moment of inertia (JG ), with respect to the rota-
tion axis, , which goes through the body center of gravity, G;
the body moment of inertia (JG ) (with respect to the rotation
axis, , which goes through G) is the body mass times the squared
distance from G to the rotation axis;
the resultant torque of external forces (˙ M  G (Fe )) is the resultant
force (˙ Fe ) times its distance to the rotation axis, , which goes
through G.
348 S. Bouisset, M.-C. Do

Figure 2 Hess’s model. The model includes three components, each being represented by a figure: the jumper (1) is standing
on the bearer’s shoulders (2), and the bearer is secured by a supporter (3) during the jump. The jumper represents the voluntary
movement, while the bearer and the supporter represent the ‘‘static’’ and ‘‘dynamic’’ equilibration reactions. Left column (a, b,
c): the task is performed efficiently, owing to adequate postural dynamics, and a ‘‘static’’ posture is recovered at the end of the
jump. Right column (d, e, f): the task is not performed efficiently, as the subject falls at the end of jump: postural stabilization is
not efficient enough to counteract transient disequilibrium (from [43]).

tion between internal and external forces in such a way that
performance fulfils task-movement constraints.
Support base and support reaction forces. Contrary to
gravity, reaction forces originating from the ground (and
more generally from the physical environment) vary as a
function of the forces developed by the muscles during the
motor act. Therefore, a clear understanding of the postural
phenomena associated with voluntary movement requires
taking the external forces into account, and the support
reaction forces in particular. This necessity requires con-
sidering the ‘‘support base’’, i.e. the body areas in contact
with the physical support.
Support base characteristics depend on two factors, the
physical properties of the support and the interface between
the body and the support. The first depends on the proper-
ties of the materials (elasticity, friction, etc.), as well as on
geometrical parameters (flatness or curvature, inclination,
etc.) and design (steadiness or oscillations, translations,
etc.). The second depends on the subject’s posture (stand-
ing, on tiptoe, on the entire foot, on one foot or both feet,
seated, with or without a backrest, etc). They also depend
on the support base perimeter.
Biomechanical principles indicate that support base char-
acteristics have an effect on the reaction forces applied
to the body (and on their torque). For example, by con-
struction, the support base perimeter delimits Centre of
Pressure (CoP) displacement, insofar as the CoP is the point
where the vertical forces resultant is applied. Now, CoG hor-
izontal acceleration has been shown to be proportional to
the difference between CoP and CoG horizontal displace-
ments [21]. Therefore, CoG horizontal acceleration may be
reduced when the support base perimeter is limited and
when the CoP initial position is close to the support base
boundaries (Fig. 4).
The human body, as a system of rigid solids
The human body is composed of anatomical ‘‘segments’’,
including bones and the living tissues around them, such as
the arm or the leg. From a biomechanical point of view,
it can also be considered a system of rigid solids, termed
‘‘links’’ [28], which are movable in relation to each other,
i.e. they form an ‘‘articulated chain’’ and interact. The
mobility of an articulated chain is a function of the num-
ber of the degrees of freedom and of the range of individual
joint movements.
Postural maintenance is an ‘‘active’’ process. A certain
equilibrium state, often termed ‘‘static’’, is associated to
given posture maintenance. According to Newton’s laws, it
corresponds to the condition when the resultant reaction
and the resultant torque of the external forces are nil. This
necessity implies two sets of requirements.
The first are global requirements: they are the same
as if the body were an invariant structure, i.e. as if the
body segment configuration were mechanically constrained.
Posture, dynamic stability, and voluntary movement 349

Figure 3 Illustration of Newton’s laws in shoulder abduction performed in two support base conditions. A. Top inset: Newton’s
laws, (1): ‘‘dynamic resultant law’’, (2) ‘‘kinetic torque law’’. Eq. (1) relates the external forces resultant to the dynamic resultant,
which is the center of gravity (CoG) linear acceleration times body mass (m). Eq. (2) relates the resultant moment of external forces
to the kinetic moment, which is the angular acceleration times the body moment of inertia (JG ) with respect to the CoG. A: lower
inset: when the subject is at rest (˙ Fe = 0 and ˙ M G (Fe ) = 0), for instance during quiet standing, he is subjected to gravity (P) and
ground reaction (R) forces. One is applied to the center of gravity (Go), and the other to the center of pressure (Io). The center of
pressure (CoP) is the barycenter of the minute ground reaction forces applied to the subject’s contact areas, such as Ri . B (middle
inset), upper part: the feet are in contact with the ground. During shoulder abduction, there is a displacement of the CoG, from Go
to G1, and of the CoP, from Io to I1. B. Lower part: external forces balance sheet: during the movement (˙ Fe =  G (Fe ) =
/ 0 and ˙ M / 0),
the subject is always subjected to gravity (P) and ground reaction (R) forces. However, the ground reaction forces display a vertical
(RN ) and a horizontal (RT ) component, which vary at each instant. C. Right inset, upper part: the feet are no longer in contact
with the ground and gravity force is nil (as in a space lab), the upper limb and the rest of the body move in opposite directions.
The body’s center of gravity no longer moves (G1 = Go) and there is no longer a center of pressure, as there is no contact between
the subject and the ground. Right inset, lower part: external forces balance sheet: the subject is no longer subjected to any force
(from [11]).

They result in the usual contention according to which gen-
eral body balance is secured when the projection on the
ground of the centre of gravity is situated within the support
base perimeter. The second are local requirements, which
are usually understated. They result from the articulated
structure of the human body: each body part must also be
balanced with respect to the underlying ones. Owing to the
configuration of the joint contact surfaces and the shape of
bones, purely mechanical factors cannot fulfill this require-
ment. This is why appropriate muscular forces, originating
from equilibration reactions, are necessary even in a nat-
ural upright posture, where the soleus is the emblematic
muscle.
Therefore, postural maintenance proceeds from an
‘‘active’’ process, insofar as it results from muscle con-
traction, in addition to inanimate ‘‘passive’’ forces, such
as gravity.
Voluntary movement is a perturbation to posture. Fur-
thermore, the body segments are rigid, and each one
is closely linked to the adjacent ones. Therefore, every
mechanical action exerted on any body point is transmit-
ted with no time lag to all the others. In particular, the
forces (and torques) developed during the movement are
transmitted from the segment(s) that the subject intention-
ally mobilizes to others, and from the distal one(s) to the
support surface(s), according to the ‘‘action and reaction
principle’’.
The initial conditions of movement are defined by body
posture, which is usually ‘‘static’’, such as standing, sitting,
crouching, etc. Therefore, the dynamic phenomena implied
by movement—–i.e. inertial forces (and torques)—–tend to
perturb balance.
Equilibrium state and dynamic stability
According to Martin [58,64], ‘‘posture denotes first the
shape of the body, i.e. the geometrical relationship of the
different parts to each other’’. In order to prevent any risk
of misunderstanding, ‘‘posture’’ would be limited to the
‘‘static’’ whole body configuration, i.e. to postural main-
tenance.
350
Figure 4 Center of gravity (CoG) and center of pressure (CoP)
displacements during upper limb elevation.
The subject performs a right shoulder flexion (indicated by the
arrow). Left: the subject is standing normally. Right: the subject
is standing on tiptoe, which limits CoP (solid line) and CoG (thick
dash line) displacements. In both cases, it is to observe that
the CoP is moving (from Io to I1) around the CoG, displacement
(from Go to G1). It can be noticed that the difference between
CoP and CoG is lower when the subject is standing on tiptoe
(right figure), which means that CoG horizontal acceleration is
lower.
Nevertheless, posture is first a physiological concept,
which can be defined in kinematic terms. Posture can also
be defined with respect to kinetics, i.e. to Newton’s laws. In
this context, postural maintenance corresponds to the con-
dition when the inertial resultant forces (and torques) are
nil (˙ Fe = 0 and ˙ M G (Fe ) = 0). On the contrary, they are
not nil during postural stabilization. However, the body is in
an equilibrium state during both postural maintenance and
postural stabilization.
The mechanical concept of stability is helpful to
accurately examine the influence of movement on body
equilibrium. According to mechanics, an equilibrium state
can be ‘‘stable’’ or ‘‘unstable’’. It is termed stable when
the structure, which is initially in equilibrium or stationary
state, returns to this state after a perturbation; for instance,
when a ‘‘static’’ equilibrium state is restored after a move-
ment. To this end, the sensory-motor system must develop
the forces necessary to oppose the perturbing forces: equi-
libration reactions must be put into play. Between the onset
and the end of a motor act, the structure is in a state of
transient disequilibrium, which is termed ‘‘dynamic equi-
librium’’.
‘‘Dynamic stability’’ is the process that allows dynamic
equilibrium at every instant, which means that ‘‘static’’
balance will be restored at the end of isolated move-
ments, or balance conditions will be achieved during
the next ‘‘stance’’ phase in periodic movements such as
walking. In other words, dynamic stability allows a task-
movement to be performed efficiently, from the onset to
the end. Dynamic stability is managed by the equilibra-
tion function, i.e. by ‘‘postural stabilization’’ as mentioned
above.
The stability limits of a system of articulated solids are
exceeded when the projection of the centre of gravity onto
the ground falls outside the support base perimeter, or if
the ground reaction force vector is not located within the
stability cone [60], which is equivalent. Within the stability
limits, a given posture is stable as long as the internal forces
S. Bouisset, M.-C. Do
are sufficient to oppose the perturbing ones. For a given set
of postural conditions, stability is a function of the intensity
of the perturbation.
General features of postural activity
associated with voluntary movement
Ever since Babinski’s observations [3], neurologists have
stressed that neurophysiologic mechanisms underlie the pos-
tural reactions associated with voluntary movement. For
instance, Babinski reported that when a standing subject
voluntarily executes a trunk extension, the lower limbs
move in the opposite direction. Several decades later,
electromyographic and biomechanical methods made it pos-
sible to deepen neurological data, allowing a broad survey
of muscular patterns and providing new data on motor
programming.
PAs associated to voluntary movement
Methodological considerations
Electromyography is unique in specifying muscle activation.
Indeed, surface EMG, i.e. primarily the electrical signals
picked up by surface electrodes, is a convenient index
of muscle excitation and allows a description of muscular
patterns. However, kinetic (or dynamic) quantities (which
refer to the force and energy associated with movement)
and kinematic ones (which refer to aspects of movement
besides the consideration of mass and force) must also be
considered. They are measured using different transducers
(goniometers, accelerometers, dynamometers, etc. . .) and
specific equipment, such as force plates and movement cap-
ture displays. Kinematic quantities (displacement, velocity
and acceleration) are used to characterize local movements,
i.e. segmental movements. Kinetic quantities (force, reac-
tion force and torque) are used to measure whole body
movement. In particular, force plates, which measure sup-
port reaction forces, allow to determine the resultant forces
and torques, as well as the centre of pressure. The resul-
tant force expresses the centre of gravity linear acceleration
to within the body mass, according to Eq. (1), and angular
acceleration to within the body moment of inertia in relation
to the centre of gravity, according to Eq. (2) (Fig. 3). Reac-
tion forces (and torques) at joint level can only be estimated
from a model (‘‘direct dynamics’’ or ‘‘inverse dynamics’’
modeling).
In addition, we must also keep in mind the conceptual
partitioning of the body into a ‘‘focal’’ and a ‘‘postural’’
chain when examining the influence of movement on pos-
ture. In particular, we must ask which body part executes the
focal movement, i.e. which one is the ‘‘primum movens seg-
ment’’ (PMS). Also, we must determine which muscle of the
focal synergy is the first to be activated, i.e. the ‘‘primum
movens muscle’’ (PMM), according to Beevor’s classification
[6].
Last, to be significant, the experimental data must be
linked to an adequate evaluation of the movement issue, in
terms of task performance (displacement, velocity, force,
precision, etc.). Indeed, EMG and biomechanical patterns,
as well as quantitative measurements (amplitude and tim-
ing) risk losing their true meaning if they are not related to
Posture, dynamic stability, and voluntary movement 351

the task constraints (specific parameters and environmental

context).

PAs and task-movement parameters

A variety of experimental situations have been studied.
Some are segmental voluntary movements, performed on
a single support base, such as shoulder flexion or exten-
sion [7,15,17,23,46,52,53,57,71], shoulder horizontal [2]
or lateral abduction [80], elbow flexion [8,35,36], push-
ing or pulling a handle [24,29,53, etc], or trunk bending
[26,67,70]. Others are still segmental movements, but
the support base is transitory, as in lower limb flex-
ions/extensions [32,66,72] or abductions [63]. Yet others
involve a whole body movement, which induces a transitory
support base transfer, such as rising up on tiptoe [48,55,65],
rocking on the heels [65], or whole body reaching [74] and
lifting [75] tasks. In these tasks, there is still no body pro-
gression even though the support base is transferred. Finally,
in motor tasks, such as locomotion, there is a periodic sup-
port base transfer, which corresponds to a succession of
balance losses and recoveries [21]. Finally, in a biomechan-
ical context, it can be considered that the experimental
situations differ according to two main conditions. First,
some are segmental movements, such as shoulder flexion
or extension, while others involve whole body movement,
such as locomotion or body reaching. Second, some are
performed on a single support base, while, in others, the
support base is transitory, such as rising up on tiptoe, or
transferred periodically, as in walking. In spite of these dif-
ferences, a survey of the literature [10,13] reveals general
features.
The primary result is that PAs associated with voluntary
movement proceed from an active process, insofar as they Figure 5 Electrical activity in the muscles of the scapular gir-
result from muscular contractions, in addition to the pas- dle during walking. Top insert: right upper limb positions during
sive joint forces, which are transferred from one segment the double step. It is interesting to note that the upper limb
to the other, according to the action and reaction law. More displacement is opposite the homonymous lower limb displace-
precisely, these studies allow us to identify several general ment. Bottom traces: the signals marked by the arrow indicate
features: the closure of contact when the heel touches the ground (right,
left and then right heel contacts). The EMGs are led off at on
the right side by means of small intramuscular electrodes (from
• Motor command is distributed to the body segments, [4]).
which are directly involved in the voluntary movement
(i.e. in the focal chain), as well as to others, which are not
are task specific: their features depend on the task
directly involved (i.e. to the postural chain). For exam-
parameters. They reveal dynamic joint fixations, or even
ple, the motor command is also addressed to lower limbs
displacements, i.e. ‘‘postural movements’’.
and trunk in upper limb movement, and to upper limbs
• PAs are adaptable, in that they are modified in relation to
and trunk in walking (Fig. 5). These are referred to as
functional states (fatigue and conditioning, growth and
PAs.
old age, microgravity, etc.). They are also modified, and
• PAs usually precede, accompany and follow the inten-
even suppressed, in impairments of the nervous, muscular
tional movement (Fig. 6): anticipatory postural adjust-
or bone and joint systems. Therefore, it has been claimed
ments (APAs) precede the voluntary movement, syn-
that PAs associated with voluntary movement are under
chronous postural adjustments (SPAs) accompany it, while
CNS control.
consecutive postural adjustments (CPAs) occur after its
end. APAs have been extensively studied, and will be
examined later on. Interest in SPAs has been mostly APAs
limited to ergonomic studies, following the studies of
Lundervold [56] and Floyd and Silver [34], and to loco- Many studies on APAs—–the PAs that precede the onset of
motion ones, following Elftman’s [33] pioneering work. voluntary movement—–have followed the pioneering studies
Last, CPAs seem to have been almost entirely neglected of Belenkii et al. [7] on shoulder flexion and of Herman et
since [17,50,25]. al. on stepping [45]. One of the advantages of the shoul-
• PAs are organized according to a well-defined sequence. der flexion paradigm is that it allows a simple distinction
Indeed, EMG as well as kinematic and kinetic patterns between the focal and the postural chain. The single sup-
352 S. Bouisset, M.-C. Do

Figure 6 Shoulder flexion kinetics time courses. Left inset: Schematic representation of the shoulder flexion paradigm. Right inset:
From left to right: OUF, unilateral shoulder flexions with no additional inertia; IUF, unilateral shoulder flexions with an additional
inertia; BF, bilateral shoulder flexion. From top to bottom: Rx, Ry and ARz, antero-posterior, lateral and vertical components of the
resultant force (Rz = Rz — P, with P being the weight of the subject; plus sign corresponds respectively to forward, right-to-left and
upward forces); Mz, resultant torque about the vertical axis (positive sign corresponds to a moment which tends to rotate the body
from the right side to the left); Awi, tangential acceleration of the upper limb recorded at wrist level; to and te correspond to the
onset and the end of the upper limb movement. Force platform data recorded on one subject performing upper limb movements,
under three experimental conditions. For each type of movement, records of five trials were superimposed by synchronizing them on
the onset of Awi. Dynamic phenomena can be observed before the onset (APAs) and after the end (CPAs) of the voluntary movement,
that is, before and after upper limb acceleration. Moreover, there is a morphological difference between the shape of the upper
limb acceleration time course and whole body kinetics (that is, center of gravity acceleration). If the body were rigid, a complete
and instantaneous transmission of the forces created by the voluntary movement would be observed. This hypothesis can be ruled
out since the time courses of the body’s biomechanical variables were different from that of the upper limb, which ascertains SPAs
(from [17]).

port base allows interpretation of the results without any
risk of misinterpretation related to unexpected support base
effects.
APA measurement
As it is well-known, the CNS triggers muscle excitation,
which in turn induces muscle activation, and hence mus-
cle contraction after a certain time (‘‘electro-mechanical
delay’’: EMD). Then, movement can occur and biomechani-
cal variables can be recorded.
Therefore, there are various possibilities for measuring
APAs. APAs can be defined with respect to EMGs, i.e. they are
measured by the time interval between postural and focal
muscle EMG onset. APAs can also be defined with respect
to biomechanical variables, i.e. the time interval between
global dynamics (or postural movement) onset and focal
movement onset. Of course, it is also possible to define APAs
by the time interval between the onset of a biomechani-
cal quantity and an EMG onset. This method raises various
problems, such as the location and parameter of the two
signals. Also, the influence of electro-mechanical delay on
the results, which differs according to the muscles and the
task parameters, cannot be disregarded.
Last, there is a technical problem, which cannot be
underestimated: how to estimate postural and focal onset.
Indeed, special care must be taken in determining EMG
onset and cessation. Also, it is less accurate to consider dis-
placements rather than accelerations, because of the slower
variation rate at the onset of movement.
APAs: movements which precede movement
In order to present APAs, it is easier to refer first to unilateral
shoulder flexions, performed with the upper limb stretched
out and on a single support base. Indeed, as it is a very simple
paradigm, one can suppose that the postural mechanisms,
which are involved, can guide the interpretation of more
complex experimental situations. Nevertheless, a compar-
ison with locomotion can be helpful, in that locomotion is
a lower limb movement, including a periodic support base
transfer.
A given sequence of EMGs was observed in the lower limbs
and trunk muscles before the onset of the Anterior Del-
toïdeus EMG, which is the PPM (Fig. 7). First, the ipsilateral
Soleus is deactivated 50 to 100 ms prior to PPM onset. Then
there is a sequence of phasic activations and deactivations,
starting with activation of the contralateral Tensor Fasciae
Latae and Rectus Femoris. This sequence is reproducible and
specific to the task under consideration [85].
A sequence of segmental accelerations corresponds to
the EMG sequence. Thus, before the onset of upper limb
acceleration, i.e. PMS onset, the lower limbs, pelvis, trunk
and shoulder display anticipatory accelerations (refer later
on to Fig. 9): the existence of accelerations indicates that
anticipatory activities are actually movements. More pre-
cisely, there is a given order in anticipatory accelerations,
starting with the contralateral lower limb and hip, the
ipsilateral ones, and ending with the ipsilateral shoulder:
it follows a bottom-up progression, as the postural seg-
ment accelerations proceed from the support surface. The
Posture, dynamic stability, and voluntary movement 353

Figure 7 EMG patterns in upper limb and lower limb movements. Top inset: electrodes location. Left inset: EMG activities of
lower limbs and pelvis recorded for shoulder flexion. From left to right: OUF, unilateral flexions with no additional inertia; IUF,
unilateral flexions with additional inertia; BF, bilateral flexions. From top to bottom: activities of anterior portion of Deltoideus
(DA), Erectores Spinae (ES), Gluteus Maximus (GM), Tensor Fasciae Latae (TFL), Semitendinosus (ST), Rectus Femoris (RF), Vastus
Lateralis (VL), Tibialis Anterior (TA), Soleus (SOL). Ipsilateral (i) and contralateral (c) activities with respect to the moving limb are
displayed in opposite directions. EMG activities are recorded for one subject performing the three types of movements. For each
type of movement, rectified and smooth EMG activities of five trials were superimposed, by synchronizing records on the onset of
DAi activities (dotted line). (From [94]) Right inset: EMG activities of lower limbs and pelvis recorded for gait initiation. From top
to bottom: activities of anterior portion of Soleus (SOL), Tibialis Anterior (TA), Rectus Femoris (RF), Semitendinosus (ST). Right (R)
and left (L) activities (gait is initiated with the right foot). TS: Soleus deactivation onset; tTA: Tibialis Anterior activation onset;
tE: ipsilateral Soleus burst onset; tRF: ipsilateral Rectus Femoris burst onset; tHO: onset of heel-off; tC: heel contact; tTO: onset
of the second step. Rectified and smooth EMG activities of five trials were superimposed by synchronizing recordings on the onset
of heel-off (tHO) (from [20]).

accelerometric sequence is also reproducible and specific
to the task under consideration. Since these results are
consistent with EMG ones, the EMGs could be related to
local accelerations: the local movements are not ‘‘passive’’
movements.
In gait initiation, the anticipatory EMG sequence (Fig. 7)
includes Soleus deactivation, followed by activation of both
Tibialis Anteriors 60 to 110 ms later. Then, the Soleus of
the forthcoming swing limb, which is the PPM of the heel-
off (HO), is activated. Thereafter, EMG bursts occur in the
Rectus Femoris of the swing limb and in the Semitendi-
nosus of the stance limb [20]. In addition, before heel-off
(HO) (i.e. PMS onset), the pelvis, trunk and shoulder display
anticipatory accelerations. The anticipatory sequence starts
at the stance leg, then at both hips and at the ipsilateral
shoulder, and afterwards at the contralateral shoulder [19].
Neck and head displacements in addition to trunk displace-
ments and their corresponding EMGs have also been reported
[81]. Finally, all the local accelerations indicate that the
movement is accelerated forwards and towards the stance
foot.
Here again, the sequence follows a bottom-up progres-
sion. But, as the postural segment accelerations proceed
from the support surface, it seems more appropriate to con-
sider that the progression order follows a ‘‘posturo-focal
gradient’’, i.e. proceeds from the support base (ground,
354 S. Bouisset, M.-C. Do

seat, etc... according to the initial posture) to the postural
chain, and then to the focal one.
Finally, APAs are postural movements. They precede vol-
untary movement. And like voluntary movement, they are a
dynamic phenomenon.
APAs and postural programming
According to Bernstein’s ideas [9], it is assumed that PAs
constitute a part of the motor program. In other words, it
is admitted that there is a ‘‘postural task’’ associated with
the focal one. APAs constitute a valuable tool in identifying
task parameters, which are programmed. Indeed, since by
definition, APAs precede the onset of focal movement, they
cannot result from re-afferentation triggered by the focal
movement: they are ‘‘pre-programmed’’. If APAs occur in
the postural chain, it can be said that the postural chain is
programmed. If APAs vary in relation to task parameters, it
can be assumed that the postural chain is programmed in
relation to these parameters.
Different specific parameters have been identified, such
as ‘‘velocity, load, direction’’, and orientation in space’’
(for a review, see [13]). For instance, in gait initiation, APAs
are graded as a function of movement peak velocity (Fig. 8).
A similar result has been found in shoulder flexion, and the
curvature of the relationship increases with additional iner-
tia [14]. Moreover, APAs depend on ‘‘postural conditions’’,
as will be discussed later on.
In addition, it must be stressed that APAs are not only
a function of the specific task-movement parameters. They
also depend on its location with respect to the body’s axes
of symmetry. Indeed, the role played by the homonymous
muscles and segments is inverted, when the movement
is executed by the contralateral limb (Fig. 9). This char-
acteristic is called ‘‘dynamic asymmetry’’ as opposed to
‘‘dynamic symmetry’’, corresponding to bilateral movement
[16]: in terms of posturo-kinetic programming, it is not
equivalent to raise both upper limbs simultaneously, or one
or the other upper limb. Therefore, since by definition,
APAs occur before the onset of voluntary movement, it can
be said that dynamic asymmetry and dynamic symmetry
are ‘‘pre-programmed’’. The organization of the postural
sequence according to the forthcoming focal limb(s) is a gen-
eral feature. For example, gait initiation displays a mirror
organization according to the stepping leg, i.e. a dynamic
asymmetry pattern, whereas it is a dynamic symmetry pat-
tern in standing jump initiation.
Finally, APAs are dynamic, polarized, and task specific.
They are programmed in relation not only to the focal
movement parameters per se, but more generally to task-
movement parameters. In addition, they are ‘‘adaptable’’,
as they depend on the functional state of the motor system,
which will be discussed later on.
Dual role of APAs
APAs as a counter-perturbation. The role of APAs may
be proposed on the basis of an analysis of forces acting
at the shoulder level and at the body’s centre of gravity
(Fig. 10), whose directions are opposite. From this analysis,
it can be assumed that ‘‘APAs tend to create inertial forces,
which, when the time comes, will counterbalance the dis-
turbance to postural equilibrium due to the intentional
forthcoming movement’’ [15]. According to this hypothesis,
the intentional movement is considered a ‘‘perturbation’’,
in accordance with the ideas put forward by several neu-

Figure 8 Gait initiation. Left part: Biomechanical quantities time course. From top to bottom: XG: forward (F) antero-posterior
CoG displacement (dotted line) and Xp: backward (B) antero-posterior CoP displacement (solid line) with ax being the peak
amplitude; Yp and YG: CoP and CoG lateral displacements to the left (L) and to the right (R); ŸG : right (R) and left (L) CoG
lateral acceleration; X G : forward (F) and backward (B) CoG antero-posterior acceleration; X’G : CoG antero-posterior velocity. to:
onset of kinetic phenomena; tHO; heel off; tV: peak velocity progression (V) instant. The movement is executed at spontaneous
speed with the left foot. It can be observed that CoP and CoG displacements precede tHO. In particular, the CoP first moves backward
and toward the forthcoming stepping foot, while the CoG moves in the opposite direction. More precisely, there is a rapid unloading
of the forthcoming swing limb with an equivalent loading of the stance limb. (From 21). Right part: relationship between APA indices
and progression velocity. APA peak amplitude (ax) and duration (tHO) are plotted against the peak of progression velocity (V) (from
[21]).
Posture, dynamic stability, and voluntary movement 355

Figure 9 Dynamic asymmetry and symmetry. Left part: Local accelerations in shoulder flexions. From left to right: OUF, unilateral
flexions with no additional inertia; IUF, unilateral flexions with additional inertia; BF, bilateral flexions. From top to bottom: Awi,
tangential acceleration of the upper limb measured at wrist level (plus sign corresponds to the acceleration phase of the movement);
Ash, Atr, Ah, At and As, antero-posterior accelerations measured at level of shoulders, trunk, hips, thighs and shanks (plus sign
corresponds to forward acceleration); i and c, ipsilateral and contralateral accelerations with respect to the moving limb. Antero-
posterior local accelerations are reported from one subject performing the three types of movement. For each type of movement,
accelerations of five trials were superimposed, by synchronizing records on the onset of Awi (dotted line). (From 94). Right part:
direction of anticipatory local accelerations. From left to right: unilateral flexions performed by the right upper limb (UFR), the
left upper limb (UFL) and both upper limbs simultaneously (BF). Notice that APA direction is inverted when the forthcoming moving
upper limb is the left instead of the right (from [84]).

rologists (see, for instance [1]), which were first revisited
by Belenkii et al. [7]. In other words, APA represents a
‘‘counter-perturbation’’, which would be responsible for
the stabilizing action.
In this context, it was proposed that one consider that the
postural perturbation, associated with voluntary movement,
as well as the counter-perturbation, might be characterized
by two biomechanical factors [16]. They correspond to the
two vectors to which the perturbation force system may be
reduced, i.e. its resultant and its torque (in reference to the
body’s centre of gravity, for example). One is linear, while
the other is rotational. Both may include three orthogonal
components.
However that may be, as APAs are triggered by a feed-
forward command and are specific to the characteristics of
the motor task, they have to be determined from previous
knowledge of its perturbing effect [84]. More precisely, APAs
are programmed according to the expected perturbation and
not to the actual one [77].
APA as a perturbation. The contention according to which
APAs represent a counter-perturbation has been argued
within the framework of segmental movements performed
on a continuous support base. But, the support base can also
change transiently, as in lower limb movements, or peri-
odically, as in locomotion. These movements will provoke
a perturbation of the whole kinematic chain, but the cor-
responding APAs have been shown to play a more complex
role.
In lower limb movements involving a transient change
of the support base perimeter, such as lower limb flexion,
APAs have been shown to be related not only to the counter-
perturbation of the forthcoming movement, but also to
‘‘body weight transfer’’ onto the forthcoming stance foot
[32]. This transfer can be interpreted as being related to
postural stabilization, as well as to the necessary unloading
of the forthcoming moving foot.
In locomotion, there is a succession of balance losses and
recoveries, which corresponds to the periodic support base
transfer. It has been demonstrated that during gait initia-
tion, the greater the forward fall of the centre of gravity,
the faster the progression velocity [21]: APAs induce postural
destabilization, which is necessary to initiate gait. In addi-
tion, the initiation phase also includes body weight transfer
towards the forthcoming stance foot, which can be inter-
preted as being related to postural stabilization.
APAs as a perturbation and a counter-perturbation.
Finally, it cannot be excluded that APAs play a dual role
simultaneously, i.e. to perturb balance and to counteract
the perturbation, as suggested by gait initiation: acceler-
ations occur at the ipsilateral hip at heel-off, the forward
direction of which is opposite to the hip perturbation at this
instant [30].
Indeed, just as in shoulder flexion initiation, lower-limb
forward displacement at heel-off will induce a perturbation
at the hip joint, which is directed backwards. It must be
compensated in order to prevent the hip from being pushed
356
Figure 10 Interpretation of the finality of anticipatory pos-
tural adjustments. The filled arrows correspond to the actual
recorded biomechanical data; the broken arrows correspond
to theoretical parameters. , angular displacement of the
upper limb(s); Aw,  r , and  tangential, radial and total upper
limb acceleration. Rx and Rz , antero-posterior and vertical
acceleration of the body’s center of gravity, G. Mz resul-
tant moment about the vertical axis crossing G. BF: bilateral
(shoulder) flexion; UF: unilateral (shoulder) flexion. It can be
observed that the perturbation (indicated by broken arrows at
the shoulder level), and the corresponding APA (indicated by
full arrows at the center of gravity) display opposite directions
(from [17]).
backwards at the very instant when it has to serve as a
fulcrum for hip rotation.
More generally, trunk and shoulder movements during
walking, which induce oscillation in the upper limbs, play
a major role in regulating body rotation with respect to the
vertical axis [33]. They diminish whole-body rotation dur-
ing the single support and favor a more gradual change in
the rotation direction during double support. In addition,
body rotation with reference to the antero-posterior axis
is modified by upper-limb oscillation, such that direction
changes are more progressive. Consequently, these upper-
body movements play a role in postural stabilization, as was
claimed, after Elftman [33], by authors such as Cappozzo
[22], Cromwell et al. [27], Jian et al. [50] and Winter [82].
Finally, APAs appear to play a dual role, that is, postural
stabilization and the creation of propulsive forces. One or
S. Bouisset, M.-C. Do
the other prevails depending on the task-movement param-
eters. For a given task-movement, it cannot be excluded
that the global role is different from the one played at the
local level.
APAs as an optimal control process
Belenkii et al. [7] have suggested that APAs have to com-
ply with the constraint of ‘‘minimizing the perturbation of
the whole kinematic chain’’. However, the APA counter-
perturbing role can recover from various constraints: is it
to fix the body centre of gravity or the joint(s) around which
the focal limb is moved, or—–for limb movements at least—–
the reference axes needed to calibrate the motor program,
such as the trunk [41] or the head [69]. Even though the
problem is obviously complex and still under discussion, the
first hypothesis seems to have been largely agreed upon for
the past two decades (see the reviews [10,59]). But the
other two possibilities, which are not mutually exclusive,
still need to be argued.
According to Brenière et al. [21] and Lepers and Brenière
[54], the APA perturbing role in gait initiation corresponds
to an economical process, insofar as the propulsive forces
at heel-off originate primarily from gravity forces, which
results in muscular energy savings. But the APA role can also
include other constraints, such as postural stabilization or
the reference axes required to calibrate the motor program
[68].
Therefore, to claim that, as a part of the entire motor
act, APAs result from an optimal control process is not eas-
ily demonstrated, insofar as a critical question must first
be answered, i.e. how many and what criteria have to
be minimized: mechanical energy, metabolic energy, neu-
ral inputs, etc. Obviously, partial responses can only be
provided, although it is tempting to consider that the APA
sequence complies with an optimal control process, whose
criteria could vary according to the task-movement param-
eters and the subject’s functional state.
Posturo-kinetic capacity: a limiting factor of
performance
Posturo-kinetic and foco-kinetic capacities
APAs have been presented as the optimal biomechanical
means to initiate voluntary movement and to approach
postural programming. In order to broaden the approach
to motor programming, it is tempting to return to the
hypothesis according to which motor control includes a
‘‘focal’’ and a ‘‘postural’’ component [38] and to con-
sider a ‘‘postural system’’ and a ‘‘focal system’’. To one
would correspond a ‘‘posturo-kinetic capacity’’ (PKC), as
proposed by Bouisset and Zattara [16], and to the other,
a ‘‘foco-kinetic capacity’’ (FKC) (Fig. 11). PKC means the
ability to develop a counter-perturbation to the posture per-
turbation induced by voluntary movement, and therefore
the ability to manage the perturbation to balance asso-
ciated with the forthcoming movement. FKC means the
ability to execute the voluntary movement per se, i.e. to
mobilize the focal segments. PKC is believed to depend
on the configuration of the postural chain, the environ-
mental context and the functional state of the postural
system.
Posture, dynamic stability, and voluntary movement
Figure 11 Posturo-kinetic capacity (PKC) and foco-kinetic
capacity (FKC). The motor command (‘‘command’’) is assumed
to result from the intent of achieving a given task, which can be
characterized by three factors: category, specific parameters,
and environmental context (refer to Fig. 1). ‘‘Performance’’
characterizes the task that is actually performed as a result of
the movement. Under the hypothesis according to which motor
command includes a ‘‘focal’’ and a ‘‘postural’’ component, it
is supposed that there is a ‘‘postural system’’ and a ‘‘focal
system’’. To one would correspond a ‘‘posturo-kinetic capac-
ity’’ (PKC), and to the other, a ‘‘foco-kinetic capacity’’ (FKC),
both of which must be seen as systemic entities. PKC means the
ability to develop a counter-perturbation to the posture per-
turbation induced by voluntary movement, and therefore the
ability to manage the perturbation to balance associated with
the forthcoming movement. FKC means the ability to execute
the voluntary movement per se, that is, to mobilize the focal
segments. PKC is assumed to depend on the configuration of the
postural chain, the environmental context and the functional
state of the postural system. FKC is assumed to depend on the
configuration of the focal chain, the environmental context and
the functional state of the focal system. It is assumed that per-
formance is the end product of the cooperation between PKC
and FKC.
It is assumed that performance is the end product of
the cooperation between PKC and FKC. In other words,
PKC would control task performance in preparing the motor
action, supporting its course and contributing to its effi-
ciency. In this view, PKC would operate throughout the entire
motor act.
Finally, both PKC and FKC must be seen as systemic enti-
ties: their properties proceed from a combination of their
functional components, mainly the muscular, bone and joint,
and sensory-motor systems. Both PKC and FKC are assumed
to be ‘‘dynamic processes’’, which vary at each instant in
relation to the current conditions.
PKC assessment
It is clear that PKC assessment allows approaching the pos-
tural strain related to a given task. The problem has already
been discussed in detail [12,13]. To summarize, the method
for assessing PKC involves evaluating the postural input to
the motor system, and the corresponding output, when max-
imal performance is required. The simplest procedure is
to relate performance (i.e. the maximal movement veloc-
ity, or the maximal muscular force, etc. depending on the
357
paradigm) to the corresponding APA (i.e. to their duration,
amplitude, or even area). However, it does not necessarily
imply that the two quantities have to be expressed in the
same unit. Finally, the ratio of performance to APA defines
PKC, and the inverse ratio, a ‘‘postural cost’’. For example,
if the maximal velocity (m/s) is lower than in the controls
and the APA lasts longer (s), it can be concluded that PKC is
reduced and postural cost increased.
This reasoning is indubitably supported by the assump-
tion that anticipatory, contemporary, and consecutive PAs
are parts of the same motor program. In other words, it is
assumed that, since the movement process is continuous,
the postural process is also continuous and part of it, such
that the APA is representative of the entire process. This
assumption is consistent with theories on motor planning.
Another possibility for assessing PKC is to consider the
biomechanical or EMG pattern. If it is assumed that the
primary synergy results from an optimization criterion, a dif-
ferent synergy can be assumed to indicate a less economical
process: performance would be achieved at the expense of
increased postural cost. EMG or biomechanical profiles have
to be examined and their main features (amplitude, dura-
tion, time to peaks, etc.) measured in order to characterize
the postural pattern. But the method has also proved to be
useful even though it is qualitative, as will be exemplified
in the case of impairments.
Finally, the influence of a given factor on PKC, such as
postural stability or impairment, is assessed by comparing
the corresponding data to the control, for example, the data
obtained for comfortable stance or able-bodied subjects.
To conclude, assessing PKC means evaluating the postu-
ral cost of a task. The method involves relating performance
to APA and considering muscular and/or biomechanical pat-
terns.
PKC and postural configuration
In the preceding sections, voluntary movement was said
to be a perturbation to body balance, and that postural
counter-perturbation must be adequate for the intended
task to be achieved efficiently. Moreover, voluntary move-
ment is usually superimposed on an initial ‘‘static’’ posture
and is followed by a final ‘‘static’’ posture as well.
According to the literature (for a review, see [13]), APAs
depend on postural conditions, i.e. mainly on the support
base area and stability. However, when the initial and final
postures are less stable [37,47], some authors find that APA
duration increases while others find it decreases. These dif-
ferences are not easy to interpret, in that the experimental
conditions are so diverse that comparisons are difficult.
Moreover, performance is seldom reported. However, two
sets of experiments help delve deeper into the influence of
the support base area on PKC.
In the first one, the voluntary movement (a pointing
task) was performed by standing subjects. The support base
configuration differed by the size of the antero-posterior
distance between the heels [83]. When the distance was
increased from 0 to 40 cm, i.e. when postural stability was
enhanced, maximal velocity increased, while the APA dura-
tion remained equivalent. Thus, the PKC was greater when
the body support size was enlarged (PKC = 0.5, as compared
to PKC = 0.2) and the postural cost was lower. Hence, postu-
ral cost is a function of postural stability.
358 S. Bouisset, M.-C. Do

In the second experiment, the voluntary movement
(a flexion or a flexion-extension of the lower limb) was
performed while various initial and final postures were
considered [32,66]. The general trend was an increase in
APA duration and a decrease in maximal velocity when
the initial and final postures were less stable: PKC was
reduced in relation to postural stability. Further, APAs were
absent when the subjects stood on one foot, from the
beginning of the lower limb movement to the end [66]:
as APAs induce dynamics, they can potentially perturb
body balance, and are no longer present when the pos-
ture is excessively unstable. Therefore, in conditions of
high stability demand, the CNS may reduce and even sup-
press APAs, as protection against their possible destabilizing
effects. Then, the focal and the postural chains are acti-
vated simultaneously, at the expense of greater forces to
be developed during the movement itself. In other words,
the lack of APAs suggests that postural stabilization is
a major constraint instead of muscular energy minimiza-
tion.
PKC and the functional state of the motor system
As stated above, performance results from a hierarchi-
cal process, associating three different components, i.e.
the central nervous, muscular, and bone and joint systems
(Fig. 1). More, or less, efficient performance depends on
their functional state, which is not easy to assess, either
clinically or scientifically. Nevertheless, it is agreed that it
varies according to impairment or rehabilitation, condition-
ing or fatigue, development or ageing [51].

Figure 12 EMG pattern in paraplegic (Pa), as compared to able-bodied (Va) subjects, performing a pick-and-place task. Top
inset Schematic representation of the experimental situation: Biomechanical traces of able-bodied and paraplegic subjects. Rz*:
vertical component of the normalized resultant of forces (vertical acceleration of the center of gravity); plus sign (+) indicates
upward acceleration; b1: first peak amplitude; tz: onset time. Rx*: horizontal component of the normalized resultant of forces
(horizontal acceleration of the center of gravity); plus sign (+) indicates forward acceleration; a1 and a2: first and second peak
amplitudes; tx: onset time; d1 and d2: duration of the first and second phase of voluntary movement. to and te: onset and end of load
transport. Notice that the load transport duration is greater for Pa than for Va. Superimposition of five records (from [31]). Bottom
inset: Location of bipolar surface electrodes for recording EMG activities. From top to bottom: EMG activities of Latissimus Dorsi
(LD), Serratus Anterior (SA), Trapezius Medialis (TM), Trapezius Superior (TS), Erectores Spinae (ES), which cannot be activated by
Pa patient; subscript R and L indicate right and left respectively. Rx* is the antero-posterior component of the normalized resultant
of forces. to and te are the onset and the end of the forward load transport. Superimposition of five records of rectified EMGs.
Posture, dynamic stability, and voluntary movement 359

For instance, it has been reported that APAs last longer in
the elderly [46,49,57,75] and in hemiplegics [46], whereas
maximal movement velocity is reduced. Also, APAs van-
ish when people are bedridden for several weeks [40],
and in Parkinsonians [5,29,18]. Moreover, postural patterns
are changed in Parkinsonians, hemiplegics, etc. Therefore,
sensory-motor impairments diminish the functional state,
i.e. ‘‘sensory-motor capacity’’, which is the resultant of PKC
and FKC.
In other words, PKC assessment in sensory-motor impair-
ments requires consideration of those impairments, which
result in a restriction of joint mobility, regardless of the
exact origin (bones, joints, tendons, muscles or nervous sys-
tem). It is particularly interesting to discuss paraplegics and
lower-limb amputees.
Paraplegia and PKC. In paraplegics performing a pick-
and-place task, it was established that maximal movement
velocity was slower than in the able-bodied controls, but
that APAs lasted much longer [31]. Thus, PKC is less in para-
plegics: PKC = 3.4, as compared to PKC = 9 for able-bodied
subjects.
Postural synergy is also different: other scapular girdle
and trunk muscles are activated, especially on the contralat-
eral side (Fig. 12). If it is assumed that the primary synergy
corresponds to an optimization criterion, the transition from
one synergy to another can be assumed to correspond to a
less economical process.
In other words, paraplegics, whose equilibrium is more
unstable than able-bodied subjects, could not develop PAs
adapted to the perturbation provoked by the motor task
without an excessive balance risk: the perturbation, and
hence the required performance, should be reduced. There-
fore, due to PKC reduction, voluntary movement is less
efficient.

Figure 13 Gait initiation by above-knee amputees. Top: biomechanical quantities time course. From left to right: above-knee
amputee subject initiating gait with the prosthetic limb (A) and with the sound limb (B), and able-bodied subject (C). Ground
reaction forces were recorded from two force platforms. to: onset of biomechanical quantities variation; FO: time of foot-off;
APA: anticipatory postural adjustment phase; EXE: swing phase; tVm: time of maximum velocity, Fx: antero-posterior ground reac-
tion force recorded on the side of the stepping limb (dashed line) and on the side of the stance limb (solid line); ˙Fx: sum of
antero-posterior reaction forces. f: forward; b: backward. (From 68). Bottom: PKC cost in above-knee and below-knee amputees
and controls. From left to right: Amputation (Amp.) level: above-knee (AK), below-knee (BK); performance: maximum speed of
progression; APA duration; PKC: ratio between speed and APA duration. It can be noted that PKC depends on the level of amputa-
tion. When the prosthetic limb is the stance limb, PKC is lower in the AK amputee, and PKC is higher in the lower amputation (BK
amputee). Moreover, when the stance limb is the sound limb, PKC is comparable between AK and BK amputees, but still lower than
controls. This gap suggests that other factors could contribute to the PKC (from [61] results).
360
Lower limb amputation and PKC. Similarly, unilateral
lower limb amputees showed many kinetic changes in gait
initiation [61,62,73,76,78,79]. Although maximal progres-
sion velocity is similar, regardless of the stepping side
(prosthetic or sound limb), it is slower that in controls. Fur-
thermore, the muscular pattern was necessarily different.
PKC can be discussed with reference to the forthcoming
stance leg [61] (Fig. 13). When above-knee (AK) amputees
initiate gait with the sound limb (the prosthetic leg is the
forthcoming stance leg), APAs are longer and, since the pro-
gression velocity is the same, the PKC is lower (PKC = 1.5).
When they initiate gait with the prosthetic limb (the sound
leg is the forthcoming stance leg), APAs are shorter while
the progression velocity remained the same: PKC is higher
(PKC = 2.7). In controls, PKC is approximately the same
as in amputees when the stance limb is the sound one
(PKC = 2.8).
In addition, PKC is less affected by below-knee ampu-
tation than by above-knee amputation. Indeed, when
below-knee (BK) amputees initiate gait with the sound limb,
PKC = 2.5, and when it is with the prosthetic limb, PKC = 2.1.
It is interesting to note that PKC values are of the same order
in below-knee amputation and in controls when the stance
limb is the sound limb. Also, PKC is drastically reduced in
above-knee amputation as compared to below-knee ampu-
tation when the stance limb was the prosthetic limb.
Finally, the results, obtained in both paraplegics and
amputees, could be interpreted in terms of PKC. The same
hypothesis also fits other results, such as those obtained
in hemiplegics [46] and the elderly [75], where maximal
movement velocity was reduced but APAs were longer. It
also proves useful in the study of low back pain subjects
[42].
Therefore, the PKC concept probably has various appli-
cations, in the clinical field at least. Its potential deserves
further investigation.
References
[1] André-Thomas. Équilibre et équilibration. Masson; 1940.
[2] Aruin AS, Latash ML. Directional specificity of postural muscles
in feed-forward postural reactions during fast voluntary arm
movements. Exp Brain Res 1995;103:323—32.
[3] Babinski J. De l’asynergie cérebelleuse. Rev Neurol 1899;7:
806—16.
[4] Ballesteros ML, Buchthal F, Rosenfalck P. The pattern of mus-
cular activity during the arm swing of natural walking. Acta
Physiol Scand 1965;63:296—310.
[5] Bazalgette D, Zattara M, Bathien N, Bouisset S, Rondot P.
Postural adjustments associated with rapid voluntary arm
movements in patients with Parkinson’s disease. In: Yarr PD,
Bergman KJ, editors. Advances in neurology, 45. Raven Press;
1986. p. 371—4.
[6] Beevor CE. Croonian lectures (for 1903) on Muscular Move-
ments. London: Adlar; 1904.
[7] Belenkii YY, Gurfinkel VS, Paltsev YI. Element of control of
voluntary movements. Biofizika 1967;12:135—41.
[8] Benvenuti F, Panzer V, Thomas S, Hallett M. Kinetimatic
and EMG analysis of postural adjustments associated with
fast elbow flexion movements. In: Brandt T, Paulus W,
Bles W, Dieterich M, Krafezyk S, Straube A, editors.
Disorders of posture and gait. G Thieme Verlag; 1990.
p. 72—5.
S. Bouisset, M.-C. Do
[9] Bernstein N. Coordination and regulation of movements. Perg-
amon Press; 1935 (Amer. translation, 1967).
[10] Bouisset S. Relation entre support postural et mouvement
intentionnel: approche biomécanique. Arch Int Physiol Bioch
Biophys 1991;99:A77—92.
[11] Bouisset S. Biomécanique et physiologie du mouvement. Paris:
Masson; 2002.
[12] Bouisset S, Do MC, Zattara M. Posturo-kinetic capacity assessed
in paraplegics and parkinsonians. In: Woolacott M, Horak F, edi-
tors. Posture and Gait; control mechanisms, vol II. University
of Oregon Books; 1992. p. 19—22.
[13] Bouisset S, Le Bozec S. Posturo-kinetic capacity and pos-
tural function in voluntary movements. In Latash ML, ed:
Progress in Motor Control, vol II: Structure-Function Relations
in Voluntary Movements. Human Kinetics. 2002, Chapter 3:
p. 25—52.
[14] Bouisset S, Richardson J, Zattara M. Do anticipatory postural
adjustments occurring in different segments of the postural
chain follow the same organisational rule for different task-
movement velocities, independently of the inertial load value?
Exp Brain Res 2000;132:79—86.
[15] Bouisset S, Zattara M. A sequence of postural movements pre-
cedes voluntary movement. Neurosci Lett 1981;22:263—70.
[16] Bouisset S, Zattara M. Anticipatory postural movements related
to a voluntary movement. In Space Physiology, Cepadues Pubs.
1983; p. 137—41.
[17] Bouisset S, Zattara M. Biomechanical study of the programming
of anticipatory postural adjustments associated with voluntary
movement. J Biomech 1987;20:735—42.
[18] Bouisset S, Zattara M. Segmental movement as a perturba-
tion to balance? Facts and concepts. In: Winters JM, Woo SLY,
editors. Multiple muscle system, biomechanics and movement
organization. Springer Verlag; 1990. p. 498—506.
[19] Brenière Y, Dietrich G. Heel-Off perturbation during gait initi-
ation: biomechanical analysis using triaxial accelerometry and
a force plate. J Biomech 1992;25(2):121—7.
[20] Brenière Y, Do MC. Modifications posturales associées au lever
du talon dans l’initiation du pas de la marche normale. J Bio-
phys Biomech 1987;11(4):161—7.
[21] Brenière Y, Do MC, Bouisset S. Are dynamic phenomena prior
to stepping essential to walking? J Motor Behav 1987;19(1):
62—76.
[22] Cappozzo A. Analysis of the linear displacement of the head
and trunk during walking at different speeds. J Biomech
1981;14(6):411—25.
[23] Clément G, Gurfinkel VS, Lestienne F, Lipshits MI, Popov KE.
Adaptation of postural control of weightlessness. Exp Brain Res
1984;57:61—72.
[24] Cordo PJ, Nashner LM. Properties of postural movements
related to a voluntary movement. J Neurophysiol 1982;47:
287—303.
[25] Crenna P, Do MC, Brenière Y. Motor programmes for the termi-
nation of gait in humans: organisation and velocity-dependent
adaptation. J Physiol Lond 2001;537:1059—72.
[26] Crenna P, Frigo C, Massion J, Pedotti A. Forward and backward
axial synergies in man. Exp Brain Res 1987;65:538—48.
[27] Cromwell RL, Newton RA, Forrest G. Head stability in older
adults during walking with and without visual input. J Vestib
Res 2001;11(2):105—14.
[28] Dempster WT. The anthropometry of body action. Annals NY
Academy of Sciences 1955;63:559—85.
[29] Dick JPR, Rothwell JC, Berardelli A, Thompson PD, Gioux
M, Benecke R, et al. Associated postural adjustments in
Parkinson’s disease. J Neurol Neurosurg Psychiatry 1986;49:
1378—85.
[30] Dietrich G, Brenière Y, Do MC. Local expressions of anticipatory
movements in single step initiation. Hum Mov Stud 1994;13:
195—210.
Posture, dynamic stability, and voluntary movement
[31] Do MC, Bouisset S, Moynot Cl. Are Paraplegics handicapped
in the execution of a manual task? Ergonomics 1985;28(9):
1363—75.
[32] Do MC, Nouillot Ph, Bouisset S. Is balance or posture at the
end of a voluntary movement programmed? Neurosci Lett
1991;130:9—11.
[33] Elftman H. The function of arms in walking. Hum Biol 1939;
11(4):529—35.
[34] Floyd WF, Silver PHS. The function of Erectores Spinae mus-
cles in certain movements and postures in man. J Physiol Lond
1955;129(1):184—203.
[35] Friedli WG, Hallett M, Simon SR. Postural adjustments
associated with rapid voluntary arm movements 1. Elec-
tromyographic data. J Neurol Neurosurg Psychiatry 1984;47(6):
611—22.
[36] Friedli WG, Cohen L, Hallett M, Stanches S, Simon SR. Postural
adjustments associated with rapid voluntary arm movements
II. Biomechanical analysis. J Neurol Neurosurg Psychiatry
1988;51:232—43.
[37] Gantchev GN, Dimitrova DM. Anticipatory postural adjust-
ments associated with arm movements during balancing
on unstable support surface. Int J Psychophysiol 1996;22:
117—22.
[38] Gelfand IM, Gurfinkel VS, Tsetlin ML, Shik ML. Problems in anal-
ysis of movements. In: Gelfand IM, Gurfinkel VS, Fomin SV,
Tsetlin ML, editors. Models of the structural functional organi-
zation of certain biological systems (Amer. translation, 1971).
Cambridge: MIT Press; 1966. p. 330—45.
[39] Gray J. Animal locomotion. Weidenfeld and Nicolson; 1968.
[40] Gurfinkel VS, Elner AM. On two types of static disturbances
in patients with local lesions of the brain. Agressologie
1973;14D:65—72.
[41] Gurfinkel VS, Lipshits MI, Mori S, Popov KE. Stabilization of body
position as the main task of postural regulation. Fiziologiya
Cheloveka 1981;7:400—10.
[42] Hamaoui A, Do MC, Bouisset S. Postural sway increase in low
back pain subjects is not related to reduced spine range of
motion. Neurosci Lett 2004;357:135—8.
[43] Hamaoui A, Do MC, Poupard L, Bouisset S. Does respiration per-
turb body balance more in chronic low back pain subjects than
in healthy subjects? Clin Biomech 2002;17:548—50.
[44] Hess WR. Teleokinetisches und ereismatisches Kräftesystem in
der. Biomotorik Helv Physiol Pharmacol Acta 1943;1:C62—3.
[45] Herman R, Cook T, Cozzens B, Freedman W. Control of postural
reactions in man: the initiation of gait. In: Stein RB, Pearson
KG, Redford JB, editors. Control of posture and locomotion.
New York: Plenum Press; 1973. p. 363—88.
[46] Horak FB, Esselman PE, Anderson ME, Lynch MK. The effect
of movement velocity, mass displaced and task certainty on
associated postural adjustments made by normal and hemi-
plegic individuals. J Neurol Neurosurg Psychiatry 1984;47:
1020—8.
[47] Horak FB, Nashner LM. Central programming of postural move-
ments: adaptation to altered support-surface configuration. J
Neurophysiol 1986;55(6):1369—81.
[48] Houtz SJ, Fisher FJ. Function of leg muscles acting on foot
as modified by body movements. J Appl Physiol 1961;16(4):
597—605.
[49] Inglin B, Woollacott M. Age-related changes in anticipa-
tory postural adjustments associated with arm movements. J
Gerontology 1988;43:M105—13.
[50] Jian Y, Winter DA, Ishac MG, Gilchrist L. Trajectory of the body
COG and COP during initiation and termination of gait. Gait
Posture 1993;1:9—22.
[51] Latash ML, Anson JG. What are « normal movements » in atyp-
ical populations? Behav Brain Sci 1996;19:55—106.
[52] Lee WA. Anticipatory control of posture and task muscles during
rapid arm flexion. J Mot Behav 1980;12:185—96.
361
[53] Lee WA, Buchanan TS, Rogers MW. Effects of arm acceleration
and behavioural conditions on the organization of postu-
ral adjustments during arm flexion. Exp Brain Res 1987;66:
257—70.
[54] Lepers R, Brenière Y. The role of anticipatory postural adjust-
ments and gravity in gait initiation. Exp Brain Res 1995;107:
118—24.
[55] Lipshits M, Mauritz K, Popov KE. Quantitative analysis of antici-
patory postural components of a complex voluntary movement.
Hum Physiol 1981;7:411—9.
[56] Lundervold A. Electromyograhic investigations of position and
manner of working in typewriting. Acta Physiol Scand suppl
1951;84:1—171.
[57] Maki BE. Biomechanical approach to quantifying anticipatory
postural adjustments in the elderly. Med Biol Eng Comput
1993;31:355—62.
[58] Martin JP. The Basal Ganglia and Posture. Pitman; 1967.
[59] Massion J. Movement, posture and equilibrium: Interaction and
coordination. Prog Neurobiol 1992;35:35—56.
[60] McCollum G, Leen TK. Form and exploration of mechanical
stability limits in erect stance. J Mot Behav 1989;21:225—44.
[61] Michel V, Do MC. Are stance ankle plantar flexor muscles neces-
sary to generate propulsive force during human gait initiation?
Neurosci Lett 2002;325(2):139—43.
[62] Michel V, Do MC*, Chong RKY. The strategies to regulate and to
modulate the propulsive forces during gait initiation in lower
limb amputees. Exp Brain Res 2004;158(3):356—65, *erratum
on http://dx.doi.org/10.1007/s00221-004-2001-3.
[63] Mouchnino L, Aurenty R, Massion J, Pedotti A. Stratégies
de contrôle simultané de l’équilibre et de la position de la
tête pendant l’élévation d’une jambe. CR Acad Sci, Paris
1991;312:225—32.
[64] Murray MP, Seireg A, Scholz RC. Center of gravity, center of
pressure and supportive forces during human activities. J Appl
Physiol 1967;23:831—8.
[65] Nardone A, Schieppati M. Postural adjustments associated with
voluntary contractions of leg muscles in standing man. Exp
Brain Res 1988;69:469—80.
[66] Nouillot P, Bouisset S, Do MC. Do fast voluntary movements
necessitate anticipatory postural adjustments even if equilib-
rium is unstable? Neurosci Lett 1992;147:1—4.
[67] Oddsson L, Thorstensson A. Fast voluntary trunk flexion
movements in standing: primary movements and associ-
ated postural adjustments. Acta Physiol Scand 1986;128(3):
341—9.
[68] Paillard J. Brain and space. Oxford University Press; 1991.
[69] Pozzo T, Berthoz A, Lefort L. Head stabilization during various
locomotor tasks in humans. I. Normal subjects. Exp Brain Res
1990;82:97—106.
[70] Pedotti A, Crenna P, Deat A, Frigo C, Massion J. Postural syner-
gies in axial movements: short and long-term adaptation. Exp
Brain Res 1989;74:3—10.
[71] Riach CL, Hayes KC. Anticipatory postural control in children.
J Mot Behav 1990;22:250—6.
[72] Rogers MW, Pai YC. Dynamic transitions in stance support
accompanying leg flexion movements in man. Exp Brain Res
1990;81:398—402.
[73] Rossi SA, Doyle W, Skinner HB. Gait initiation of persons with
below-knee amputation: the characterization and comparison
of force profiles. J Rehabil Res Dev 1995;32(2):120—7.
[74] Stapley P, Pozzo T, Grishin A. The role of anticipatory postural
adjustments during whole body reaching movements. Neurore-
port 1998;9:395—401.
[75] Stelmach GE, Populin L, Friedemann M. Postural muscle
onset and voluntary movement in the elderly. Neurosci Lett
1990;117:188—93.
[76] Tokuno CD, Sanderson DJ, Inglis JT, Chua R. Postural
and movement adaptations by individuals with a unilateral
362
below-knee amputation during gait initiation. Gait Posture
2003;18(3):158—69.
[77] Toussaint HM, Michies YM, Baber MN, Commissaris ACM, van
Dieën JH. Scaling anticipatory postural adjustments dependent
on confidence of load estimation in a bimanual whole-body
lifting task. Exp Brain Res 1998;120:85—94.
[78] Van Keeken HG, Vrieling AH, Hof AL, Halbertsma JP, Schoppen T,
Postema K, et al. Controlling propulsive forces in gait initiation
in transfemoral amputees. J Biomech Eng 2008;130(1): 011002.
[79] Vrieling AH, van Keeken HG, Schoppen T, Otten E, Halbertsma
JP, Hof AL, et al. Gait initiation in lower limb amputees. Gait
Posture 2008;27(3):423—30.
[80] Vernazza S, Cincera M, Pedotti A, Massion J. Balance con-
trol during lateral arm raising in humans. NeuroReport
1996;7:1543—8.
S. Bouisset, M.-C. Do
[81] Waters RL, Morris JM, Perry J. Transitional motion of head and
trunk during walking. J Biomech 1973;6:167—72.
[82] Winter DA. Human balance and posture control during standing
and walking. Gait Posture 1995;3:193—214.
[83] Yiou E, Hamaoui A, Le Bozec S. Influence of sup-
port base on arm pointing performance and associated
anticipatory postural adjustments. Neurosci Lett 2007;423:
29—34.
[84] Zattara M, Bouisset S. Chronometric analysis of the posturo-
kinetic programming of the voluntary movement. J Mot Behav
1986;18:215—23.
[85] Zattara M, Bouisset S. Posturo-kinetic organization during
the early phase of voluntary upper limb movement 1.
Normal subjects. J Neurol Neurosurg Psychiatry 1988;51:
956—65.