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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 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
Figure 1 From the task to perform to the real task: outline rotation of body segments produced by the muscles leads, in
of a system analysis. The ‘‘task to perform’’ is the task that the turn, to movements of the centre of gravity (or around it).
subject intends to execute. It can be characterized by three fac- In other words, if a muscular contraction provokes a joint
tors: category, specific parameters and environmental context. movement—–upper limb flexion for instance—–the rest of the
These different factors are taken into account at the program- body tends to move in the opposite direction. As expressed
ming level. The central nervous system issues a command that by Gray [39]: ‘‘In order to subject the body to a propul-
triggers inputs to the muscular system. The muscles mobilize sive force, it is necessary to exert an exactly equal but
the bone and joint system, which produces movement. The opposite force against his external environment. The reac-
‘‘real task’’ is the task, which is actually performed. In order tion is equal to the subject’s action («action and reaction
to characterize motor activity in a non invasive manner, periph- principle»)’’.
eral variables are measured at the muscular (electromyographic In other words, voluntary movement occurs as the result
activity: EMG) and movement (biomechanical quantities: kine- of a combination of muscle torques and ground reactions.
matic and kinetic ones) levels. Movement efficiency is evaluated However, even though they are internal, muscular forces
by the actual specific parameters values (speed, precision, etc, play a major role. As they are under the control of the
from [11]). CNS, they are the only ones that can be used to manage
the fatal effects of the external forces, and adjust coopera-

structures, both inanimate and animate ones, such as the


human body. →
1
 = ˙M
Eq. (2) (JG −−  G (Fe )) refers to a solid, rotating about a fixed
Newton’s laws are expressed by two equations (Fig. 3): axis. In this equation:
one (the dynamic resultant law) refers to linear movements,
→ →
and the other (the kinetic torque law), to rotations. They • the kinetic momentum (JG −−  ) is the angular acceleration (−−
)
relate forces to movement at every instant. Indeed, on the times the body moment of inertia (JG ), with respect to the rota-
left side of the equal sign, there is an acceleration (to within tion axis, , which goes through the body center of gravity, G;
a constant coefficient) and on the right side, an expression • the body moment of inertia (JG ) (with respect to the rotation
of external forces. axis, , which goes through G) is the body mass times the squared
More precisely, Eq. (1) relates the external forces resul- distance from G to the rotation axis;
tant (˙ Fe ) to the dynamic resultant (mG ), with G being • the resultant torque of external forces (˙ M  G (Fe )) is the resultant
the centre of gravity (CoG) linear acceleration and m, body force (˙ Fe ) times its distance to the rotation axis, , which goes
mass. Eq. (2) relates the resultant torque of external forces 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 where the vertical forces resultant is applied. Now, CoG hor-
performance fulfils task-movement constraints. izontal acceleration has been shown to be proportional to
Support base and support reaction forces. Contrary to the difference between CoP and CoG horizontal displace-
gravity, reaction forces originating from the ground (and ments [21]. Therefore, CoG horizontal acceleration may be
more generally from the physical environment) vary as a reduced when the support base perimeter is limited and
function of the forces developed by the muscles during the when the CoP initial position is close to the support base
motor act. Therefore, a clear understanding of the postural boundaries (Fig. 4).
phenomena associated with voluntary movement requires
taking the external forces into account, and the support The human body, as a system of rigid solids
reaction forces in particular. This necessity requires con- The human body is composed of anatomical ‘‘segments’’,
sidering the ‘‘support base’’, i.e. the body areas in contact including bones and the living tissues around them, such as
with the physical support. the arm or the leg. From a biomechanical point of view,
Support base characteristics depend on two factors, the it can also be considered a system of rigid solids, termed
physical properties of the support and the interface between ‘‘links’’ [28], which are movable in relation to each other,
the body and the support. The first depends on the proper- i.e. they form an ‘‘articulated chain’’ and interact. The
ties of the materials (elasticity, friction, etc.), as well as on mobility of an articulated chain is a function of the num-
geometrical parameters (flatness or curvature, inclination, ber of the degrees of freedom and of the range of individual
etc.) and design (steadiness or oscillations, translations, joint movements.
etc.). The second depends on the subject’s posture (stand- Postural maintenance is an ‘‘active’’ process. A certain
ing, on tiptoe, on the entire foot, on one foot or both feet, equilibrium state, often termed ‘‘static’’, is associated to
seated, with or without a backrest, etc). They also depend given posture maintenance. According to Newton’s laws, it
on the support base perimeter. corresponds to the condition when the resultant reaction
Biomechanical principles indicate that support base char- and the resultant torque of the external forces are nil. This
acteristics have an effect on the reaction forces applied necessity implies two sets of requirements.
to the body (and on their torque). For example, by con- The first are global requirements: they are the same
struction, the support base perimeter delimits Centre of as if the body were an invariant structure, i.e. as if the
Pressure (CoP) displacement, insofar as the CoP is the point 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- mechanical action exerted on any body point is transmit-
eral body balance is secured when the projection on the ted with no time lag to all the others. In particular, the
ground of the centre of gravity is situated within the support forces (and torques) developed during the movement are
base perimeter. The second are local requirements, which transmitted from the segment(s) that the subject intention-
are usually understated. They result from the articulated ally mobilizes to others, and from the distal one(s) to the
structure of the human body: each body part must also be support surface(s), according to the ‘‘action and reaction
balanced with respect to the underlying ones. Owing to the principle’’.
configuration of the joint contact surfaces and the shape of The initial conditions of movement are defined by body
bones, purely mechanical factors cannot fulfill this require- posture, which is usually ‘‘static’’, such as standing, sitting,
ment. This is why appropriate muscular forces, originating crouching, etc. Therefore, the dynamic phenomena implied
from equilibration reactions, are necessary even in a nat- by movement—–i.e. inertial forces (and torques)—–tend to
ural upright posture, where the soleus is the emblematic perturb balance.
muscle.
Therefore, postural maintenance proceeds from an Equilibrium state and dynamic stability
‘‘active’’ process, insofar as it results from muscle con- According to Martin [58,64], ‘‘posture denotes first the
traction, in addition to inanimate ‘‘passive’’ forces, such shape of the body, i.e. the geometrical relationship of the
as gravity. different parts to each other’’. In order to prevent any risk
Voluntary movement is a perturbation to posture. Fur- of misunderstanding, ‘‘posture’’ would be limited to the
thermore, the body segments are rigid, and each one ‘‘static’’ whole body configuration, i.e. to postural main-
is closely linked to the adjacent ones. Therefore, every tenance.
350 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
Figure 4 Center of gravity (CoG) and center of pressure (CoP) voluntarily executes a trunk extension, the lower limbs
displacements during upper limb elevation. move in the opposite direction. Several decades later,
The subject performs a right shoulder flexion (indicated by the electromyographic and biomechanical methods made it pos-
arrow). Left: the subject is standing normally. Right: the subject sible to deepen neurological data, allowing a broad survey
is standing on tiptoe, which limits CoP (solid line) and CoG (thick of muscular patterns and providing new data on motor
dash line) displacements. In both cases, it is to observe that programming.
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 PAs associated to voluntary movement
(right figure), which means that CoG horizontal acceleration is
lower. Methodological considerations
Electromyography is unique in specifying muscle activation.
Indeed, surface EMG, i.e. primarily the electrical signals
Nevertheless, posture is first a physiological concept, picked up by surface electrodes, is a convenient index
which can be defined in kinematic terms. Posture can also of muscle excitation and allows a description of muscular
be defined with respect to kinetics, i.e. to Newton’s laws. In patterns. However, kinetic (or dynamic) quantities (which
this context, postural maintenance corresponds to the con- refer to the force and energy associated with movement)
dition when the inertial resultant forces (and torques) are and kinematic ones (which refer to aspects of movement
nil (˙ Fe = 0 and ˙ M G (Fe ) = 0). On the contrary, they are besides the consideration of mass and force) must also be
not nil during postural stabilization. However, the body is in considered. They are measured using different transducers
an equilibrium state during both postural maintenance and (goniometers, accelerometers, dynamometers, etc. . .) and
postural stabilization. specific equipment, such as force plates and movement cap-
The mechanical concept of stability is helpful to ture displays. Kinematic quantities (displacement, velocity
accurately examine the influence of movement on body and acceleration) are used to characterize local movements,
equilibrium. According to mechanics, an equilibrium state i.e. segmental movements. Kinetic quantities (force, reac-
can be ‘‘stable’’ or ‘‘unstable’’. It is termed stable when tion force and torque) are used to measure whole body
the structure, which is initially in equilibrium or stationary movement. In particular, force plates, which measure sup-
state, returns to this state after a perturbation; for instance, port reaction forces, allow to determine the resultant forces
when a ‘‘static’’ equilibrium state is restored after a move- and torques, as well as the centre of pressure. The resul-
ment. To this end, the sensory-motor system must develop tant force expresses the centre of gravity linear acceleration
the forces necessary to oppose the perturbing forces: equi- to within the body mass, according to Eq. (1), and angular
libration reactions must be put into play. Between the onset acceleration to within the body moment of inertia in relation
and the end of a motor act, the structure is in a state of to the centre of gravity, according to Eq. (2) (Fig. 3). Reac-
transient disequilibrium, which is termed ‘‘dynamic equi- tion forces (and torques) at joint level can only be estimated
librium’’. from a model (‘‘direct dynamics’’ or ‘‘inverse dynamics’’
‘‘Dynamic stability’’ is the process that allows dynamic modeling).
equilibrium at every instant, which means that ‘‘static’’ In addition, we must also keep in mind the conceptual
balance will be restored at the end of isolated move- partitioning of the body into a ‘‘focal’’ and a ‘‘postural’’
ments, or balance conditions will be achieved during chain when examining the influence of movement on pos-
the next ‘‘stance’’ phase in periodic movements such as ture. In particular, we must ask which body part executes the
walking. In other words, dynamic stability allows a task- focal movement, i.e. which one is the ‘‘primum movens seg-
movement to be performed efficiently, from the onset to ment’’ (PMS). Also, we must determine which muscle of the
the end. Dynamic stability is managed by the equilibra- focal synergy is the first to be activated, i.e. the ‘‘primum
tion function, i.e. by ‘‘postural stabilization’’ as mentioned movens muscle’’ (PMM), according to Beevor’s classification
above. [6].
The stability limits of a system of articulated solids are Last, to be significant, the experimental data must be
exceeded when the projection of the centre of gravity onto linked to an adequate evaluation of the movement issue, in
the ground falls outside the support base perimeter, or if terms of task performance (displacement, velocity, force,
the ground reaction force vector is not located within the precision, etc.). Indeed, EMG and biomechanical patterns,
stability cone [60], which is equivalent. Within the stability as well as quantitative measurements (amplitude and tim-
limits, a given posture is stable as long as the internal forces 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 APAs: movements which precede movement
risk of misinterpretation related to unexpected support base In order to present APAs, it is easier to refer first to unilateral
effects. shoulder flexions, performed with the upper limb stretched
out and on a single support base. Indeed, as it is a very simple
APA measurement paradigm, one can suppose that the postural mechanisms,
As it is well-known, the CNS triggers muscle excitation, which are involved, can guide the interpretation of more
which in turn induces muscle activation, and hence mus- complex experimental situations. Nevertheless, a compar-
cle contraction after a certain time (‘‘electro-mechanical ison with locomotion can be helpful, in that locomotion is
delay’’: EMD). Then, movement can occur and biomechani- a lower limb movement, including a periodic support base
cal variables can be recorded. transfer.
Therefore, there are various possibilities for measuring A given sequence of EMGs was observed in the lower limbs
APAs. APAs can be defined with respect to EMGs, i.e. they are and trunk muscles before the onset of the Anterior Del-
measured by the time interval between postural and focal toïdeus EMG, which is the PPM (Fig. 7). First, the ipsilateral
muscle EMG onset. APAs can also be defined with respect Soleus is deactivated 50 to 100 ms prior to PPM onset. Then
to biomechanical variables, i.e. the time interval between there is a sequence of phasic activations and deactivations,
global dynamics (or postural movement) onset and focal starting with activation of the contralateral Tensor Fasciae
movement onset. Of course, it is also possible to define APAs Latae and Rectus Femoris. This sequence is reproducible and
by the time interval between the onset of a biomechani- specific to the task under consideration [85].
cal quantity and an EMG onset. This method raises various A sequence of segmental accelerations corresponds to
problems, such as the location and parameter of the two the EMG sequence. Thus, before the onset of upper limb
signals. Also, the influence of electro-mechanical delay on acceleration, i.e. PMS onset, the lower limbs, pelvis, trunk
the results, which differs according to the muscles and the and shoulder display anticipatory accelerations (refer later
task parameters, cannot be disregarded. on to Fig. 9): the existence of accelerations indicates that
Last, there is a technical problem, which cannot be anticipatory activities are actually movements. More pre-
underestimated: how to estimate postural and focal onset. cisely, there is a given order in anticipatory accelerations,
Indeed, special care must be taken in determining EMG starting with the contralateral lower limb and hip, the
onset and cessation. Also, it is less accurate to consider dis- ipsilateral ones, and ending with the ipsilateral shoulder:
placements rather than accelerations, because of the slower it follows a bottom-up progression, as the postural seg-
variation rate at the onset of movement. 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 anticipatory accelerations. The anticipatory sequence starts
to the task under consideration. Since these results are at the stance leg, then at both hips and at the ipsilateral
consistent with EMG ones, the EMGs could be related to shoulder, and afterwards at the contralateral shoulder [19].
local accelerations: the local movements are not ‘‘passive’’ Neck and head displacements in addition to trunk displace-
movements. ments and their corresponding EMGs have also been reported
In gait initiation, the anticipatory EMG sequence (Fig. 7) [81]. Finally, all the local accelerations indicate that the
includes Soleus deactivation, followed by activation of both movement is accelerated forwards and towards the stance
Tibialis Anteriors 60 to 110 ms later. Then, the Soleus of foot.
the forthcoming swing limb, which is the PPM of the heel- Here again, the sequence follows a bottom-up progres-
off (HO), is activated. Thereafter, EMG bursts occur in the sion. But, as the postural segment accelerations proceed
Rectus Femoris of the swing limb and in the Semitendi- from the support surface, it seems more appropriate to con-
nosus of the stance limb [20]. In addition, before heel-off sider that the progression order follows a ‘‘posturo-focal
(HO) (i.e. PMS onset), the pelvis, trunk and shoulder display 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 is executed by the contralateral limb (Fig. 9). This char-
chain, and then to the focal one. acteristic is called ‘‘dynamic asymmetry’’ as opposed to
Finally, APAs are postural movements. They precede vol- ‘‘dynamic symmetry’’, corresponding to bilateral movement
untary movement. And like voluntary movement, they are a [16]: in terms of posturo-kinetic programming, it is not
dynamic phenomenon. 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
APAs and postural programming be said that dynamic asymmetry and dynamic symmetry
According to Bernstein’s ideas [9], it is assumed that PAs are ‘‘pre-programmed’’. The organization of the postural
constitute a part of the motor program. In other words, it sequence according to the forthcoming focal limb(s) is a gen-
is admitted that there is a ‘‘postural task’’ associated with eral feature. For example, gait initiation displays a mirror
the focal one. APAs constitute a valuable tool in identifying organization according to the stepping leg, i.e. a dynamic
task parameters, which are programmed. Indeed, since by asymmetry pattern, whereas it is a dynamic symmetry pat-
definition, APAs precede the onset of focal movement, they tern in standing jump initiation.
cannot result from re-afferentation triggered by the focal Finally, APAs are dynamic, polarized, and task specific.
movement: they are ‘‘pre-programmed’’. If APAs occur in They are programmed in relation not only to the focal
the postural chain, it can be said that the postural chain is movement parameters per se, but more generally to task-
programmed. If APAs vary in relation to task parameters, it movement parameters. In addition, they are ‘‘adaptable’’,
can be assumed that the postural chain is programmed in as they depend on the functional state of the motor system,
relation to these parameters. which will be discussed later on.
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 Dual role of APAs
are graded as a function of movement peak velocity (Fig. 8). APAs as a counter-perturbation. The role of APAs may
A similar result has been found in shoulder flexion, and the be proposed on the basis of an analysis of forces acting
curvature of the relationship increases with additional iner- at the shoulder level and at the body’s centre of gravity
tia [14]. Moreover, APAs depend on ‘‘postural conditions’’, (Fig. 10), whose directions are opposite. From this analysis,
as will be discussed later on. it can be assumed that ‘‘APAs tend to create inertial forces,
In addition, it must be stressed that APAs are not only which, when the time comes, will counterbalance the dis-
a function of the specific task-movement parameters. They turbance to postural equilibrium due to the intentional
also depend on its location with respect to the body’s axes forthcoming movement’’ [15]. According to this hypothesis,
of symmetry. Indeed, the role played by the homonymous the intentional movement is considered a ‘‘perturbation’’,
muscles and segments is inverted, when the movement 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 In lower limb movements involving a transient change
by Belenkii et al. [7]. In other words, APA represents a of the support base perimeter, such as lower limb flexion,
‘‘counter-perturbation’’, which would be responsible for APAs have been shown to be related not only to the counter-
the stabilizing action. perturbation of the forthcoming movement, but also to
In this context, it was proposed that one consider that the ‘‘body weight transfer’’ onto the forthcoming stance foot
postural perturbation, associated with voluntary movement, [32]. This transfer can be interpreted as being related to
as well as the counter-perturbation, might be characterized postural stabilization, as well as to the necessary unloading
by two biomechanical factors [16]. They correspond to the of the forthcoming moving foot.
two vectors to which the perturbation force system may be In locomotion, there is a succession of balance losses and
reduced, i.e. its resultant and its torque (in reference to the recoveries, which corresponds to the periodic support base
body’s centre of gravity, for example). One is linear, while transfer. It has been demonstrated that during gait initia-
the other is rotational. Both may include three orthogonal tion, the greater the forward fall of the centre of gravity,
components. the faster the progression velocity [21]: APAs induce postural
However that may be, as APAs are triggered by a feed- destabilization, which is necessary to initiate gait. In addi-
forward command and are specific to the characteristics of tion, the initiation phase also includes body weight transfer
the motor task, they have to be determined from previous towards the forthcoming stance foot, which can be inter-
knowledge of its perturbing effect [84]. More precisely, APAs preted as being related to postural stabilization.
are programmed according to the expected perturbation and APAs as a perturbation and a counter-perturbation.
not to the actual one [77]. Finally, it cannot be excluded that APAs play a dual role
APA as a perturbation. The contention according to which simultaneously, i.e. to perturb balance and to counteract
APAs represent a counter-perturbation has been argued the perturbation, as suggested by gait initiation: acceler-
within the framework of segmental movements performed ations occur at the ipsilateral hip at heel-off, the forward
on a continuous support base. But, the support base can also direction of which is opposite to the hip perturbation at this
change transiently, as in lower limb movements, or peri- instant [30].
odically, as in locomotion. These movements will provoke Indeed, just as in shoulder flexion initiation, lower-limb
a perturbation of the whole kinematic chain, but the cor- forward displacement at heel-off will induce a perturbation
responding APAs have been shown to play a more complex at the hip joint, which is directed backwards. It must be
role. compensated in order to prevent the hip from being pushed
356 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
Figure 10 Interpretation of the finality of anticipatory pos- be minimized: mechanical energy, metabolic energy, neu-
tural adjustments. The filled arrows correspond to the actual ral inputs, etc. Obviously, partial responses can only be
recorded biomechanical data; the broken arrows correspond provided, although it is tempting to consider that the APA
to theoretical parameters. , angular displacement of the sequence complies with an optimal control process, whose
upper limb(s); Aw,  r , and  tangential, radial and total upper criteria could vary according to the task-movement param-
limb acceleration. Rx and Rz , antero-posterior and vertical eters and the subject’s functional state.
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
Posturo-kinetic capacity: a limiting factor of
observed that the perturbation (indicated by broken arrows at performance
the shoulder level), and the corresponding APA (indicated by
full arrows at the center of gravity) display opposite directions Posturo-kinetic and foco-kinetic capacities
(from [17]). APAs have been presented as the optimal biomechanical
means to initiate voluntary movement and to approach
postural programming. In order to broaden the approach
backwards at the very instant when it has to serve as a to motor programming, it is tempting to return to the
fulcrum for hip rotation. hypothesis according to which motor control includes a
More generally, trunk and shoulder movements during ‘‘focal’’ and a ‘‘postural’’ component [38] and to con-
walking, which induce oscillation in the upper limbs, play sider a ‘‘postural system’’ and a ‘‘focal system’’. To one
a major role in regulating body rotation with respect to the would correspond a ‘‘posturo-kinetic capacity’’ (PKC), as
vertical axis [33]. They diminish whole-body rotation dur- proposed by Bouisset and Zattara [16], and to the other,
ing the single support and favor a more gradual change in a ‘‘foco-kinetic capacity’’ (FKC) (Fig. 11). PKC means the
the rotation direction during double support. In addition, ability to develop a counter-perturbation to the posture per-
body rotation with reference to the antero-posterior axis turbation induced by voluntary movement, and therefore
is modified by upper-limb oscillation, such that direction the ability to manage the perturbation to balance asso-
changes are more progressive. Consequently, these upper- ciated with the forthcoming movement. FKC means the
body movements play a role in postural stabilization, as was ability to execute the voluntary movement per se, i.e. to
claimed, after Elftman [33], by authors such as Cappozzo mobilize the focal segments. PKC is believed to depend
[22], Cromwell et al. [27], Jian et al. [50] and Winter [82]. on the configuration of the postural chain, the environ-
Finally, APAs appear to play a dual role, that is, postural mental context and the functional state of the postural
stabilization and the creation of propulsive forces. One or system.
Posture, dynamic stability, and voluntary movement 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
Figure 11 Posturo-kinetic capacity (PKC) and foco-kinetic assumption is consistent with theories on motor planning.
capacity (FKC). The motor command (‘‘command’’) is assumed Another possibility for assessing PKC is to consider the
to result from the intent of achieving a given task, which can be biomechanical or EMG pattern. If it is assumed that the
characterized by three factors: category, specific parameters, primary synergy results from an optimization criterion, a dif-
and environmental context (refer to Fig. 1). ‘‘Performance’’ ferent synergy can be assumed to indicate a less economical
characterizes the task that is actually performed as a result of process: performance would be achieved at the expense of
the movement. Under the hypothesis according to which motor increased postural cost. EMG or biomechanical profiles have
command includes a ‘‘focal’’ and a ‘‘postural’’ component, it to be examined and their main features (amplitude, dura-
is supposed that there is a ‘‘postural system’’ and a ‘‘focal tion, time to peaks, etc.) measured in order to characterize
system’’. To one would correspond a ‘‘posturo-kinetic capac- the postural pattern. But the method has also proved to be
ity’’ (PKC), and to the other, a ‘‘foco-kinetic capacity’’ (FKC), useful even though it is qualitative, as will be exemplified
both of which must be seen as systemic entities. PKC means the in the case of impairments.
ability to develop a counter-perturbation to the posture per- Finally, the influence of a given factor on PKC, such as
turbation induced by voluntary movement, and therefore the postural stability or impairment, is assessed by comparing
ability to manage the perturbation to balance associated with the corresponding data to the control, for example, the data
the forthcoming movement. FKC means the ability to execute obtained for comfortable stance or able-bodied subjects.
the voluntary movement per se, that is, to mobilize the focal To conclude, assessing PKC means evaluating the postu-
segments. PKC is assumed to depend on the configuration of the ral cost of a task. The method involves relating performance
postural chain, the environmental context and the functional to APA and considering muscular and/or biomechanical pat-
state of the postural system. FKC is assumed to depend on the terns.
configuration of the focal chain, the environmental context and
the functional state of the focal system. It is assumed that per- PKC and postural configuration
formance is the end product of the cooperation between PKC In the preceding sections, voluntary movement was said
and FKC. 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-
It is assumed that performance is the end product of ment is usually superimposed on an initial ‘‘static’’ posture
the cooperation between PKC and FKC. In other words, and is followed by a final ‘‘static’’ posture as well.
PKC would control task performance in preparing the motor According to the literature (for a review, see [13]), APAs
action, supporting its course and contributing to its effi- depend on postural conditions, i.e. mainly on the support
ciency. In this view, PKC would operate throughout the entire base area and stability. However, when the initial and final
motor act. postures are less stable [37,47], some authors find that APA
Finally, both PKC and FKC must be seen as systemic enti- duration increases while others find it decreases. These dif-
ties: their properties proceed from a combination of their ferences are not easy to interpret, in that the experimental
functional components, mainly the muscular, bone and joint, conditions are so diverse that comparisons are difficult.
and sensory-motor systems. Both PKC and FKC are assumed Moreover, performance is seldom reported. However, two
to be ‘‘dynamic processes’’, which vary at each instant in sets of experiments help delve deeper into the influence of
relation to the current conditions. the support base area on PKC.
In the first one, the voluntary movement (a pointing
PKC assessment task) was performed by standing subjects. The support base
It is clear that PKC assessment allows approaching the pos- configuration differed by the size of the antero-posterior
tural strain related to a given task. The problem has already distance between the heels [83]. When the distance was
been discussed in detail [12,13]. To summarize, the method increased from 0 to 40 cm, i.e. when postural stability was
for assessing PKC involves evaluating the postural input to enhanced, maximal velocity increased, while the APA dura-
the motor system, and the corresponding output, when max- tion remained equivalent. Thus, the PKC was greater when
imal performance is required. The simplest procedure is the body support size was enlarged (PKC = 0.5, as compared
to relate performance (i.e. the maximal movement veloc- to PKC = 0.2) and the postural cost was lower. Hence, postu-
ity, or the maximal muscular force, etc. depending on the ral cost is a function of postural stability.
358 S. Bouisset, M.-C. Do

In the second experiment, the voluntary movement vated simultaneously, at the expense of greater forces to
(a flexion or a flexion-extension of the lower limb) was be developed during the movement itself. In other words,
performed while various initial and final postures were the lack of APAs suggests that postural stabilization is
considered [32,66]. The general trend was an increase in a major constraint instead of muscular energy minimiza-
APA duration and a decrease in maximal velocity when tion.
the initial and final postures were less stable: PKC was
reduced in relation to postural stability. Further, APAs were PKC and the functional state of the motor system
absent when the subjects stood on one foot, from the As stated above, performance results from a hierarchi-
beginning of the lower limb movement to the end [66]: cal process, associating three different components, i.e.
as APAs induce dynamics, they can potentially perturb the central nervous, muscular, and bone and joint systems
body balance, and are no longer present when the pos- (Fig. 1). More, or less, efficient performance depends on
ture is excessively unstable. Therefore, in conditions of their functional state, which is not easy to assess, either
high stability demand, the CNS may reduce and even sup- clinically or scientifically. Nevertheless, it is agreed that it
press APAs, as protection against their possible destabilizing varies according to impairment or rehabilitation, condition-
effects. Then, the focal and the postural chains are acti- 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 velocity was slower than in the able-bodied controls, but
the elderly [46,49,57,75] and in hemiplegics [46], whereas that APAs lasted much longer [31]. Thus, PKC is less in para-
maximal movement velocity is reduced. Also, APAs van- plegics: PKC = 3.4, as compared to PKC = 9 for able-bodied
ish when people are bedridden for several weeks [40], subjects.
and in Parkinsonians [5,29,18]. Moreover, postural patterns Postural synergy is also different: other scapular girdle
are changed in Parkinsonians, hemiplegics, etc. Therefore, and trunk muscles are activated, especially on the contralat-
sensory-motor impairments diminish the functional state, eral side (Fig. 12). If it is assumed that the primary synergy
i.e. ‘‘sensory-motor capacity’’, which is the resultant of PKC corresponds to an optimization criterion, the transition from
and FKC. one synergy to another can be assumed to correspond to a
In other words, PKC assessment in sensory-motor impair- less economical process.
ments requires consideration of those impairments, which In other words, paraplegics, whose equilibrium is more
result in a restriction of joint mobility, regardless of the unstable than able-bodied subjects, could not develop PAs
exact origin (bones, joints, tendons, muscles or nervous sys- adapted to the perturbation provoked by the motor task
tem). It is particularly interesting to discuss paraplegics and without an excessive balance risk: the perturbation, and
lower-limb amputees. hence the required performance, should be reduced. There-
Paraplegia and PKC. In paraplegics performing a pick- fore, due to PKC reduction, voluntary movement is less
and-place task, it was established that maximal movement 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 S. Bouisset, M.-C. Do

Lower limb amputation and PKC. Similarly, unilateral [9] Bernstein N. Coordination and regulation of movements. Perg-
lower limb amputees showed many kinetic changes in gait amon Press; 1935 (Amer. translation, 1967).
initiation [61,62,73,76,78,79]. Although maximal progres- [10] Bouisset S. Relation entre support postural et mouvement
sion velocity is similar, regardless of the stepping side intentionnel: approche biomécanique. Arch Int Physiol Bioch
(prosthetic or sound limb), it is slower that in controls. Fur- Biophys 1991;99:A77—92.
[11] Bouisset S. Biomécanique et physiologie du mouvement. Paris:
thermore, the muscular pattern was necessarily different.
Masson; 2002.
PKC can be discussed with reference to the forthcoming [12] Bouisset S, Do MC, Zattara M. Posturo-kinetic capacity assessed
stance leg [61] (Fig. 13). When above-knee (AK) amputees in paraplegics and parkinsonians. In: Woolacott M, Horak F, edi-
initiate gait with the sound limb (the prosthetic leg is the tors. Posture and Gait; control mechanisms, vol II. University
forthcoming stance leg), APAs are longer and, since the pro- of Oregon Books; 1992. p. 19—22.
gression velocity is the same, the PKC is lower (PKC = 1.5). [13] Bouisset S, Le Bozec S. Posturo-kinetic capacity and pos-
When they initiate gait with the prosthetic limb (the sound tural function in voluntary movements. In Latash ML, ed:
leg is the forthcoming stance leg), APAs are shorter while Progress in Motor Control, vol II: Structure-Function Relations
the progression velocity remained the same: PKC is higher in Voluntary Movements. Human Kinetics. 2002, Chapter 3:
(PKC = 2.7). In controls, PKC is approximately the same p. 25—52.
[14] Bouisset S, Richardson J, Zattara M. Do anticipatory postural
as in amputees when the stance limb is the sound one
adjustments occurring in different segments of the postural
(PKC = 2.8). chain follow the same organisational rule for different task-
In addition, PKC is less affected by below-knee ampu- movement velocities, independently of the inertial load value?
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