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Motor Control Theories and Their Applications
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Medicina (Kaunas). 2010 ; 46(6): 382–392.
Motor Control Theories and Their Applications
Mark L. Latash1, Mindy F. Levin2, John P. Scholz3, and Gregor Schöner4

1Department of Kinesiology, The Pennsylvania State University, University Park, PA, USA
2Department of Physical Therapy, McGill University, Montreal, Canada

3Department of Physical Therapy, University of Delaware, Newark, DE, USA
4Institute for Neuroinformatics, Ruhr University, Bochum, Germany
Summary
We describe several influential hypotheses in the field of motor control including the equilibrium-
point (referent configuration) hypothesis, the uncontrolled manifold hypothesis, and the idea of
synergies based on the principle of motor abundance. The equilibrium-point hypothesis is based
on the idea of control with thresholds for activation of neuronal pools; it provides a framework for
analysis of both voluntary and involuntary movements. In particular, control of a single muscle
can be adequately described with changes in the threshold of motor unit recruitment during slow
muscle stretch (threshold of the tonic stretch reflex). Unlike the ideas of internal models, the
equilibrium-point hypothesis does not assume neural computations of mechanical variables. The
uncontrolled manifold hypothesis is based on the dynamic system approach to movements; it
offers a toolbox to analyze synergic changes within redundant sets of elements related to
stabilization of potentially important performance variables. The referent configuration hypothesis
and the principle of abundance can be naturally combined into a single coherent scheme of control
of multi-element systems. A body of experimental data on healthy persons and patients with
movement disorders are reviewed in support of the mentioned hypotheses. In particular,
movement disorders associated with spasticity are considered as consequences of an impaired
ability to shift threshold of the tonic stretch reflex within the whole normal range. Technical
details and applications of the mentioned hypotheses to studies of motor learning are described.
We view the mentioned hypotheses as the most promising ones in the field of motor control, based
on a solid physical and neurophysiological foundation.
Keywords
Motor control; equilibrium-point hypothesis; synergy; uncontrolled manifold hypothesis; motor
disorders
1. Introduction
Motor control is a relatively young field of research. It may be defined as an area of natural
science exploring how the central nervous system (CNS) produces purposeful, coordinated
movements in its interaction with the rest of the body and with the environment. Hence, the
main goal of motor control research is to create a formal description, operating with exactly
defined variables, of the physical and physiological processes that make such movements
possible. Progress in motor control over the recent years has been slowed down by the lack
Address for correspondence: Mark L. Latash, Department of Kinesiology, Rec.Hall-268, Penn State University, University Park, PA
16802, USA, mll11@psu.edu.
Latash et al. Page 2
of a broadly accepted and exactly defined set of notions that would be specific for typical
problems of motor control, an adequate language for this area of research (see Gelfand &
Latash 1998).
It is very hard to look for an adequate set of notions in an area that does not have them, but it
is also very challenging and exciting. It is much more simple (and tempting) to borrow one
of the developed approaches from another field that shares “key words” with motor control,
for example classical mechanics, control theory, and engineering. One should keep in mind,
however, that such approaches have strict limitations. They can provide tools that may help
find answers to questions after the questions have been formulated. But they cannot offer an
adequate formulation of questions in a field that differs from the areas for which these
approaches have been developed.
Biological objects belong to the physical world and they are alive. So, help with formulating
questions may be expected to come from physics and biology (physiology), not from control
theory and engineering developed to deal with objects in the inanimate world. Physics of the
inanimate nature, while being a highly developed science, has troubles dealing with typical
problems of motor control. First, in contrast to movements in the inanimate world,
movements of biological objects are intentional and purposeful. These two notions cannot be
easily incorporated into physics. Another problem is that the body is a very complex system,
maybe too complex to be studied with the currently available physical tools, and many
crucial variables are not directly measurable or even identifiable. Physics of living systems,
unfortunately, does not exist so far, although one of the authors of this paper (MLL)
graduated some 30+ years ago from the Moscow Institute of Physics and Technology, and
his major was indeed the non-existing Physics of Living Systems.
Two aspects of motor control have been traditionally considered separately from each other.
The first relates to the nature of physiological variables that are used by the brain to control
muscles. The second relates to the problem of motor redundancy (Bernstein 1967): How
does the brain select particular solutions from infinite sets afforded by the redundant design
of the neuromotor system at all levels of its analysis? Within this brief paper, we will review
two theories that, to our opinion, are most promising in moving the field of motor control
closer to physics of living systems.
The first of these theories is the equilibrium-point hypothesis (referent configuration

hypothesis, Feldman 1966, 1986, 2007), which is based on Galileo’s principle of relativity
and the fact that neurons are threshold elements. The second one is the uncontrolled
manifold hypothesis (Schöner 1995; Scholz & Schöner 1999), which, in combination with
the principle of abundance (Gelfand & Latash 1998, 2002), suggest a novel approach to the
problem of motor redundancy. There have been several recent attempts to move towards
combining the two approaches into a single theory of motor control (Martin et al. 2009;
Latash et al. 2010).
2. Equilibrium-point theory
Explanation of normal and disordered motor control
The equilibrium-point (EP) theory was first described by Feldman in a series of papers in the
1960’s and 70’s. Over the past 50 years, the EP theory has been revised and refined from a
hypothesis describing the control of a simple single-joint system to a theory addressing the
production of complex movements, such as multi-joint movement and locomotion, and
uniting the processes underlying movement production and perception.

Fundamental to the EP theory is the notion that threshold position control underlies
intentional motor actions. To perform such actions, electrochemical influences descending
from the brain in the presence of proprioceptive feedback to motoneurons are transformed
into changes in the threshold muscle lengths or joint angles at which these motoneurons
begin to be recruited, thus setting the spatial activation range in reference to the body
geometry. This allows control levels of the CNS to specify where, in spatial coordinates,
muscles are activated without being concerned about exact details on when and how they are
activated. In the most advanced formulation of the EP theory, activity of each muscle
emerges, without any programming, depending on the difference between the actual
configuration of the body and its threshold (referent) configuration, as well as on the rate of
changes of this difference (Feldman, in press). A hallmark of the EP theory is that it
describes motor control based on neurophysiological and physical principles.
In a series of experiments on the elbow joint in humans combining involuntary movement

elicited by unloading of the pre-loaded arm (the unloading reflex), voluntary changes in the
joint angle, and full relaxation of the elbow muscles while they were stretched, Asatryan and
Feldman (1965) identified invariant and modifiable neurophysiological variables involved in
these motor actions.
Involuntary behaviour was analyzed by unloading experiments in which the forearm was
placed on a horizontal manipulandum and subjects resisted a specific load torque at a
specific position with the elbow flexors. The elbow position, load torque and flexor and
extensor electromyographic (EMG) activity were measured in this initial state, called the
equilibrium point (EP) of the system (Fig. 1, filled circle a). The EP is thus comprised of
both an equilibrium position and its associated equilibrium torque. In subsequent trials, from
this initial EP, the elbow flexors were unloaded to varying amounts. Subjects were
instructed not to intervene voluntarily when the unloading occurred. This means that they
had to let the arm move to its natural new position after the unloading and not try to make a
correction, to return the arm to the initial position or to relax completely.
After each time that the arm was partially unloaded, the arm naturally found a different final
EP (open circles), depending on the amount of unloading that occurred. These final EPs
were plotted on a torque-angle graph and together with the initial EP, they formed a smooth
non-linear torque-angle characteristic (upper left solid curve in Fig. 1).
After that, the subject was asked to voluntarily change the initial position against a load and
the procedure was repeated from a new initial EP, yielding a new torque-angle characteristic
(Fig. 1, right solid curve). In this way, a family of torque-angle characteristics was recorded.
Finally, subjects were asked to fully relax their muscles while the elbow was extended by
the manipulandum and a torque-angle characteristic of the subject’s passive arm muscles
was obtained (dashed curve in Fig. 1). The first two unloading characteristics (starting from
points a and b) were similar: for each of them, the torque was non-linearly related to the arm
position, and EMG activity changed depending on the load. The characteristics were
somewhat different in terms of shape which may have resulted from differences in the
mechanical properties of muscles in different parts of the angular range, rather than from a
voluntary action. Each unloading curve merged with the passive joint characteristic at a
specific joint angle (R). At these points R, muscles became silent and ceased to generate
active torque. These threshold angles were different for different characteristics (Fig. 1).
From these experiments, Asatryan and Feldman (1965) concluded that the threshold angle,
R, at which muscles ceased to be active was invariant for a given initial set point or
command. When the subject intentionally changed the initial arm position, a new R value
was specified. The experiments also showed that to fully relax the arm muscles, the R had to
be shifted beyond the upper biomechanical limit of the elbow joint (Fig. 1, R+) so that the

muscles could remain silent in the entire biomechanical range of the elbow joint angle. In
contrast, to fully activate the muscle, even at the shortest muscle length, the R had to be
shifted beyond the lower limit of the biomechanical range (Fig. 1, R−). Thus, the CNS
specifies R and its associated torque-angle characteristic and regulates the range of R within
or beyond the biomechanical range of the joint. These empirical results underlie the EP
theory.
It should be noted that for each torque-angle characteristic, neither the arm position, muscle
torque, force, EMG, nor stiffness (the slope) was invariant. The EMG activity, for example,
simply scaled with the magnitude of the load (Fig. 1, vertical segments near EPs), in
accordance with the known EMG-force relationship (Bigland & Lippold 1954). The
invariant variable is the R.
Disorders in motor function following lesions to the CNS have been attributed to deficits in
the range of regulation of R (Levin and Feldman 1994). Instead of using an unloading
procedure, R has been estimated in patients with stroke or cerebral palsy (CP) as the joint
angle at which the tonic stretch reflex threshold (TSRT) is reached when the system is at
rest (Fig. 2). Stretch-reflex thresholds depend on velocity (dynamic stretch reflex threshold;
DSRT). One can determine DSRTs (asterisks in Fig. 2) from stretches at various velocities
and extrapolate these value to zero velocity to estimate the TSRT (Levin 2000). In non-
disabled individuals, DSRTs can usually only be evoked in muscles at rest if the stretch is
performed at a high velocity (> 300°/s; Levin 2000; Thilmann et al. 1991; Fig. 2, dashed
diagonal line). However, in adults with stroke and children with CP, muscles stretches at
speeds as low as 8°/s, applied to the elbow joint, can generate DSRTs (Fig. 2, solid diagonal
line).
Previous studies in patients suggest that: 1) the TSRT lies within the physiological range of
motion of the elbow (Fig. 2, TSRTs); 2) the TSRT is inversely correlated with the degree of
clinical spasticity (i.e. the lower the TSRT the higher the spasticity); 3) the TSRT is a better
measure of spasticity than the gain of the stretch reflex (Levin 2000;Jobin & Levin 2000).
Jobin and Levin (2000) quantified the TSRT in the elbow flexors of children with CP and
tested the stability of this measure as a possible estimate of spasticity. Fourteen children
with CP and eight typically developed children participated in the trial. DSRTs were
evaluated by performing eight sets of stretches at seven, randomly selected velocities
between 8 and 160°/s using a torque motor. The joint angle and velocity corresponding to
the onset of the EMG in elbow flexors or extensors were recorded during each stretch. These
points were the DSRTs for each stretch velocity. A linear regression line was computed
through the DSRTs and the TSRT, or the SRT at 0°/s was extrapolated. For the elbow
flexors, TSRTs expressed in angular coordinates occurred later (closer to full elbow
extension) for slow velocities of stretch compared to fast velocities of stretch. The test–retest
reliability of the computed TSRTs was estimated to be good (ICC=0.73, p<0.001). An
important conclusion of this study was that the TSRT is a valid measure of spasticity,
presently unavailable in clinical practice.
The use of this fundamental concept of the EP theory has led to a new and more in depth
understanding of the mechanism underlying motor control deficits in patients with CNS
lesions. Limitations in the ranges of regulation of the TSRT have been shown to correspond
to the appearance of abnormal muscle activation patterns when patients attempt to make
voluntary movements, such as excessive co-activation (Levin et al. 2000). Also, ranges in
which typical patterns of muscle activation can occur, such as reciprocal activation, have
also been described using the TSRT approach. Using this basic notion of the EP theory,

clinicians can identify specific deficits in motor control and work with patients to increase
the range of regulation of TSRT in order to improve motor function.
3. The concept of the uncontrolled manifold

A general principle of movement generation in organisms is what Gelfand and Latash
(1998) have referred to as the principle of abundance: there are many more degrees-of-
freedom available for any specific task than strictly needed. For instance, most tasks
performed with our arms involve positioning, perhaps orienting the hand in space. That
requires between 3 and 6 degrees-of-freedom. Our arm has at least 10 degrees of freedom,
however, and even more if we allow for movements of the upper body. Similarly, almost all
joints are actuated by many more muscles than a pair of agonist and antagonist muscles that
would be minimally required. When we grasp objects, our four fingers generate surface
forces and moments which could minimally be brought about by just two fingers that oppose
the thumb.
The principle of abundance or task-relative redundancy gives rise to the question of how the
many degrees-of-freedom are harnessed to achieve a given task. This question has been
known for a long time as the degree-of-freedom problem (Bernstein 1967) and is central to
much research in motor control. Researchers have asked, for instance, which variables the
CNS "uses" to plan, time, and control movement.
There is consensus now that movement planning is best characterized in external, task-
relevant coordinates, such as the direction of movement of the end-effector in external
space. Response times, for instance, are functions of spatial task parameters. Knowing end-
effector movement direction beforehand leads to a stronger reduction of response time than
knowing movement extent (Rosenbaum 1980). Although such dependences do not preclude
that movement preparation may also reflect effector-level processes, a coherent and
comprehensive account of movement preparation can be provided at the level of end-
effector variables (Erlhagen & Schöner 2002). This is also consistent with the
neurophysiology of motor and premotor cortex. The majority of cells in these areas display
relatively uniform tuning curves to such spatial movement parameters as the direction in
space of end-effector motion or the direction in space of forces that act on the end-effector
(Georgopolous et al. 1982, 1992). These tuning curves may depend also on joint
configuration (Scott & Kalaska 1995), but largely retain their uniform shape, again
suggesting an account in terms of this spatial parameter.
Many different mechanism seem to be involved in controlling the timing of movements,

from spinal pattern generators, cerebellar estimators, to cortical mechanisms (Ivry et al.
2004). The abstract functional description of timing as a form of neuronal oscillation
accounts for coordination among multiple timed movements through the coupling among
such oscillators (Schöner & Kelso 1988). Such coupling establishes stable patterns of
relative timing. Stability, the capacity to restore the pattern following stochastic or external
perturbations, is constitutive for such patterns of coordination. This is best seen when
stability is lost. Generically, the pattern in which homologous limbs alternate ("anti-phase")
becomes unstable for higher movement frequencies, leading to a switch to the in-phase
pattern. Although it is tempting to think of the mechanisms that establish stable patterns of
coordination as residing at a level close to the neural control of the effectors (Grossberg et
al. 1997), it turns out that this is not appropriate. In a series of elegant experiments,
Mechsner and colleagues (2001) exploited the instability of anti-phase to establish whether
anti-phase is defined anatomically (homologous muscles alternating) or spatially (movement
of two end-effectors toward each other). It turned out, that it is the spatially defined anti-

phase pattern that becomes unstable at higher frequencies, even though it is anatomically
close to the stable in-phase pattern.
The control of limb movement cannot be achieved entirely at a level of task-relevant spatial
variables. At some point, muscles must be activated, forces must be generated, and joints
must be moved. Even so, since Nikolai Bernstein's studies of movements in skilled workers,
the notion has been around that the control of limb movements gives priority to the spatial
shape of the movement over the trajectories of the individual joints (see account in Zatsiorky
1998). For many years, research efforts were aimed at establishing which variables the CNS
controls when it generates a movement trajectory. Two candidates were spatial end-effector
variables or joint-level variables. One way to ask the question was by looking for invariance,
that is, finding out if movement trajectories are simpler and depend less on task parameters
when represented in end-effector vs. in joint coordinates (Morasso 1981). Although spatial
representations of movement seem to capture more of the simplicity and invariance
(Lacquaniti 1989), overall no clear pattern emerged. Moreover, most of this work used tasks
in which the effector did not have abundant degrees-of-freedom, so in principle, spatial and
joint-level trajectories were strictly transformable into each other. This weakened the case
for identifying any one of the variables as being “more controlled" than any other.
Today there is consensus that the appropriate operational definition of a variable being
"controlled" is that it is stabilized against perturbations (Won & Hogan 1995). The
variability across trials is then a possible measure of stability: highly stable states resist
stochastic perturbations more than less stable states. Even variance, however, cannot help to
decide which variable is the preferred one in human motor control when a non-abundant
task setting is used: it is true also at the level of variance that a one-to-one mapping between
joint angles and spatial end-effector coordinates precludes any decision about a control
priority.
In a task-effector system that has abundant solutions, or is redundant in more conventional

parlance, such a decision can be made, however. When the CNS has the "choice" among an
ensemble of joint-level realizations of the same spatial movement of the end-effector, we
may investigate if it stabilizes these choices at the joint level as much as it stabilizes the end-
effector motion. Indeed, Bernstein had partly argued on this basis, postulating that the end-
effector paths were less variable than the joint level trajectories. That comparison is, alas,
difficult to make. End-effector variance is measured in centimeters (squared), joint variance
in radians or degrees (squared). So a direct comparison is meaningless. What is needed is a
shared space in which all variables can be embedded and a shared metric enables
comparisons.
This is what the concept of the uncontrolled manifold achieves (Schöner 1995; Scholz &
Schöner 1999; Latash et al. 2007). The idea is to use joint space as the embedding space in
which all variance is measured. The structure of the variance in joint space is now
interpreted relative to spatial task variables. This is done by defining a subset, the
uncontrolled manifold (UCM), that contains all those combinations of joint angles that are
consistent with one particular end-effector position. There is such an UCM for any position
of the end-effector. The hypothesis is then that, at any given point during the movement,
joint configurations vary primarily within that subset rather than outside of it.
An operational formulation linearizes the UCM, which is typically possible given the limited
range of joint variance. Trials in the same task setting are aligned in time based on
movement initiation and termination. Time is recoded as percent of the mean movement
time. At any given moment of this warped time, the variability in joint space is analyzed by
decomposing it into a component that lies within the UCM and a component that lies

orthogonal to it. The component within the UCM is consistent with no variance at the level
of the spatial task variable. The component orthogonal to the UCM is consistent with a
variable spatial task variable. The UCM hypothesis about the structure of variance is thus
that there is more variance within the UCM than orthogonal to it. In other words, the CNS
preferentially stabilizes those combinations of joint angles that matter for the stabilization of
the spatial task variable than those combinations of joint angles that do not.
This conception has been operationalized and applied to a large set of movement systems
and movement tasks (e.g., Scholz et al. 2000; Tseng et al. 2002). The ideas have been
generalized to the control of isometrical force generation by the fingers of the hand (e.g,
Latash et al., 2002) as well as to the control of muscular activation to achieve particular
location of the center of pressure by a standing person (Krishnamoorthy et al., 2003). In all
cases, strong structure of variance was observed at the level of the elemental variables (joint
angles, individual finger forces, or muscle activations) that was consistent with the preferred
stabilization of spatial task variables (end-effector position, spatial pointing directions,
spatial force vector, or center of pressure coordinate).
Thus, the CNS controls all elemental degrees-of-freedom, including joint angles, digit
forces, and muscle activations. But it does so in a way that reflects the spatial task
constraints by preferentially stabilizing task-relevant directions in joint (or finger or muscle)
space. What does this mean mechanistically? A theoretical model (Martin et al. 2009; see
also Goodman & Latash, 2006 for a related model) proposes that the neuronal command
signals sent to all joints are selectively coupled to the descending motor commands. Only
those combinations of joint angles that affect a spatial task variable are driven by the
descending movement and timing plan while combinations of joint angles orthogonal to
these combinations are free to vary, to accomodate other task constraints, or to receive back-
coupling from the sensed effector position. The model makes use of the conception of an
equilibrium point for every joint-muscle system, which accounts for how initial and terminal
joint configuration differ.
4. Practical applications of the uncontrolled manifold theory

The UCM approach (Scholz & Schöner 1999) has proven to be a powerful tool to
investigate the flexibility/stability aspect of synergies (Latash et al. 2007). This has been
performed in a variety of contexts, including reaching (Reisman & Scholz 2003; Tseng &
Scholz 2005; Tseng et al. 2002), finger force production (Latash et al 2001, 2002; Li et al
1998; Olafsdottir et al. 2007; Shim et al. 2003; Zhang et al. 2008), and postural control
(Freitas et al. 2006; Hsu et al. 2007; Krishnamoorthy et al. 2003, 2004, 2007; Reisman et al
2002a; Scholz et al. 2001) among other tasks. Most studies to date have used the approach to
investigate motor abundance, analyzing the variance of motor elements across repetitions of
a task, or across time in tasks with during relatively steady-state performance. This section
addresses a number of important practical issues that should be considered to apply and
interpret properly the results of this analysis.
Number of data points required

When applying the UCM approach to analyze variance in the space of a set of motor
elements, the number of data points used in the analysis is an important consideration. As
discussed below, for tasks where the configuration of the elements changes substantially
throughout the movement (e.g., joint motion when reaching to a target), analysis needs to be
performed across trials at comparable points in the movement time. Ideally, the more data
points or number of trials in this reaching example, the better. Depending on the nature and
complexity of the experimental conditions, however, the number of trials may be limited by
the ability of subjects to perform a large number of trials per condition. This is particularly

an issue for studies of patients who may be limited in their ability to perform many trials
because of fatigue (Reisman & Scholz 2006). We have performed informal tests of how the
variance structure changes with the number of trials included when analyzing a reaching
task. Figure 3 illustrates the results of one such test where 50 trials of reaching were
performed by a subject with mild hemiparesis resulting from a stroke. Depending on the
portion of the reach examined, UCM analysis performed on different blocks of 10 randomly
selected trials could result in significant quantitative differences in the variance components
within the UCM and orthogonal to the UCM (VUCM and VORT), although the relative
magnitudes of VUCM and VORT did not differ much across blocks. The quantitative
differences diminished with the number of trials included in each block. This is illustrated in
Figure 3.1, which shows the standard deviation of the variance components across the trial
blocks of a given number of trials. The standard deviation stabilizes somewhat with 20 or
more trials included. One might expect the standard deviation to approach zero, but trial-to-
trial variability in performance is not unusual, especially in stroke survivors. This is
certainly not a definitive test, but it suggests loosely that one should include at least 20 trials
for UCM analysis to increase the chance of having a stable estimate of the variance
components. As noted, this is not always possible if multiple conditions are studied and the
experiments involve patients. With fewer trials, however, caution should be used in
interpreting the data across conditions. Fortunately, this is not an issue when the analysis can
be performed across time during the performance of relatively steady-state tasks such as
postural control, where a large number of data points can be included in the analysis (Hsu et
al. 2007).
Relating task space to the space of motor elements

The UCM approach requires being able to relate changes in a hypothesized task variable to
changes in the space of the motor elements (Scholz & Schöner 1999). This requires a formal
model relating how variability of the motor elements affects the task variable. This is usually
not a problem for kinematic studies, where a formal geometric model linking typically can
be written down. For example, changes in the hand’s position can be related directly to small
changes in the current joint configuration through a model of link lengths and trigonometric
functions of the joint angles (Tseng & Scholz 2005). This allows computing the Jacobian of
the system that allows to represent task-pecific variables in the space of elemental variables
(joint angles).
This may not be as trivial for studies involving analysis of the relationship between changes
in muscle activation patterns, as estimated by EMG activity, to a task-relevant variable such
as the center of pressure (CoP) coordinate (Krishnamoorthy et al. 2003, 2004, 2007).
Although a formal model relating EMG signals to such task variables is theoretically
possible, available models are extremely complicated, involve a large number of parameter
estimates that are themselves a potential source of error, and are generally limited to a few
degrees-of-freedom. One way to circumvent this problem has been to use regression
methods to estimate an analog of the Jacobian that relates changes of the task variable to
small changes in the elemental variables. Successful application of such methods has been
performed in studying postural reactions in a variety of contexts (Krishnamoorthy et al.
2003, 2004). The comparability of this experimental approach for estimating the Jacobian to
one based on a formal geometric model has been verified recently in two separate studies of
kinematic redundancy (Freitas & Scholz 2010; Freitas et al. 2010), although future
confirmation in a paradigm relating muscle activation to a task-relevant variable would be
useful.
A related issue is the determination of the elemental variables. In studies relating EMG
activity to kinetic variables such as the CoP, it is typically not possible to record activity
from every muscle involved in the task. It has also been accepted that the brain does not

specify activation levels of each muscle separately. Hence, such studies typically assume
that muscles are grouped and the brain uses a few variables to modify in parallel the
activation levels within each group. Nonetheless, conclusions based on such studies require
caution and the more of the motor elements that can be included, the better.
In addition, including motor elements that have little or no effect on the task being studied
will artificially inflate the variance estimates, especially VUCM. For example, depending on
the context and experimental question, spurious conclusions could result by including
rotational degrees-of-freedom about the long axes of the humerus and forearm in a study of
planar reaching. Consider, for example, reaching with the hand, from an initial position
where the forearm rests on a table, elbow flexed to 90°, and hand pointing forward, to a
target located at a fixed distance immediately above the hand. These rotational degrees-of-
freedom have little or no effect on the vertical position of the hand along its path to the
target. Therefore, any variation in these angles will contribute only to VUCM, although this
additional variance may have no relationship to the task. Caution should be used, therefore,
when determining what degrees-of-freedom to included. At the very least, this issue should
be considered when interpreting the experimental results.
Identification of task variables

UCM analysis is related to other methods such as principal component analysis (PCA),
where the structure of variance among a combination of motor elements is analyzed. PCA
may be considered an objective approach because it is performed without direct connection

to a hypothesis about what task variable might be generating the observed structure in the
space of the motor elements. In contrast, for UCM analysis, the structure of variance in the
motor element space is always related to a hypothesized task variable through a formal
model linking the two spaces. The decision about what task variables to investigate certainly
has a subjective element. Including the analysis of a number of relevant task variables may
be important if one is investigating questions about what the nervous system “cares” most
about in a given task.
Thus, application of the UCM method of analysis is useful only if one has an explicit
experimental question to address related to the structure of variance of the elemental
variables in relation to changes in a task variable and/or when performing a task under
different constraints. For a given task, one has to hypothesize what task-level variables are
“most important” for the nervous system. Indeed, by analyzing how the variance of the
elemental variables is structured relative to different task variables, it is possible to
differentiate among the relative importance of such variables for the task being investigated.
In some cases, it may even be useful to differentiate among different dimensions of the same
task variable. For example, in studies of the sit-to-stand action under different task
constraints, it was shown that joint variance was structured mostly to stabilize the horizontal
position of the center of mass (CoM) in contrast to the vertical CoM position (Scholz &
Schöner 1999). Stability of the vertical CoM position through the use of flexible
combinations of joint postures was more evident, however, when sitting down, where vision
of the seat was limited (Reisman et al. 2002b).
An additional caution is required when interpreting differences in the variance components

after UCM analysis among different task variables if the model relating task space to
elemental space is very different in the two cases. For example, in kinematic studies, the
geometry of the system will influence the degree to which variance in joint space affects
variance of a task variable. Because the models relating different task variables to joint
variance may be quite different, it would be important to examine the extent to which VORT
actually affects the task variable in both cases. That is, the same amount of VORT may affect

one variable more than the other. This relationship can be identified by examining the
singular values of the Jacobian in each case, as demonstrated by Freitas et al. (2010).
Indices of the strength of a synergy

Different methods of combining the two variance components obtained from UCM analysis,
i.e. VUCM and VORT, have been used in previous studies to provide an index of the strength
of a synergy stabilizing the task variable for which the analysis was run. Initially this
involved examining the ratio of the two components. More recent investigations have used
the relative difference between the components (Gorniak et al. 2007, 2009b), in part because
the ratio has a floor effect and is not normally distributed. The idea is that a larger value of
this measure indicates a stronger synergy. Concluding that one person or task exhibits a
stronger synergy than another may be valid depending on the values of the individual
variance components. Thus, the interpretation of such a measure requires caution.
Consider as an example emergency responders pressing with the index and middle fingers to
generate chest compressions in an infant having cardiac arrest. In such an incident, one
needs to produce a stable force that is large enough to restore the heart beat while
minimizing injury to the infant. This could be achieved, in principle, by pressing with a
fixed amount of force by each finger on successive repetitions. However, biological systems
are variable and it is difficult to generate the identical amount of force by each finger on
each compression. If one were to plot the two finger forces against each other across all
repetitions (Figure 4A), then this independent control strategy would yield a circular cloud
of points, the diameter of which would indicate how successful the strategy was in
stabilizing the total force. Notice, however, that, because the forces act independently of
each other, the total force may frequently fall below the desired compressive force level, or
even injure the infant’s chest because the total force is too high. A more desirable solution
would be to co-vary the finger forces through a synergy that links their output to the task
requirements. Plotting the results of such a control strategy would result in a cigar shaped
cloud of points with a negative slope, indicating that when one finger’s force increases too
much, the other finger’s force is reduced by a comparable amount and vice versa to preserve
the total force. Coupling the natural variability of the fingers’ forces in this way may reflect
the fact that the nervous system has available to it flexible ways to combine the motor
elements when needed. Perfect compensation between the fingers would result in all data
points falling along the diagonal line in Figure 4B. If this compensation is not perfect, as
likely is the case, the cloud of points will become fatter and more cigar-shaped. This
fattening of the diagonal cloud of finger forces indicates greater VORT when performing
UCM analysis. Comparing results across two such emergency responders, if the repetitive
force structure was fatter for one of them with no comparable difference in the diagonal
length of the two clouds of finger forces, i.e. no difference in VUCM, then one certainly
could conclude that one responder exhibits a stronger finger force synergy than the other.
It is problematic to make such a conclusion without examining the magnitude of the

individual components, however. Consider the difference in the two clouds of points in
Figure 4B. Although the cloud of points in the right panel of this figure (responder #2) is
fatter than in the left panel (responder #1) (i.e. responder #2 has larger VORT), the force
variance also extends substantially further along the diagonal representing the UCM for
responder #2 (much larger VUCM). The typically used synergy index in this case may
indicate a stronger synergy for responder #2. However, only VORT affects the task variable’s
variance. Therefore, responder #1‘s performance would be associated with less task
variability. It would be odd to conclude that this individual exhibits a weaker synergy if the
notion of a functional synergy is associated with successful task performance. A similar
result was reported by Reisman and Scholz (2003) in a study comparing reaching between
healthy control subjects and persons with mild hemiparesis who had suffered a stroke. The

latter individuals had significantly greater overall joint variance that was distributed across
both subspaces of joint space compared to the control subjects. Thus, a similar synergy
index would have suggested an equally strong synergy in these individuals as the controls
despite having significantly higher VORT and much greater task error.
Effects of practice and motor disorders studied with the UCM method
Several studies have demonstrated that the method of the UCM hypothesis can be used to
explore changes in motor coordination that accompany movement disorders, atypical
development, aging, and practice. In particular, the mentioned study of Reisman and Scholz
(2003) of hemiparetic subjects after stroke documented similar indices of multi-joint
synergy (the relative amounts of VUCM and VORT in the total joint configuration variance)
stabilizing the endpoint trajectory during reaching movements by the affected arm
(contralateral to the stroke site) of the patients and similar movements performed by healthy
subjects. This unexpected result underscored the importance of exploring both variance
indices, VUCM and VORT, in clinical studies.
Persons with Down syndrome show low indices of multi-finger synergy in multi-finger
accurate force production tasks (Latash et al. 2002b; Scholz et al. 2003). Then tend to show
positive co-variation of individual finger forces as if they used the hand as a fork turned
upside down and scaled efforts of individual fingers in parallel. However, a relatively brief
training session led to a significant improvement in accuracy of their performance
accompanied by an increase in the synergy index and more flexible use of individual fingers
of the hand.
Elderly persons are known to show a decline in the hand function over a range of everyday
tasks. This decline is accompanied by an impaired coordination of the fingers reflected in
lower indices of synergies stabilizing total force and total moment of force produced by the
fingers (Shinohara et al. 2004; Olafsdottir et al. 2007). A six-week strength training protocol
resulted in improved accuracy of performance in force production tasks which correlated
with an increase in the synergy index (Olafsdottir et al. 2008). So, this study has shown that
synergy index may be a predictive factor of accurate performance.
Several studies of the effects of practice on synergy indices in young, healthy persons
(Latash et al. 2003; Kang et al. 2004; reviewed in Latash 2010) have suggested a two-stage
process of performance improvement with practice. These stages correspond to (1)
discovery and strengthening of motor synergies stabilizing salient performance variable(s);
and (2) their weakening when other aspects of motor performance are optimized. Within the
equilibrium-point hypothesis, the first stage may be viewed as consisting of two step, the
elaboration of an adequate referent configuration trajectory and the elaboration of multi-joint
(multi-muscle) synergies stabilizing the referent configuration trajectory. Both steps are
expected to lead to more variance in the space of elemental variables that is compatible with
a desired time profile of the salient performance variable (VUCM). Adjusting control to other
aspects of performance during the second stage (for example, esthetics, energy expenditure,
time, fatigue, etc.) may lead to a drop in VUCM.
5. Concluding Comments
A few recent studies attempted to link the equilibrium-point hypothesis, the principle of
abundance, and the uncontrolled manifold hypothesis into a single, coherent scheme on how
natural voluntary movements are produced. These involved a theoretical study by Martin et
al. (2009) and two experimental studies (Gorniak et al. 2009b; Latash et al. 2010). These
first attempts show that creating such a coherent scheme based on the reviewed hypothesis is
feasible. Moreover, this seems to be the only currently available direction of research that

promises success in achieving the main goal of motor control as stated in the Introduction:
To create a formal description, operating with exactly defined variables, of the physical and
physiological processes that make coordinated voluntary movements possible.
References
Asatryan DG, Feldman AG. Functional tuning of nervous system with control of movement or
maintenance of a steady posture. I. Mechanographic analysis of the work of the joint on execution
of a postural task. Biophysics 1965;10:925–935.
Bernstein, NA. The coordination and regulation of movements. London: Pergamon Press; 1967.
Bigland B, Lippold OCJ. The relation between force, velocity and integrated electrical activity in
human muscles. J Physiol 1954;123:214–224. [PubMed: 13131257]
Erlhagen W, Schöner G. Dynamic field theory of movement preparation. Psychol Rev 2002;109:545–
572. [PubMed: 12088245]
Feldman AG. Functional tuning of the nervous system with control of movement or maintenance of a
steady posture. II. Controllable parameters of the muscle. Biophysics 1966;11:565–578.
Feldman AG. Once more on the equilibrium-point hypothesis (λ-model) for motor control. J Mot
Behav 1986;18:17–54. [PubMed: 15136283]
Feldman, AG. Space and time in the context of the equilibrium point theory. Wiley Cognitive
Sciences; In press
Feldman, AG.; Adamovich, SV.; Ostry, Dl; Flanagan, IR. The origin of electromyograms -
explanations based on the equilibrium point hypothesis. In: Winters, IM.; Woo, SL-Y., editors.
Multiple Muscle Systems. Berlin, Heidelberg, New York: Springer; 1990. p. 195-312.
Feldman AG, Goussev V, Sangole A, Levin MF. Threshold position control and the principle of
minimal interaction in motor actions. Prog Brain Res 2007;165:267–281. [PubMed: 17925252]
Freitas SM, Duarte M, Latash ML. Two kinematic synergies in voluntary whole-body movements
during standing. J Neurophysiol 2006;95:636–645. [PubMed: 16267118]
Freitas SM, Scholz JP. A comparison of methods for identifying the Jacobian for uncontrolled
manifold variance analysis. J Biomech 2010;43:775–777. [PubMed: 19922938]
Freitas SM, Scholz JP, Latash ML. Analyses of joint variance related to voluntary whole-body
movements performed in standing. J Neurosci Methods 2010;188:89–96. [PubMed: 20105441]
Gelfand IM, Latash ML. On the problem of adequate language in motor control. Motor Control
1998;2:306–313. [PubMed: 9758883]
Gelfand, IM.; Latash, ML. On the problem of adequate language in biology. In: Latash, ML., editor.
Progress in Motor Control. vol. 2: Structure-Function Relations in Voluntary Movement. Urbana,
IL: Human Kinetics; 2002. p. 209-228.
Georgopoulos AP. Population activity in the control of movement. Int Rev Neurobiol 1994;37:103–
119. [PubMed: 7883475]
Georgopoulos AP, Ashe J, Smyrnis N, Taira M. The motor cortex and the coding of force. Science
1992;245:1692–1695. [PubMed: 1609282]

Georgopoulos AP, Kalaska JF, Caminiti R, Massey JT. On the relations between the direction of two-
dimensional arm movements and cell discharge in primate motor cortex. J Neurosci 1982;2:1527–
1537. [PubMed: 7143039]
Goodman SR, Latash ML. Feed-forward control of a redundant motor system. Biol Cybern
2006;95:271–280. [PubMed: 16838148]
Gorniak SL, Zatsiorsky VM, Latash ML. Hierarchies of synergies: an example of two-hand, multi-
finger tasks. Exp Brain Res 2007;179:167–180. [PubMed: 17103206]
Gorniak SL, Feldman AG, Latash ML. Joint coordination during bimanual transport of real and
imaginary objects. Neurosci Lett 2009a;456:80–84. [PubMed: 19429138]
Gorniak SL, Zatsiorsky VM, Latash ML. Hierarchical control of static prehension: II. Multi-digit
synergies. Exp Brain Res 2009b;194:1–15. [PubMed: 19048236]
Grossberg S, Pribe C, Cohen MA. Neural control of interlimb oscillations. I. Human bimanual
coordination. Biol Cybern 1997;77:131–140. [PubMed: 9323862]

Hsu WL, Scholz JP, Schöner G, Jeka JJ, Kiemel T. Control and estimation of posture during quiet
stance depends on multijoint coordination. J Neurophysiol 2007;97:3024–3035. [PubMed:
17314243]
Ivry, RB.; Diedrichsen, J.; Spencer, R.; Hazeltine, E.; Semjen, A. A cognitive neuroscience
perspective on bimanual coordination and interference. In: Swinnen, SP.; Duysens, J., editors.
Neuro-Behavioral Determinants of Interlimb Coordination — An interdisciplinary aproach.
Dordrecht, The Netherlands: Kluwer Academic Publishers; 2004. p. 259-295.
Jobin A, Levin MF. Regulation of stretch reflex threshold in elbow flexors in children with cerebral
palsy: a new measure of spasticity. Develop Med Child Neurol 2000;42:531–540. [PubMed:
10981931]
Kang N, Shinohara M, Zatsiorsky VM, Latash ML. Learning multi-finger synergies: An uncontrolled
manifold analysis. Exp Brain Res 2004;157:336–350. [PubMed: 15042264]
Krishnamoorthy V, Latash ML, Scholz JP, Zatsiorsky VM. Muscle synergies during shifts of the
center of pressure by standing persons. Exp Brain Res 2003;152:281–292. [PubMed: 12904934]
Krishnamoorthy V, Latash ML, Scholz JP, Zatsiorsky VM. Muscle modes during shifts of the center
of pressure by standing persons: effect of instability and additional support. Exp Brain Res
2004;157:18–31. [PubMed: 14985897]
Krishnamoorthy V, Scholz JP, Latash ML. The use of flexible arm muscle synergies to perform an
isometric stabilization task. Clin Neurophysiol 2007;118:525–537. [PubMed: 17204456]
Lacquaniti F. Central representations of human limb movement as revealed by studies of drawing and
handwriting. Trends Neurosci 1989;12:287–292. [PubMed: 2475946]
Latash ML. Stages in learning motor synergies: A view based on the equilibrium-point hypothesis.
Hum Move Sci. 2010 (in press).
Latash ML, Friedman J, Kim SW, Feldman AG, Zatsiorsky VM. Prehension synergies and control
with referent hand configurations. Exp Brain Res 2010;202:213–229. [PubMed: 20033397]
Latash ML, Kang N, Patterson D. Finger coordination in persons with Down syndrome: Atypical
patterns of coordination and the effects of practice. Exp Brain Res 2002b;146:345–355. [PubMed:
12232691]
Latash ML, Scholz JF, Danion F, Schöner G. Structure of motor variability in marginally redundant
multifinger force production tasks. Exp Brain Res 2001;141:153–165. [PubMed: 11713627]
Latash ML, Scholz JF, Danion F, Schöner G. Finger coordination during discrete and oscillatory force
production tasks. Exp Brain Res 2002;146:419–432. [PubMed: 12355270]
Latash ML, Scholz JP, Schöner G. Toward a new theory of motor synergies. Motor Control
2007;11:276–308. [PubMed: 17715460]
Latash ML, Yarrow K, Rothwell JC. Changes in finger coordination and responses to single pulse
TMS of motor cortex during practice of a multi-finger force production task. Exp Brain Res
2003;151:60–71. [PubMed: 12740728]
Levin MF. Sensorimotor deficits in patients with central nervous system lesions: explanations based on
the λ model of motor control. Hum Move Sci 2000;19:107–137.
Levin MF, Feldman AG. The role of stretch reflex threshold regulation in normal and impaired motor
control. Brain Res 1994;657:23–30. [PubMed: 7820623]
Levin MF, Selles RW, Verheul MHG, Meijer OG. Deficits in the coordination of agonist and
antagonist muscles in stroke patients: implications for normal motor control. Brain Res
2000;853:352–369. [PubMed: 10640634]
Li ZM, Latash ML, Zatsiorsky VM. Force sharing among fingers as a model of the redundancy
problem. Exp Brain Res 1998;119:276–286. [PubMed: 9551828]
Martin V, Scholz JP, Schöner G. Redundancy, self-motion and motor control. Neural Comput
2009;21:1371–1414. [PubMed: 19718817]
Mechsner F, Kerzel D, Knoblich G, Prinz W. Perceptual basis of bimanual coordination. Nature
2001;414:69–73. [PubMed: 11689944]
Morasso P. Spatial control of arm movements. Exp Brain Res 1981;42:223–227. [PubMed: 7262217]

Olafsdottir HB, Zatsiorsky VM, Latash ML. The effects of strength training on finger strength and
hand dexterity in healthy elderly individuals. J Appl Physiol 2008;105:1166–1178. [PubMed:
18687981]
Olafsdottir H, Zhang W, Zatsiorsky VM, Latash ML. Age-related changes in multifinger synergies in
accurate moment of force production tasks. J Appl Physiol 2007;102:1490–1501. [PubMed:
17204576]
Reisman DS, Scholz JP. Aspects of joint coordination are preserved during pointing in persons with
post-stroke hemiparesis. Brain 2003;126:2510–2527. [PubMed: 12958080]
Reisman DS, Scholz JP. Workspace location influences joint coordination during reaching in post-
stroke hemiparesis. Exp Brain Res 2006;170:265–276. [PubMed: 16328275]
Reisman DS, Scholz JP, Schoner G. Coordination underlying the control of whole body momentum
during sit-to-stand. Gait and Posture 2002a;15:45–55. [PubMed: 11809580]
Reisman DS, Scholz JP, Schoner G. Differential joint coordination in the tasks of standing up and
sitting down. J Electromyogr Kinesiol 2002b;12:493–505. [PubMed: 12435546]
Rosenbaum DA. Human movement initiation: Specification of aim, direction, and extent. J Exp
Psychol: General 1980;109:444–474.
Scholz JP, Kang N, Patterson D, Latash ML. Uncontrolled manifold analysis of single trials during
multi-finger force production by persons with and without Down syndrome. Exp Brain Res
2003;153:45–58. [PubMed: 12928761]
Scholz JP, Reisman D, Schoner G. Effects of varying task constraints on solutions to joint coordination
in a sit-to-stand task. Exp Brain Res 2001;141:485–500. [PubMed: 11810142]
Scholz JP, Schöner G. The uncontrolled manifold concept: identifying control variables for a
functional task. Exp Brain Res 1999;126:289–306. [PubMed: 10382616]
Scholz JP, Schoner G, Hsu WL, Jeka JJ, Horak F, Martin V. Motor equivalent control of the center of
mass in response to support surface perturbations. Exp Brain Res 2007;180:163–179. [PubMed:
17256165]
Schöner G. Recent developments and problems in human movement science and their conceptual
implications. Ecol Psychol 1995;7:291–314.
Schöner G, Kelso JAS. Dynamic pattern generation in behavioral and neural systems. Science
1988;239:1513–1520. [PubMed: 3281253]
Scott SH, Kalaska JF. Changes in motor cortex activity during reaching movements with similar hand
paths but different arm postures. J Neurophysiol 1995;73:2563–2567. [PubMed: 7666162]
Shim, v; Latash, ML.; Zatsiorsky, VM. Prehension synergies: trial-to-trial variability and hierarchical
organization of stable performance. Exp Brain Res 2003;152:173–184. [PubMed: 12898101]
Shinohara M, Scholz JP, Zatsiorsky VM, Latash ML. Finger interaction during accurate multi-finger
force production tasks in young and elderly persons. Exp Brain Res 2004;156:282–292. [PubMed:
14985892]
Thilmann AF, Fellows SJ, Garms E. The mechanisms of spastic muscle hypertonus. Variation in reflex
gain over the time course of spasticity. Brain 1991;114:233–244. [PubMed: 1998884]
Tseng YW, Scholz JP. The effect of workspace on the use of motor abundance. Motor Control
2005;9:75–100. [PubMed: 15784951]
Tseng YW, Scholz JP, Schöner G. Goal-equivalent joint coordination in pointing: affect of vision and
arm dominance. Motor Control 2002;6:183–207. [PubMed: 12122226]
Won J, Hogan N. Stability properties of human reaching movements. Exp Brain Res 1995;107:125–
136. [PubMed: 8751070]
Zatsiorsky, VM. Kinematics of human motion. Urbana, IL: Human Kinetics; 1998.
Zhang W, Scholz JP, Zatsiorsky VM, Latash ML. What do synergies do? Effects of secondary
constraints on multidigit synergies in accurate force-production tasks. J Neurophysiol
2008;99:500–513. [PubMed: 18046000]

Figure 1.
Three experimental procedures leading to the notion that central shifts in the threshold
position (R) underlie voluntary motor actions. Procedures include: involuntary motor action
– unloading reflex – resulting from sudden unloading of the forearm to different final
equilibrium points (EPs, open circles) of pre-loaded elbow flexors from an initial EP (filled
circle, a); voluntary motor action when the subject changed the initial EP a to EP b; stretch
of the fully relaxed elbow flexors (dashed curve). For each unloading characteristic (solid
curves), the tonic EMG activity decreased with the decreasing residual torque (vertical lines)
and became zero when the unloading characteristic reached the characteristic of the passive
muscle at position R, called the threshold elbow position. For different unloading
characteristics, threshold positions were different (ΔR). Therefore, the intentional motor
action associated with the transition from EP a to EP b (or from one unloading characteristic
to another), was accomplished by a change in the threshold angle R. Horizontal line at
bottom of figure shows that the range of regulation of R (R− to R+) is greater than the
biomechanical range of the joint.

Figure 2.
Schematic illustration of tonic stretch reflex thresholds (TSRT) determined by extrapolation
of a linear regression line through dynamic stretch reflex thresholds (DSRTs). DSRTs are
evoked by stretching elbow flexors at different velocities. In healthy subjects, DRSTs can
only be evoked by high velocity stretches and the TSRT lies beyond the biomechanical
range of the joint (TSRTh). In patients with spasticity, DSRTs occur earlier (at smaller joint
angles), and the TSRTs lies within the biomechanical range of the joint.

Figure 3.
Standard deviation of variance components, VUCM and VORT across five blocks of 10, 15,
20, or 25 trials. Variance is computed as rad2 per dimension of each subspace. Trials were
randomly selected from a group of 50 trials collected for this subject. The largest reduction
in standard deviation occurs between 10 and 15 trials.

Figure 4.
In a two-finger force production task, independent control of each finger leads to
uncorrelated variance across repetitions (forming a circle, stars). Negative covariation of
finger forces decreases the total force variance. B. Comparison of the VUCM and VORT
components of variance in two hypothetical first responders performing infant CPR. The
often used synergy index (e.g., [VUCM − VORT] / [(VUCM + VORT)/2]) would result in a
larger value for responder #2 than responder #1, despite the fact that VORT, the only
variance component that can lead to task level variance, is smaller for responder #1. The
comparison emphasizes the need to examine each component of variance as well as any
combined index when making conclusions about the usefulness of a synergy.
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Motor Control Theories and Their Applications

Article in Medicina (Kaunas, Lithuania) · January 2010

Mindy F Levin John Scholz

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Medicina (Kaunas). 2010 ; 46(6): 382–392.

Motor Control Theories and Their Applications

Mark L. Latash1, Mindy F. Levin2, John P. Scholz3, and Gregor Schöner4

2Department of Physical Therapy, McGill University, Montreal, Canada

The first of these theories is the equilibrium-point hypothesis (referent configuration

Medicina (Kaunas). Author manuscript; available in PMC 2011 January 9.

In a series of experiments on the elbow joint in humans combining involuntary movement

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3. The concept of the uncontrolled manifold

Many different mechanism seem to be involved in controlling the timing of movements,

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In a task-effector system that has abundant solutions, or is redundant in more conventional

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4. Practical applications of the uncontrolled manifold theory

Number of data points required

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Relating task space to the space of motor elements

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Identification of task variables

may be considered an objective approach because it is performed without direct connection

An additional caution is required when interpreting differences in the variance components

Medicina (Kaunas). Author manuscript; available in PMC 2011 January 9.

Indices of the strength of a synergy

It is problematic to make such a conclusion without examining the magnitude of the

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Medicina (Kaunas). Author manuscript; available in PMC 2011 January 9.

1992;245:1692–1695. [PubMed: 1609282]

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Medicina (Kaunas). Author manuscript; available in PMC 2011 January 9.

Medicina (Kaunas). Author manuscript; available in PMC 2011 January 9.

Medicina (Kaunas). Author manuscript; available in PMC 2011 January 9.

Medicina (Kaunas). Author manuscript; available in PMC 2011 January 9.

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Medicina (Kaunas). Author manuscript; available in PMC 2011 January 9.

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