Motor Control
These studies gave further evidence for the validity of
the model (for details see Gubler and Bischof 1991,
Gubler et al. 1994).
4. Future Directions
While the functioning of the security and the arousal
systems is sufficiently supported by empirical evidence,
there are still several open questions for future
research. Although the existing data also support the
assumptions made about the autonomy system, the
causal link of the autonomy claim to dependency and
enterprise still has to be established. Since the Zurich
Model was originally conceived from the point of view
of a child, more adult motives such as altruism have to
be included as well. Furthermore, the study of the
developmental course taken by the different reference
variables or motives also has to be extended to older
age groups in order to get a model explaining the
motivational development of the whole life span.
See also: Academic Achievement Motivation, Devel-
opment of; Intrinsic Motivation, Psychology of;
Motivation and Actions, Psychology of; Motivation:
History of the Concept; Motivation, Learning, and
Instruction
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of Attachment: A Psychological Study of the Strange Situation.
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Bischof N 1975 A systems approach towards the functional
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Bischof N 1985 Das RaW tsel OedipusDie biologischen Wurzeln
des Urkonfliktes on IntimitaW t und Autonomie. Piper, Munich,
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Bischof N 1993 Untersuchungen zur Systemanalyse der sozialen
Motivation I: Die Regulation der sozialen DistanzVon der
Feldtheorie zur Systemtheorie. Zeitschrift fuW r Psychologie
201: 543
Bischof N 1995 Struktur und Bedeutung. Eine EinfuW hrung in die
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Bowlby J 1969 Attachment and Loss: Vol. 1. Attachment. Basic
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95132
Hyland M E 1988 Motivational control theory: An integrative
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Klein H J 1989 An integrated control theory model of work
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M. E. Schneider
Motor Control
The rich varieties of human and non-human animal
movement are the final product of motor mechanisms
within the nervous system and the muscles they
innervate. The selective contraction and relaxation of
these muscle groups support the organism (posture),
mediate internal functions such as heartbeat, breath-
ing, and peristalsis of the digestive system, and propel
the creature through its external world. The motor
neurons typically work in concert with interneurons
and sensory mechanisms that are orchestrated to-
gether through intricate patterns of activity. The
motor control pathways in whole organism actions
read-out these patterns from many parts of the brain
10125
Motor Control

and its contact with the sensory world. It has been

argued (Lashley, 1951 Sperry 1952) that movement,
and thus motor control, is the most fundamental
property of nervous systems. Without movement there
is no behavior as the term is normally defined.
Movement also provides overt and objective measures
of the rich symphonies of behavior we see in the
natural world.

1. Commonalities and Specializations in

Mechanism
Motor control in all vertebrate species (animals with
backbones) operates from a similar ground plan, but
with important differences in detail. For all vertebrates
spinal and brain stem mechanisms mediate a number
of basic forms of movement, such as locomotion and
postural adjustment (see Behaior, Hierarchical Organ-
ization of; Eye Moement, Control of (Oculomotor
Control)). In mammals, and especially in humans,
higher cortical areas along with the basal ganglia and
cerebellum contribute importantly to the acquisition,
planning, flexibility, and fine-tuning of movement
sequences (Fig. 1). Major properties of these structures
are discussed below. In invertebrates (animals without
backbones) the basic building blocks of motor control
are diverse and distinct from those found in vertebrate
species, yet there is a remarkable convergence in terms
of overall solutions to motor control. For example,
relatively small groups of neurons, and in some species
single cells, generate rhythms of an endogenous
(intrinsic to the cells) rhythms without the need for
molding by afferent influences. These central pattern
generators were first documented clearly in inver-
tebrates, and then determined for vertebrates (Fentress Figure 1
1976, Pearson 1993; see Comparatie Neuroscience). The motor systems have three levels of controlthe
In the intact animal these patterns of spontaneous spinal cord, brain stem, and forebrainorganized both
activity are typically modulated by other neural and serially and in parallel. Source: Kandel et al. 2000
endocrine influences, however.
Most investigators today believe that many of the tion in Virtual Space: Psychological and Neural
most important questions of motor control rest at the Aspects; Naigation in Spatial Enironments, Neural
interface between intrinsic properties of neurons and Basis of; Neural Representations of Intended Moement
their sensitivity to extrinsic factors. Motor commands in Motor Cortex).
may remain latent until triggered by descending
influences, rather as dropping a needle onto a phon-
ograph record triggers patterns of sound programmed 2. Early Models and Current Perspecties
within the chosen record. The analogy to music turns
out to be a useful one, for it reminds us that we wish to Early models of vertebrate movement stressed in-
account for the notes (basic units) of movement, the dividual reflexes and reflex chains as the basic building
construction of chords of simultaneously activated blocks in the organization of action patterns
motor elements, melodies in time, rhythmic principles, (Sherrington 1906). Spinal reflexes operate through
volume (intensity) and timbre (fine tuning), and even relatively simple chains of neurons, where sensory and
possible transpositions from one moment to the next. interneurons link up with motor neurons that in turn
Questions of movements guided toward specific feat- project to a select group of muscles. Interest in reflex
ures of the environment are also of obvious impo- behavior (such as the well-known knee jerk reflex
rtance, and are discussed in other chapters (see elicited by your local physician) remains intense, and
Behaior, Hierarchical Organization of; Neural Repre- reflexes are clearly essential for the production of
sentations of Direction (Head Direction Cells); Naiga- posture, basic movements, and defensive reactions.

10126

Their speed and reliability of production reflect what
are often called automated properties of motor
control, that is movement profiles that are rapid,
essentially ballistic, and not mediated by conscious
processes. Even these reflexes can be modulated by
higher neural activities, however. As in nearly all
aspects of neural function it is thus a matter of looking
at intrinsic circuits together with extrinsic influences.
These occur at many levels of neural control.
2.1 Moement Units and Their Limits
One of the basic challenges in the study of motor
control is the dissection of fundamental movement
units. In this terminology a unit is a relatively invariant
pattern of muscular contractions that are typically
elicited together. Reference to one of the most complex
forms of human movement control, speech, is illustra-
tive (Neville 1995). When we speak we articulate a
broadly invariant set of phonemes (basic sound
patterns) in sequence. This allows for the articulation
of speech to be relatively thought free in normal
discourse, and allows the listener to decode the
message through a complex array of perceptual and
higher-order cognitive processes (see Syntactic Aspects
of Language, Neural Basis of; Motor Control; Sign
Language: Psychological and Neural Aspects; Lexical
Processes (Word Knowledge): Psychological and
Neural Aspects; Speech Production, Neural Basis of).
The example is useful in that in the smooth flow of
speech individual phonemes are often co-articulated
with the fine details of one sound unit reflecting the
production of preceding and subsequent sound pat-
terns. So even here the intrinsic properties of a speech
unit can be sensitive to extrinsic influences as defined
by other speech units. This is what gives speech its
smooth and flowing nature (as opposed, for example,
to most computer generated speech).
Thus, in some sense movement and their motor
control units are abstractions that frequently blur at
the edges, and which can be defined from a number of
complementary perspectives (Golani 1992). In com-
plex forms of movement, such as playing a sport or
musical instrument, or dancing with a moving partner,
many neural pathways are orchestrated together in
dazzling patterns that are widely distributed, serially
ordered, and parallel in their operations. It is for this
reason that the very distinction between motor control
systems and other properties of the nervous system are
often difficult to untangle (see Vision for Action: Neural
Mechanisms; Cognitie Control (Executie Functions):
Role of Prefrontal Cortex).
2.2 Moement Notation
A number of movement notation systems have helped
clarify the organization of motor patterns at many
Motor Control
different levels. For example, the EshkolWachman
movement notation system (Golani 1992) has helped
investigators separate specific parameters of move-
ment, such as movements in a plane (flat surface) and
those that involve rotations or conical movements
around a single limb. These notational systems,
derived from human choreography, also remind us
that movement is in its essence both relational and
dynamic. Its description also depends upon the frame
of reference adopted. For example, we can move our
bodies with respect to internal coordinates and yet
maintain a fixed relationship to external events, such
as a moving social partner. Often the units of control
involve multiple muscular systems, such as when we
produce relatively invariant motions during the final
phases of feeding or brushing our teeth via com-
plementary adjustments of individual muscular acts
(such as those involved in arm movements or turning
the head).
2.3 Dynamic Systems
Many workers today have chosen to supplement the
language of motor units and experimentally isolated
mechanisms with insights derived from dynamic
systems theory (Kelso 1995, Thelen 1985). These
approaches remind us that behavior (movement, and
thus motor control) is fundamentally imbedded in
time, and involves the cooperation of a rich multitude
of processes whose boundaries are often difficult to
define. Further, there has been an increased appr-
eciation for the flexibility of underlying mechanism
operations. Even in invertebrates circuits that con-
tribute to a given movement may be shared and
change their internal properties from one occasion of
expression to the next (Marder and Calabrese 1996).
This remolding of circuit properties adds impor-
tantly to the challenges of deep understanding of the
processes of motor control.
3. Diersity and Plasticity in Human Moement
The human species is especially remarkable in its
ability to master skills of an enormous diversity. One
needs to reflect just for a moment on actions such as
speech, sports, dance, driving an automobile, or
musical performance to appreciate this fact. Indeed
while each species is the master of its own behavioral
niche, it is in humans that we find the most rich variety
of forms of expression. Our nervous systems, through-
out much of our lives, are remarkably plastic, meaning
that they can be refined and remolded into an often
dizzying array of specialized talents (see Neural Plas-
ticity in Auditory Cortex; Neural Plasticity;
Dysexecutie Syndromes; Neural Plasticity in Visual
Cortex).
10127
Motor Control
We have all had the experience that in the early
learning of a new motor skill our attention is fully
absorbed. Later we produce the actions in an auto-
mated manner, without any conscious awareness.
When we seek to modify this skill, or apply it to a new
context, the executie parts of our nervous system are
recaptured, with thought and planning once again
playing a major role. It is now well established that
different neural circuits are employed in the early and
late stages of skill acquisition (Posner et al. 2001).
3.1 Basic Pathways
It remains possible to separate particular neural
pathways that contribute to different properties of
movement. The human motor system can be visualized
in part as being composed of descending pathways
derived from the cerebral cortex and brainstem that
are fundamental to the suprasegmental control of
movement. The term segmental refers to spinal
segments that control different muscle groups in an
orderly way. Suprasegmental thus refers to actions
that incorporate muscles of different extremities such
as the head, arms, trunk, and legs. The basal ganglia
and cerebellum contribute to the planning, sequencing
and fine-tuning of movement. The basal ganglia are
embedded deep within the forebrain and also include
midbrain pathways. These pathways do not connect
directly to motor pathways, but exert important
indirect influences that are mediated through cortical
and thalamic loops. Disorders of the basal ganglia
greatly compromise effective movement, such as seen
in Parkinsons Disease with its characteristic tremor
and poverty of movement, and Huntingtons Disease
with its characteristic choreoform (sporadic and bal-
listic) movements. Each of these diseases is also
marked by cognitive (thought, planning) deficits as the
disease progresses, highlighting once again how motor
systems are intimately linked with other neural
functions.
The cerebellum has multiple sensory and motor
functions, among which the timing and fine-tuning of
motor control are among the most apparent. It is
now known that lesions of the lateral cerebellum also
compromise the perception of temporal events (Ivry
1996), thus illustrating yet again how distinctions
between motor and other neural functions can be
misleading. Individual circuits, and even individual
neurons, can mediate a variety of functions (see
Cerebellum: Associatie Learning; Cerebellum: Cog-
nitie Functions).
The spinal cord, which receives descending inputs
from the brain, is neatly organized with anatomically
arranged segments influencing and controlling actions
of the upper and lower body. These richly inter-
connected pathways work together with the neuro-
muscular system which is also composed of nuclei of
the motor cranial nerves and the muscles that they
innervate.
10128
At every level of organization major afferent path-
ways project to the motor control mechanisms, but
their influence is a function of such factors as speed of
movement and the extent to which movement patterns
have become automated (as in well practiced tasks).
Often the influence of afferent pathways is relatively
small in well practiced and simple automated move-
ments, but sensitivity to sensory influences can attain
major importance when actions have to be adjusted,
such as in the confrontation with unexpected cir-
cumstances.
Activities from extrinsic sources are gated to varying
degrees as a function of the demands placed upon the
organism at the moment. Routine activities run more
or less autonomously, but are able to be corrected if
circumstances change. In this way the production of
movement sequences can be both automated and
responsive to novel circumstances. There are many
known human motor diseases that can selectively
affect both the automated performance of movement
and the ability of the individual to adjust to novel
situations (Linas 2001). A skillful neurologist can,
by examining detailed properties of movement per-
formance, often pinpoint parts of the nervous system
that are damaged.
Much of what we call voluntary movement (as
opposed, to the elicitation of simple reflexes) is
mediated by connections that in a sense wander
through a complex neuronal maze from the cerebral
cortex to the spinal cord. Thus, the primary motor
cortex, which is somatotopically organized as is the
somatic sensory cortex (meaning that different body
parts are connected to specific neural regions), influ-
ences specific muscle groups. The cortex itself is
layered, with neurons forming groups that are rather
like a series of pies stacked upon one another. It is in
what anatomists call layer V (counting from the top of
the cortex) of the primary motor cortex that one finds
projections directly to motor neurons and inter-
neurons, in the ventral (bottom) level of the spinal
cord, via what is called the corticospinal tract. In
humans there are approximately one million axons
(those processes of nerves that carry signals out from
the neurons cell body) and just under 50 percent of
these originate from the motor cortex. Other parts of
the cortex, such as the premotor areas and the
supplementary motor areas, also exert important and
distinct influences upon our movement skills. Thus
our cortices, the highest parts of the human brain,
connect us more or less directly to the world through
movement.
From the cortex the neural axons pass through
white matter of the sub-cortex, through the internal
capsule and cerebral peduncles on their way to the
spinal cord, and hence the muscles that make move-
ment possible (see Fig. 1). The term white matter
refers to the fact that these neurons have a fatty
insulating sheath that promotes rapid and reliable
transmission of signals from one neuron to another.

These tracts in turn form the medullary pyramids deep
within the brain (a region called the medulla). The
projection encompasses what has classically been
called the pyramidal tract. In addition to the pyra-
midal tract there is the extrapyramidal system, which
is composed of other nuclei, such as those found in the
basal ganglia. These extrapyramidal systems are for
the most part more indirect in their motor control
functions than are the pyramidal pathways, but their
importance should not be underestimated. For exam-
ple, linkages between emotional and cognitive pro-
cesses and the motor system are largely mediated
through these extrapyramidal systems.
The left side of the brain primarily is involved with
movements on the right side of the body, and vice
versa, but the crossing of fibers is incomplete. Ulti-
mately, the coordination of movements from each side
of the body involves unified actions of neural circuits
that are in themselves primarily concerned with one
body side.
The corticospinal tract itself makes monosynaptic
(i.e., direct) connections with motor neurons, which in
turn control muscular contractions. There are also
many connections to interneurons in the spinal cord.
The importance of these indirect connections is that
they allow a number of influences from other brain
regions that are essential for adaptive motor per-
formance. Throughout, the activities of the cortico-
spinal tract and related pathways incorporate a rich
variety of descending modulating influences.
3.2 Moement Disorders and Deelopment
In addition to the often tragic consequences of neural
damage (e.g., stroke) the neurologist and behavioral
neurobiologist can gain important insights into the
foundations of motor control by examining the de-
velopmental progression of motor skills. There are
predictable milestones in the construction of human
motor skills. A relatively recent perspective has stres-
sed the theme of ontogenetic adaptations in motor
control (Oppenheim 1982). In the past it was cust-
omary to think of nervous systems starting out as
imperfect adult systemsrather like a linear prog-
ression from less than fully adequate to fully adequate.
This view has now changed. Think, for example, of the
newborn infant who must swallow (which involves a
complex coordination of 20 plus muscles in the throat;
Doty 1976), and breathe and suckle. There is no room
for error or practice. Indeed, many motor circuits are
formed in a prefunctional manner. This means that the
circuits are established without the benefit of overt
motor practice. At a more detailed neuronal level these
circuits develop in what has been called a retrograde
manner, with the establishment of final neuromuscular
and other motor circuits preceding the establishment
of higher-order interneuronal influences and thus
Motor Control
responses to sensory events from the internal and
external environments (Fentress and Gadbois 2001).
This reminds us of the work of ethologists (e.g.,
Hinde 1970, Lorenz 1950, Tinbergen 1963) who
stressed the instinctive nature of many actions in
animals and humans. The basic idea is that many of
the fundamental properties in animal and human
behavior are a relatively direct reflection of our genetic
make-up. People, for example, speak; other species do
not. Many basic human and animal capacities for
movement emerge relatively independently of spec-
ialized experiences. In some deep sense we are indeed
preprogrammed to be human, just as the bird, and
bat, and insect are preprogrammed to be members of
their own species. What is perhaps most remarkable
about humans is that our genetic structure allows for
enormous flexibility in the detailed organization of our
movements. In that respect we stand alone among the
animal species. Yet, for all animal species the delicate
interplay between genetic and experiential factors
(broadly defined, across many levels of organization)
remains a major area of importance for further
investigation. An essential point is that even the most
basic processes of differentiation in development
involve genes and experience working in concert
(Hinde 1970, Edelman 1987).
How is it that we learn motor skills, and how are
these skills mediated? At the present time our knowl-
edge of these essential questions is very limited.
Certainly many of the mechanisms we have touched
upon contribute richly. Thus, cerebellar circuits are
important in the establishment of newly acquired
reflexes as well as more complex cognitive skills
(Thompson, Thach). The basal ganglia as well as
cortical mechanisms also contribute to perfections of
motor performance through experience (Graybiel).
The importance of cortical mechanisms is perhaps
predominant, but a challenge for future research is to
determine how various neural mechanisms operate
together at various stages of skill acquisition and
performance (see Neural Plasticity in Auditory Cortex;
Neural Plasticity; Neural Plasticity of Spinal Reflexes;
Neural Plasticity in Visual Cortex).
4. The Need to Combine Mechanistic and
Systems Perspecties
A challenge of increasing importance in both the
neural and behavioral sciences is how to capture the
dynamic as well as modular properties of motor
performance. Excellent overviews that cross-
complementary levels of organization can be found in
Gazzinga 2000, Kandel et al. 2000. Most investiga-
tors in the neurobehavioral sciences today take the
position that any level of analysis must make contacts
with levels that are both more molar and more
molecular.
10129
Motor Control
5. Challenges for Future Research
Human motor control systems are awesome in their
richness. It is through these motor control systems
that we interact as individuals, yet individuals deeply
connected to other human individuals and to our
animal brethren by solutions that evolution has
enabled us with. We talk, sing, and dance. In spite of
recent progress at many levels of research, precisely
how we do these basic things remains deeply mys-
terious. Yet, piece by piece, we are beginning to
understand the operation of individual mechanisms
and their coordinated actions. For example, the rapid
progress in brain imaging techniques has helped us
understand how the final processes of motor control
are connected to higher-order cognitive functions
(see Vision for Action: Neural Mechanisms; Cognitie
Control (Executie Functions): Role of Prefrontal
Cortex; Functional Brain Imaging; Neural Represent-
ations of Intended Moement in Motor Cortex). The
further understanding of motor control processes,
including not just their performance on a short-term
basis, but also their development and evolution, offers
riches into the nature of the human psyche itself.
In recent years a number of model systems for
motor control have been offered (see Computational
Neuroscience). One of the most promising is that there
are variable foci of activity that occur in conjunction
with variable expanses of collateral inhibition, that is,
those events that block competing activity (Fentress
1976). For example, both the cerebellum and basal
ganglia have lateral inhibitory (collateral) circuits that
suppress action properties that would otherwise inter-
fere with the primary focus of adaptive movement.
There is now growing evidence that these collateral
pathways can expand in their extent of influence as the
core of motor performance becomes more tightly
focused (Fentress 1976). What this means is that
systems in the control of movement can be both
broadly and much more finely tuned as circumstances
dictate. Extrinsic influences are either accepted or
rejected as the needs of the organism change.
In terms of the establishment of motor behavior
during development, a number of genetic analysis that
focus upon specific neural circuits are now being
combined with experiments designed to clarify the
roles and limits of specific sources of ontogenetic
experience. The old nature vs. nurture debate is now
considered passeT by most active investigators; the
challenge is to see how genes and experience work as
partners during the construction of brains and their
behavioral manifestations.
In motor control we have much to learn about the
detailed operation of specific mechanisms, but equally
much to learn how these specific mechanisms become
established and integrated into the rich symphonies of
human and animal action. Whatever the details that
will be revealed by future research, it is clear that
motor control mechanisms not only reflect what we
10130
mean by behavior, but also give us insights into the
inner workings of nervous systems that remain a
major source of inspiration and mystery for all those
who are fortunate enough to study them.
Major components of the human motor system.
Spinal cord, brain stem, and motor areas of the
cerebral cortex work together with direct and indirect
connections of the basal ganglia and cerebellum in the
planning and execution of integrated movement pat-
terns.
See also: Basal Ganglia; Behavior, Hierarchical Organ-
ization of; Cerebellum: Associative Learning; Cereb-
ellum: Cognitive Functions; Eye Movement, Control
of (Oculomotor Control); Motor Control Models:
Learning and Performance; Motor Cortex; Neural
Representations of Intended Movement in Motor
Cortex; Neuromuscular System; Short-term Memory,
Cognitive Psychology of
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J. C. Fentress
Motor Control Models: Learning and
Performance
Modeling the way in which humans learn to co-
ordinate their movements in daily life or in more
demanding activities is an important scientific topic
from many points of view, such as medical, psycho-
logical, kinesiological, cybernetic. The article analyzes
the complexity of this problem and reviews the variety
of experimental and theoretical techniques that have
been developed for this purpose.
With the advent of technical means for capturing
motion sequences and the pioneering work of Marey
and Muybridge in this area, the attempt at describing,
modeling, and understanding the organization of
movement has become a scientific topic. The fact that
human movements are part of everyday life para-
doxically hides their intrinsic complexity and justifies
initial expectations that complete knowledge could be
achieved simply by improving the measurement tech-
niques and carrying out a few carefully designed
experiments. Unfortunately, this is not the case. Each
experiment is frequently the source of more questions
than answers and thus the attempt to capture the
complexity of purposive action and adaptive behavior,
after a century of extensive multidisciplinary research,
is far from over.
The conventional view is based on a separation of
perception, movement, and cognition and the seg-
regation of perceptual, motor, and cognitive processes
in different parts of the brain, according to some kind
of hierarchical organization. This view is rooted in the
empirical findings of neurologists of the nineteenth
century, such as J. Hughlings Jackson, and has a
surprising degree of analogy with the basic structure of
Copyright # 2001 Elsevier Science Ltd. All rights reserved.
International Encyclopedia of the Social & Behavioral Sciences
Motor Control Models: Learning and Performance
a modern PC that typically consists of input and
output peripherals connected to a central processor.
Perhaps the analogy with modern technology justifies
why this old-fashioned attitude still has its supporters,
in spite of the massive empirical and conceptual
challenge to this view and its inability to explain the
range of skills and adaptive behaviors that characterize
biological organisms.
Let us consider perception, which is the process
whereby sensory stimulation is translated into organ-
ized experience. That experience, or percept, is the
joint product of the stimulation and of the process
itself, particularly in the perception and representation
of space. An early theory of space perception put forth
by the Anglican bishop G. Berkeley at the beginning of
the eighteenth century was that the third dimension
(depth) cannot be directly perceived in a visual way
since the retinal image of any object is two-
dimensional, as in a painting. He held that the ability
to have visual experiences of depth is not inborn but
can only result from logical deduction based on
empirical learning through the use of other senses.
The first part of the reasoning (the need of a
symbolic deductive system for compensating the fal-
lacy of the senses) is clearly wrong and the roots of
such misconception can be traced back to the neo-
platonic ideas of the Italian Renaissance, in general,
and to Albertis window metaphor, in particular.
Also, the Cartesian dualism between body and mind is
just another face of the same attitude and such
Descartes error, to quote Damasio (1994), is on a par
with the Berkeleys error above and is at the basis of
the intellectualistic effort to explain the computational
complexity of perception which characterizes a great
part of the classic artificial intelligence approach.
However, the latter part of Berkeleys conjecture (the
emphasis on learning and intersensory integration) is
surprisingly modern and agrees, on one hand, with the
modern approach to neuropsychological development
pioneered by Piaget and, on another, with the so-called
connectionist point of view, originated in the 1980s as
a computational alternative to classic artificial in-
telligence (see Artificial Neural Networks: Neuro-
computation).
An emergent idea is also the motor theory of
perception, well illustrated by Berthoz (1997); that is,
the concept that perception is not a passive mechanism
for receiving and interpreting sensory data but is the
active process of anticipating the sensory conse-
quences of an action and thereby binding the sensory
and motor patterns in a coherent framework. In
computational terms, this implies the existence in the
brain of some kind of internal model, as a bridge
between action and perception. As a matter of fact, the
idea that the instructions generated by the brain for
controlling a movement are utilized by the brain for
interpreting the sensory consequences of the move-
ment is already present in the pioneering work of
Helmholtz and von Uexku$ ll and its influence has
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