Motor Cortex
von Holst E, Mittelstaedt H 1950 Das Reafferenzprinzip.
Wechselwirkungen zwischen Zentralnervensystem und Peri-
pherie. Naturwissenschaften 37: 46476
Jeannerod M 1988 The Neural and Behaioral Organisation of
Goal-directed Arm Moements. Clarendon Press, Oxford, UK
Kawato M, Gomi H 1992 A computational model of four
regions of the cerebellum based on feedback error learning.
Biological Cybernetics 69: 95103
Marey E J 1894 Le mouement. E; dition Masson, Paris
Morasso P 1981 Spatial control of arm movements. Exper-
imental Brain Research 42: 2237
Morasso P, Sanguineti V 1997 Self-organization, Cortical Maps
and Motor Control. North Holland, Amsterdam
Muybridge E 1957 The Human Figure in Motion. Dover Press,
New York
Paillard J 1993 Brain and Space. Oxford University Press,
Oxford, UK
Piaget J 1963 The Origin of Intelligence in Children. Norton
Press, New York
Ramo! n y Cajal S 1928 Regeneration in the Vertebrate Central
Nerous System. Oxford University Press, Oxford, UK
Rizzolatti G, Luppino G, Matelli M 1998 The organization of
the cortical motor system: New concepts. Electrencephalo-
graphy and Clinical Neurophysiology. 106: 28396
Shepherd G M 1998 The Synaptic Organization of the Brain.
Oxford University Press, Oxford, UK
Sutton R S, Barto A G 1998 Reinforcement Learning. MIT
Press, Cambridge, MA
Wolpert D M, Kawato M 1998 Internal models of the cer-
ebellum. Trends in Cognitie Science 2: 33847
P. G. Morasso
Motor Cortex
The term motor cortex has both a broad and a narrow
sense. The broad meaning refers to a number of
functionally distinct cortical areas, which collectively
control voluntary action (Wise 1999). The narrow
sense, adopted here, pertains exclusively to one of
those areas, the primary motor cortex (Fig. 1). The
primary motor cortex comprises only a small pro-
portion of the cerebral cortex but plays a crucial role in
controlling movement. Some of its neurons have axons
that terminate directly on motor neurons in the spinal
cord, giving motor cortex the most direct possible
access to muscles (Porter and Lemon 1993).
1. History of Motor Cortex
The motor cortex has a short history by evolutionary
standards. Before the advent of mammals, about 200
million years ago, the motor cortex did not exist. Only
mammals evolved a motor cortex in the sense used
here. Accordingly, it probably functions to supple-
ment the remainder of the vertebrate motor system
(see Cerebellum: Cognitie Functions; Motor Control),
which evolved more than 500 million years ago.
In contrast to its relatively short evolutionary
history, on a geological timescale, the motor cortex
has a long intellectual history by the standards of
neurobiology. In the fourth and fifth centuries BC,
Hippocratic physicians observed that damage to one
side of the brain caused motor disabilities on the
opposite side of the body (Lassek 1954). In the
nineteenth century, G. Theodor Fritsch and Eduard
Hitzig in Germany and David Ferrier in England
confirmed experimentally the existence of a localized
motor cortex. They showed that electrical stimulation
of the motor cortex in dogs and monkeys produced
movements on the opposite side of the body. Although
condemned by antivivisectionists of the time as in-
applicable to humans, this late nineteenth-century
animal research established the principles of cortical
organization for all mammals, including our species.
2. Description of Motor Cortex
Like other areas of the cerebral cortex, the motor
cortex consists of several cell layers. However, the
motor cortex exceeds other areas in thickness. Four
other characteristics collectively identify the motor
cortex: electrically stimulating it evokes movements
more easily than for other cortical areas; it lacks a
layer common to most cortical areas (layer 4); it
receives relatively direct inputs from the basal ganglia
and cerebellum; and it lies immediately in front of the
somatosensory cortex. The neurons that send axons
directly to the spinal cord all arise from layer 5,
including some of the largest neurons in the cerebral
cortex. Named after Vladimir Betz, these giant neu-
rons can reach 60 m in cell-body width in monkeys
and 120 m in people. Cells in the motor cortex send
outputs to and receive inputs from somatosensory and
premotor cortex. Subcortical targets include the basal
ganglia, cerebellum, red nucleus, and other parts of the
brain stem and spinal motor system. According to
contemporary theory, individual cortical cells function
as components of distributed neural networks incorp-
orating a number of anatomically distinct structures,
rather than as part of a localized center performing a
certain function.
2.1 Nature of the Body Map
By noting the progression of epileptic seizures from
one body part to another, the British neurologist J.
Hughlings Jackson surmised that the motor cortex
was arranged in a rough map of the body. Electrical
brain stimulation confirmed that conclusion. Nearest
the top of the brain (Fig. 1), the leg is represented.
Representation, in this sense, implies that this part of
motor cortex processes information about leg move-
10137
Motor Cortex
ments. The leg representation sends some of its axons
to motor neurons that control leg muscles. Adjacent to
the leg representation, in roughly the middle of the
hatched zone in Fig. 1 (from top to bottom), lies an
arm representation. The part of motor cortex that
controls the face, head, tongue, and lips is situated yet
further from the top of the brain. The finer organiza-
tion of the motor cortex remains controversial and,
although many detailed maps have been proposed,
none have stood the test of time. In this respect, the
organization of motor cortex contrasts with that of the
somatosensory, visual, and auditory areas of the
cerebral cortex, which have finer and more orderly
maps of the receptors for touch, sight, and hearing,
respectively.
3. Function of Motor Cortex
The earliest knowledge about the function of motor
cortex arose from observing the effects of damage to it
or its main output pathway, the pyramidal tract.
Unfortunately, the medical literature often lacks an
accurate assessment of the exact site and extent of
brain damage. It is clear, however, that a number of
severe motor disabilities follow such damage, includ-
ing weakness, difficulty in controlling the muscles
individually, and loss of fine motor control. It is
commonly held that the motor cortex functions mainly
to permit motor fractionation, i.e., the independent
control of muscle groups that usually work in concert.
In this sense, the motor cortex can be viewed as
supplementing the subcortical motor system. Accord-
ing to this idea, motor fractionation adds an important
degree of flexibility to the motor system. Although in
Figure 1
The cerebral cortex of a macaque monkey (left) and human (right), to different scales. Each of the lines within the
outline depicts the typical location of the sulci, or grooves, in the cortex. The arrows show the distance from the
front to the back of the cortex. The zone with the oblique hatching is the primary motor area. Source: Passingham
(1993), by permission of Oxford University Press. Passingham R E 1993 The Frontal Lobes in Voluntary Action.
Oxford University Press, Oxford, UK
10138
many primates, including humans, independent move-
ment of the fingers typifies fractionation, the ability to
fractionate movements is by no means confined to the
digits. Whatever the motor cortex does, it does for the
body as a whole.
In the early 1960s, the American neurophysiologist
Edward Evarts developed a method for recording the
activity of individual neurons in the motor cortex of
awake, behaving monkeys. He began by investigating
a question first posed by Jackson in the nineteenth
century. Does the motor cortex control muscles or
movements? This seemingly innocuous question en-
gaged researchers for many years. It was finally
resolved by dividing the question into two compon-
ents, which yielded contradictory answers. One ques-
tion concerns the addressing of muscles by motor
cortex. Do cells in motor cortex address individual
muscles (the muscle view) or several muscles, which
may act together to produce a coherent action (the
movement view). Early brain-stimulation studies
suggested single-muscle addressing. However, with
more careful research the multiple-muscle movement
hypothesis prevailed for the specific issue of address-
ing. The other question concerns the signal communi-
cated to these muscle groups. Does motor cortex
control the level and pattern of force generated by
muscles (the muscle view) or, alternatively, joint
angles, limb position, and the direction of movement
(the movement view)? Neurophysiologists have
found that most cells in the motor cortex control
muscle activity rather than movement. Motor cortex
emits a force signal, which usually causes movement,
but may instead generate force without movement
(e.g., when one pushes against an immovable object).

A smaller number of cells control a combination of
force and movement.
3.1 Population Coding
Individual neurons have their greatest activity for
force in one direction, with systematically less activity
as the direction of force diverges from that preferred
direction. The cells are thus said to be tuned for
direction. On the assumption that these cells generate
force in their preferred direction, it is possible to
estimate a collective force signal emanating from a
neuronal population. This computation, termed the
population ector, matches closely the direction of
movement and force generated in a straight line, as
well as both spiral and sinusoidal trajectories
(Georgopoulos 1991).
3.2 Motor Cortex, Volition, and Awareness
Voluntary actions are those that are learned, attended,
and based on a comparison among alternatives
(Passingham 1993). Other kinds of action include
reflexes and other innate behaviors. It seems likely that
the motor cortex controls voluntary actions, which are
typically based upon consciously perceived informa-
tion. However, this contention implies neither that it
lacks a role in reflexes nor that all voluntary move-
ments depend upon consciousness.
Recently, the distinction between explicit (or de-
clarative, perceived) knowledge and implicit (or pro-
cedural, unperceived) knowledge has gained wide
acceptance. The former has access to conscious aware-
ness; the latter does not. Both explicit and implicit
knowledge can guide voluntary action. For example,
people with blindsight can point to a visual stimulus
while denying that they see it (Weiscrantz 1986).
Similarly, one well-studied neurological patient can
orient her hand in order to put it through a narrow
slot, but cannot report the orientation of the same slot
(Milner and Goodale 1995). These are examples of
implicit, unperceived knowledge guiding voluntary
action. It seems likely that the motor cortex is involved
in both explicitly and implicitly guided voluntary
movements.
It has become increasingly clear that the processing
of sensory information for perception differs from that
for movement. In some respects, the performance of
the motor system surpasses the neural systems under-
lying perception. For example, people can make
movements that accurately match the size of an object
to be grasped, but report incorrectly the size of the
same object due to a visual illusion (Milner and
Goodale 1995). The motor system thus has access to
somewhat different sensory information than the brain
systems underlying perception. As a consequence,
one cannot assume that peoples actions invariably
reflect their perceived knowledge.
Motor Cortex
3.3 Motor Learning and Plasticity
Motor learning and memory include both adaptation
and skill acquisition. Adaptations involve changes in
motor performance without an enhancement in capa-
bilities. For example, if a prism shifts the visual image
of a target, it takes several attempts to correct the error
induced in reaching toward that target. Such ad-
aptation leaves the system able to reach or throw
accurately, but no more so than before adaptation. By
contrast, learning a skill involves obtaining a new
capability or improved performance, beyond the
systems prior limits. Evidence from brain-imaging
studies suggests that the motor cortex changes the way
it processes information during both skill learning
(e.g., learning a movement sequence) and adaptation
(e.g., tracking a rotating target). Other methods lead
to the same conclusion. During sequence learning, for
example, the movements caused by stimulating motor
cortex become larger and easier to evoke (Hallett
1999). And during motor adaptation, the activity
levels and other properties of neurons change.
Nearly 1 percent of the population has suffered a
stroke causing motor disability. Many of these patients
survive for years, 35 percent for a decade or more.
Recovery of function usually proceeds slowly and
often remains incomplete. Recent research has shown
that on account of the brains plasticity, important
organizational changes occur after damage to motor
cortex. Practice that leads to such plasticity enhances
recovery of function (Nudo et al. 1997). Future
research will likely focus on drug treatments and other
therapies aimed at regulating cortical plasticity. The
advent of treatments to ameliorate damage to the
motor cortex could have effects resembling the deve-
lopment of correctional optics in the thirteenth cent-
ury. Eyeglasses extended the productivity of artisans
by freeing them from the limitations of middle-age
farsightedness (Landes 1998). Extending the prod-
uctivity of patients with motor cortex damage could
have similar benefits for both the individual and
society.
See also: Cerebellum: Cognitive Functions; Motor
Control; Motor Control Models: Learning and Per-
formance; Motor Skills, Psychology of; Pre-motor
Cortex
Bibliography
Georgopoulos A P 1991 Higher order motor control. Annual
Reiew of Neuroscience 14: 36177
Hallett M 1999 Plasticity in the human motor system. Neuro-
scientist 5: 32432
Landes D S 1998 The Wealth and Poerty of Nations: Why Some
are so Rich and Some are so Poor. 1st edn. W W Norton, New
York
10139
Motor Cortex
Lassek A M 1954 The Pyramidal Tract: Its Status in Medicine.
Thomas, Springfield, IL
Milner A D, Goodale M A 1995 The Visual Brain in Action.
Oxford University Press, Oxford, UK
Nudo R J, Plautz E J, Milliken G W 1997 Adaptive plasticity in
primate motor cortex as a consequence of behavioral expe-
rience and neuronal injury. Seminars in Neuroscience 9: 1323
Passingham R E 1993 The Frontal Lobes in Voluntary Action.
Oxford University Press, Oxford, UK
Porter R, Lemon R 1993 Corticospinal Function and Voluntary
Moement. Clarendon Press, Oxford, UK
Weiscrantz L 1986 Blindsight. A Case Study and Implications.
Oxford University Press, Oxford, UK
Wise S P 1999 Motor cortex. In: Adelman G, Smith B H (eds.)
Encyclopedia of Neuroscience, 2nd edn. Elsevier,
Amsterdam
S. P. Wise
Motor Skills, Psychology of
The history of the psychology of motor skills starts at
the end of the nineteenth century. Among the land-
marks are Woodworths studies of aimed movements,
which established a fundamental distinction between a
first ballistic movement phase and a later visually
guided homing-in phase. Another landmark are Bryan
and Harters studies of the telegraphic language,
which established the notion that during the acqui-
sition of motor skills elementary units are integrated
to form higher-level units. In the early twentieth
century, Thorndikes law of effect motivated a
number of studies on the effects of knowledge of
results on skill acquisition, and later Hulls concepts of
reactive and conditioned inhibition formed the theor-
etical foundation for studies of massed and distributed
practice. Among the discoveries emanating from
European schools of psychology were the covariations
between components of motor skills and the almost
constant duration of movements when identical figures
of different sizes are drawn.
From the 1960s on there was a noticeable increase in
research on motor skills. Concepts of cognitive psy-
chology turned out to be better suited for the analysis
of motor behavior than, for example, concepts of
behaviorism. In addition, many researchers came to
entertain the idea that cognitive processes primarily
serve the control of action rather than cognition per se.
Finally, there was a merging of different academic
disciplines such as neurobiology, physiology, neur-
ology, biomechanics, psychology, and physical edu-
cation to form what is called movement science or
kinesiology.
The psychology of motor skills can be organized in
two fundamentally different ways. One organization is
according to different skills such as standing, walking,
10140
Copyright # 2001 Elsevier Science Ltd. All rights reserved.
International Encyclopedia of the Social & Behavioral Sciences
speaking, and writing. The other relates to basic
concepts that are applicable to real-life skills, but
typically studied by means of artificial tasks that are
particularly suited for their purpose. Here some major
concepts will be described.
1. Autonomous and Stimulus-based Processes of
Motor Control
There is a kind of classical controversy which, his-
torically, came in at least two different formats. In
both formats it is about the question of how critical
sensory stimulation is for the production of move-
ment. In the historically earlier format, the alternative
conceptions were that of a reflex chain, according to
which stimuli which arise from one elementary move-
ment trigger the next elementary movement, and that
of autonomous activity of the central nervous system
in generating the basic motor patterns. In the his-
torically later format, the reflex chain was replaced by
the notion of a continuous processing of sensory
feedback.
The controversy about the roles of autonomous and
stimulus-based processes in motor control has dis-
solved. The surviving problem is how sensory in-
formation is integrated with autonomous processes.
Basically, sensory information can be used continu-
ously as a feedback signal or as a reference signal,
intermittently for the updating of parameters of
autonomous processes or for the triggering of new
autonomous processes, and finally sensory informa-
tion can be used before a movement is initiated.
There is good reason to assume that autonomous or
feed-forward processes of motor control run in parallel
to closed-loop processes. Such an arrangement can
actually be identified anatomically in the spinal control
of muscle length, and it helps in understanding some
fundamental characteristics of human movements.
First, closed-loop systems have a tendency to become
unstable when the feedback is processed with some
delay and the gain in the loop is sufficiently high. In
biological systems feedback is always processed with
some delay, and the high velocities of human move-
ments would require a high gain of a closed-loop
system. Nevertheless, instabilities are extremely rare.
With open-loop control in parallel with closed-loop
control, the closed-loop system can operate with a
small gain. Hence instabilities can be avoided, but the
inaccuracies of open-loop control can nevertheless be
compensated.
Second, primate movements are astonishingly little
affected by elimination of peripheral feedback, even by
more or less complete deafferentation. Such observa-
tions suggest that peripheral feedback is not processed
at all. On the other hand, the processing of peripheral
feedback is clearly indicated by strong effects of
ISBN: 0-08-043076-7